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XIV Annual Meeting of the European Association of Vertebrate Palaeontologists

6-10 July 2016, Haarlem, The Netherlands

Programme and Abstract Book

Edited by:

EAVP 2016 Programme & Abstract Committee Femke Holwerda, Anneke Madern, Dennis Voeten, Anneke van Heteren, Jeff Liston, Hanneke Meijer, Natasja den Ouden

EAVP 2016 Programme & Abstract Crew Stephan Spiekman, Tom Trapman, Feiko Miedema, Sifra Bijl, Mart Smeets, Pim Kaskes, Tim Rietbergen, Juliën Lubeek

XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Contents

Map of Haarlem 2

Map of Teylers Museum 3

Programme at a glance 4

Posters at a glance 5

Programme 6

Abstracts 18

List of authors 220

1 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Map of Haarlem

2 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Map of Teylers museum and venue information

Teylers Museum, Spaarne 16, 2011 CH Haarlem, The Netherlands

Phone: 0031 023 51 60 960 Phone (emergencies only!!): 0031 06 483 22 755

3 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Maasvlakte Pleistocene Sunday July 10 Limburg Maastrichtian den Ouden/Reumer small hall Sander Krahl Neenan Heijne During Czepiński Marzola Nieuwland Monsch Micklich Kaskes Czepiński Slowiak Veitschegger Stinnesbeck Kummel Eck spare intro: Mol Jagt Mulder coffee coffee coffee LUNCH symposium Germanic Basin Germanic Basin Germanic Basin Germanic Basin Germanic Basin Germanic Basin Germanic Basin plenary plenary plenary plenary plenary plenary plenary plenary plenary plenary plenary North Sea Basin North Sea Basin North Sea Basin North Sea Basin Barrett Butler Delfino de Vos main hall keynote 5: Maidment keynote 6: Intro: Lordkipanidze Amkreutz Lister Aytek Alpagut Marcé-Nogué Sier Joordens spare/discussion Unwin Cincotta Moreno-Azanza keynote 7: Saturday July 9 symposium plenary plenary plenary Hominins Hominins Hominins Hominins Hominins Hominins Hominins Hominins Hominins Hominins Hominins Hominins Hominins Hominins plenary plenary plenary plenary Hoch small hall Rivera Kaskes Meyer Belvedere Razzollini Castanera Bronzati Diez Diaz Stein Tortosa Rodrigues Marcé-Nogué De Esteban-Trivigno Belvedere Marchetti Rabi Tong Joyce Foth Göhlich Voeten Pimiento Muchagata keynote 4: Liston (Cafetaria) symposium plenary plenary plenary plenary plenary plenary plenary plenary plenary plenary plenary plenary plenary plenary plenary plenary plenary plenary plenary plenary plenary plenary plenary plenary coffee coffee coffee LUNCH Conference dinner main hall Henderson Challands Smithson Clack Arbez Marjanovic Gruntmejer Teschner Chakravorti Sengupta Steyer Fortuny Perez Ben Elsler Mazza vd Hoek Ostende Masters Rijsdijk/Meijer van Heteren Winkler Bethune Palombo Savorelli Buffetaut Codrea Wings break, remove posters, change for dinner Programme at a glance Programme Friday July 8 symposium Tetrapods Tetrapods Tetrapods Tetrapods Tetrapods Tetrapods Tetrapods Tetrapods Tetrapods Tetrapods Tetrapods Tetrapods Tetrapods Tetrapods Islands Islands Islands Islands Islands Islands Islands Islands Islands Islands Islands Islands Kostopoulos-Konidaris small hall intro: Vasile Pazonyi Bukhsianidze Konidaris Demirel Mayda Belmaker Pandolfi Baigusheva Madurell-Malapeira Maniakas Berlioz Tesakov Adams Martin van Kolfschoten discussion Palombo Martínez-Navarro Sardella Croitor Pawłowska : history and collections of Teylers) coffee coffee LUNCH symposium Biogeography Biogeography Biogeography Biogeography Biogeography Biogeography Biogeography Biogeography Biogeography Biogeography Biogeography Biogeography Biogeography Biogeography Biogeography Biogeography Biogeography Biogeography Biogeography Biogeography Biogeography Biogeography Biogeography coffee/registration Trienke van der Spek AGM Auction main hall keynote 1: Meyer Baez keynote 2: Maxwell keynote 3: Joyce intro: Liston Martin Frey Unwin Martill Rodrigues Maltese Siber Barrett Rauhut Albersdörfer Stead Liston roundtable welcome address ( Thursday July 7 symposium plenary plenary plenary plenary Fossillegal Fossillegal Fossillegal Fossillegal Fossillegal Fossillegal Fossillegal Fossillegal Fossillegal Fossillegal Fossillegal Fossillegal Fossillegal Fossillegal Fossillegal Fossillegal (Rotonde) general hall registration Science Workshop tour Teylers Transmitting Wednesday July 6 museum misc 4 Icebreaker party 08:00 08:15 08:30 08:45 09:00 09:15 09:30 09:45 10:00 10:15 10:30 10:45 11:00 11:15 11:30 11:45 12:00 12:15 12:30 12:45 13:00 13:15 13:30 13:45 14:00 14:15 14:30 14:45 15:00 15:15 15:30 15:45 16:00 16:15 16:30 16:45 17:00 17:15 17:30 17:45 18:00 18:15 18:30 18:45 19:00 19:15 19:30 19:45 day time XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Posters at a glance

5 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Thursday July 7

Main Hall

08:30 Welcome by EAVP President Dino Frey and Introduction by Trienke van der Spek, Teylers Museum

Plenary

08:45 Christian Meyer: TRACKING IN SWITZERLAND - PAST AND PRESENT (keynote)

09:15 Ana Baez: NEW EVIDENCE FROM THE LOWER OF PATAGONIA SHEDS LIGHT ON THE HISTORY OF SOME OF THE EARLIEST KNOWN

09:30 Erin Maxwell: NEW ICHTHYOSAURS FROM THE EARLY OF COLOMBIA, WITH IMPLICATIONS FOR DIVERSITY IN THE PALAEOTROPICS (keynote)

10:00 Walter Joyce: NEW INSIGHTS INTO THE EVOLUTION OF (keynote)

10:30 - 11:00 coffee break 1 and poster session

Symposium ‘Fossillegal’ hosted by Jeff Liston

11:00 Jeff Liston: introduction

11:15 John Martin: INTERNATIONAL LEGISLATION, THE SEMANTICS OF ‘CULTURAL PROPERTY’, AND WHY THEY ARE BAD FOR PALAEONTOLOGY AND PEOPLE (keynote)

11.45 Eberhard ‘Dino’ Frey: : NATIONAL MONUMENTS, TRADING OBJECTS, OR BOTH?

12:00 Dave Unwin: HANDS OFF, IT’S MY COLLECTION! FOSSILS, MUSEUMS AND THE PROBLEMS OF ACCESS

12:15 Dave Martill: SHOULD PALAEONTOLOGISTS BEHAVE ETHICALLY? PROTECT THE FOSSILS OR PROTECT THE COMMUNITIES WHO DIG THEM?

12:30 - 14:00 lunch break and poster session

14:00 Taissa Rodrigues: BRAZILIAN PALAEONTOLOGICAL LEGISLATION

14:15 Anthony Maltese: COLLECTION AND PLACEMENT OF FOSSILS IN COMMERCIAL PALAEONTOLOGY: PRACTICAL AND ETHICAL CONCERNS

14:30 Hans-Jacob ‘Kirby’ Siber: 40 YEARS AS A PROSPECTOR, COLLECTOR AND EXHIBITION MAKER

6 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

14:45 Paul Barrett: PUBLISH OR PROHIBIT? THE ETHICS OF PUBLISHING ON PRIVATELY-OWNED FOSSILS

15:00 Oliver Rauhut: PRIVATE COLLECTIONS: PROBLEM OR BONUS FOR SCIENCE?

15:15 - 15:45 coffee break 2 and poster session

15:45 Raymund Albersdörfer: FOSSIL LEGISLATION – PROTECTION OR DESTRUCTION?

16:00 Nicola Stead: FOSSILS, PUBLISHING AND OPEN ACCESS

16:15 Jeff Liston: AN INNOCENT ACADEMIC ABROAD?: EASY FOSSILS AND HARD CHOICES, CHINA AS A MICROCOSM OF THE CHALLENGES OF 21ST CENTURY PALAEONTOLOGY

16:30- 17:00 roundtable discussion

Small Hall

Symposium ‘Towards a Biogeographic Synthesis for the European Early Pleistocene based on the Record’ hosted by Dimitris Kostopoulos and George Konidaris

08:45 Dimitris Kostopoulos and George Konidaris: introduction

09:00 Stefan Vasile: NEW DATA ON THE EARLY PLEISTOCENE VERTEBRATE ASSEMBLAGES FROM COPĂCENI (DACIAN BASIN, SOUTHERN ROMANIA)

09:15 Piroska Pazonyi: TAXONOMICAL, TAPHONOMICAL AND PALEOECOLOGICAL ASPECTS OF A LATE EARLY PLEISTOCENE VERTEBRATE FAUNA FROM THE SOMSSICH HILL 2 SITE (SOUTH HUNGARY)

09:30 Maia Bukhsianidze: DMANISI ARTIODACTYL ASSEMBLAGE

09:45 George Konidaris: TSIOTRA VRYSSI: A NEW VERTEBRATE LOCALITY FROM THE EARLY PLEISTOCENE OF MYGDONIA BASIN (MACEDONIA, GREECE)

10:00 Fatma Demirel: KOCAKIR-2: FIRST RECORD OF EARLY PLEISTOCENE CANIS AND HIPPOPOTAMUS FROM BURDUR BASIN (SW TURKEY)

10:15 Serdar Mayda: SYSTEMATIC, BIOSTRATIGRAPHIC AND PALEOBIOGEOGRAPHIC REEVALUATION OF THE MAMMALIAN FAUNAS FROM THE TURKISH VILLAFRANCHIAN LOCALITIES OF THE KAMISLI, GULYAZI AND Y. SOGUTONU

10:30 - 11:00 coffee break 1 and poster session

11:00 Miriam Belmaker: BIOGEOGRAPHIC ORIGINS OF EARLY PLEISTOCENE MAMMALIAN FAUNA IN THE SOUTHERN LEVANT 7 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

11:15 Luca Pandolfi: PALEOBIOGEOGRAPHY AND CHRONOLOGY OF STEPHANORHINUS ETRUSCUS (FALCONER, 1858) (MAMMALIA, RHINOCEROTIDAE) IN EURASIA

11:30 V. Baigusheva and Vadim Titov: SOME BIOLOGICAL FEATURES OF ELASMOTHERIUM CAUCASICUM (MAMMALIA, RHINOCEROTIDAE) FROM THE LATE EARLY PLEISTOCENE OF EASTERN

11.45 Joan Madurell-Malapeira: LATEST EARLY PLEISTOCENE LARGE MAMMAL ASSEMBLAGES FROM THE IBERIAN PENINSULA AND THE EARLY-MIDDLE PLEISTOCENE TRANSITION

12:00 Ioannis Maniakas: BIOGEOGRAPHICAL AND PALAEOECOLOGICAL ASSOCIATIONS OF WESTERN PALAEARCTIC BISON POPULATIONS

12:15 Emilie Berlioz: DENTAL MICROWEAR TEXTURE ANALYSIS OF EARLY PLEISTOCENE CERVIDS: A PALEOENVIRONMENTAL APPROACH AT THE EUROPEAN SCALE

12:30 - 14:00 lunch break and poster session

14:00 Alexey Tesakov: EARLY PLEISTOCENE OF EASTERN EUROPE AND WESTERN : BIOCHRONOLOGY AND BIOGEOGRAPHY

14:15 Neil Adams: ADDRESSING THE ‘ALLOPHAIOMYS GAP’ IN BRITISH BIOSTRATIGRAPHY: A NEW EARLY PLEISTOCENE FAUNA FROM WESTBURY CAVE, SOMERSET,

14:30 Robert Martin: A NEW ARVICOLID RODENT FROM THE EARLY PLEISTOCENE OF THE BAZA BASIN, SOUTHERN SPAIN

14:45 Thijs van Kolfschoten: EARLY PLEISTOCENE ARVICOLINES FROM ZUURLAND (THE NETHERLANDS): NEW INSIGHTS INTO HOLARCTIC BIOCHRONOLOGY AND BIOGEOGRAPHY

15:00 roundtable discussion

15:15 - 15:45 coffee break 2 and poster session

15:45 Maria Rita Palombo: FAUNAL DYNAMICS IN SW EUROPE DURING THE LATE EARLY PLEISTOCENE: IMPLICATIONS FOR THE BIOCHRONOLOGICAL ASSESSMENT AND CORRELATION OF MAMMALIAN FAUNAS

16:00 Bienvenido Martinez-Navarro: THE LATE VILLAFRANCHIAN SITES OF ORCE (SPAIN) AND THE EARLIEST HUMAN RECORD OF WESTERN EUROPE

16:15 Raffaele Sardella: BEFORE AND AFTER EARLIEST HOMO DISPERSAL IN EUROPE: EVIDENCE FROM THE EARLY PLEISTOCENE SITES OF THE ITALIAN PENINSULA

16:30 Roman Croitor: PALEOBIOGEOGRAPHIC AND ECOLOGICAL CONTEXT OF EARLY HUMAN DISPERSALS IN WESTERN EURASIA 8 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

16:45 Kamilla Pawlowska: LARGE MAMMALS AFFECTED BY HOMININS: A BIOGEOGRAPHIC SYNTHESIS OF BUTCHERING FOR THE EUROPEAN EARLY AND MIDDLE PLEISTOCENE

Main Hall

17:00 - 18:00 AGM, Annual report of the EAVP Board

18:00 - 20:00 Annual EAVP Auction

9 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday July 8

Main Hall

Symposium ‘Early Tetrapods Awaken’ hosted by Sebastien Steyer, Josep Fortuny and Thomas Arbez

08:30 Struan Henderson: SHAPE-SHIFTING ENDOCASTS: 3D MORPHOMETRICS AND TRENDS IN PALAEOZOIC ENDOCASTS

08:45 Tom Challands: EARLIEST DIPNOI: POST-HANGENBERG RECOVERY AND THE DAWN OF A NEW ERA OF

09:00 Tim Smithson: PREDICTING THE LIKELY LOCATION OF NEW EARLY SITES: LESSONS FROM RECENT DISCOVERIES IN ROMER’S GAP

09:15 Jennifer Clack: MORPHOLOGICAL INNOVATIONS IN THE EARLIEST POST- TETRAPODS: ADAPTATIONS FOR TERRESTRIALITY?

09:30 Thomas Arbez: AN EXCEPTIONALLY WELL-PRESERVED OF STANOCEPHOSAURUS (: CAPITOSAURIA) LEADS TO A NEW HYPOTHESIS ON THE OPERATION OF THE TEMNOSPONDYL AUDITORY SYSTEM

09:45 David Marjanovic: TEMNOSPONDYLS, LISSAMPHIBIANS, AND MIDDLE EARS

10:00 Kamil Gruntmejer: COMBINED OSTEOHISTOLOGICAL AND COMPUTED FINITE ELEMENT ANALYSIS OF KRASIEJOWENSIS (AMPHIBIA, TEMNOSPONDYLI) SKULL BIOMECHANICS

10:15 Ella Teschner: HISTOLOGICAL STUDIES AS THE KEY TO POPULATION ANALYSIS – THE EXAMPLE OF METOPOSAURUS KRASIEJOWENSIS FROM KRASIEJOW, SE POLAND

10:30 - 11:00 coffee break 1 and poster session 2

11:00 Sanjukta Chakravorti: INDIAN METOPOSAURID : MORPHOMETRY, AND DISPERSAL

11:15 Dhurjati Prasad Sengupta: TEMNOSPONDYL AMPHIBIANS OF : THEIR DIVERSITY, DISPARITY AND NON-ENDEMISM

11:30 Sebastien Steyer: THE AMPHIBIANS STRIKE BACK – THE FIRST TEMNOSPONDYL FROM THE PALEOZOIC OF ZAMBIA

11.45 Josep Fortuny: SWIMMING WITH TEMNOSPONDYLS: A NEW APHANERAMMA (TEMNOSPONDYLI, TREMATOSAURIDAE) IN THE OF 10 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

12:00 Celeste Pérez Ben: MINIATURIZATION AND PAEDOMORPHOSIS IN TEMNOSPONDYL AMPHIBIANS

12:15 Armin Elsler: DIVERSITY AND BODY SIZE EVOLUTION IN PARAREPTILIA

12:30 - 14:00 lunch break and poster session 2

14:00 Lorenzo Marchetti: NON-DIAPSID EUREPTILIAN EVOLUTION: TRACK-TRACKMAKER CORRELATION OF AN IMPORTANT GROUP OF PALAEOZOIC TERRESTRIAL TETRAPODS

Symposium ‘Insular Vertebrate Palaeontology’ hosted by Lars van den Hoek Ostende and Paul Mazza

14:00 Paul Mazza: OVERSEA DISPERSAL TO MADAGASCAR: A GAMBLING MIRACLE?

14:15 Lars van den Hoek Ostende: THE INSULAR PARADOX

14:30 Judith Masters: EVOLUTION OF THE SOUTHERN OCEANS WITH A FOCUS ON MADAGASCAR AND ITS RIF-RAF INHABITANTS

14:45 Hanneke Meijer: A HYDROTAPHONOMIC MODEL FOR THE MARE AUX SONGES DODO-LAGERSTÄTTE IN MAURITIUS: MIRED IN TOXIC LOW- VISCOSITY MUD

15:00 Anneke van Heteren: THE FEEDING BIOMECHANICS OF HAAST’S EAGLE (HARPAGORNIS MOOREI)

15:15 - 15:45 coffee break 2 and poster session 2

15:45 Daniela Winkler: HUMAN IMPACT CHANGES WEAR OF MALAGASY BLACK RATS

16:00 Elehna Bethune: MULTI PROXY DIETARY TRAIT RECONSTRUCTION IN PLEISTOCENE HIPPOPOTAMIDAE FROM THE MEDITERRANEAN ISLANDS (CETARTIODACTYLA, MAMMALIA)

16:15 Maria Rita Palombo: LIGHTS AND SHADOWS IN THE EVOLUTIONARY PATTERNS OF INSULAR ELEPHANTINI

16:30 Andrea Savorelli: NEW UNEXPECTED BREAKTHROUGHS FROM THE MIOCENE ENDEMIC FAUNA OF THE APULIA PLATFORM (ITALY): A FAR MORE COMPLEX SCENARIO

16:45 Eric Buffetaut: THE GIANT BIRD GARGANTUAVIS: A CASE OF INSULAR EVOLUTION IN LATE CRETACEOUS EUROPE ?

17:00 Vlad Codrea: LATEST CRETACEOUS MICROVERTEBRATE DIVERSITY OF THE TRANSYLVANIAN LANDMASS

17:15 Oliver Wings: THE LATE JURASSIC MAMMAL ASSEMBLAGE FROM THE LANGENBERG QUARRY (LOWER SAXONY, ) 11 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Small Hall

Plenary

08:30 Hector Rivera: DIVERSITY OF MEXICAN CERATOPSIDS

08:45 Pim Kaskes: THE DUTCH TYRANNOSAURUS REX SPECIMEN AND ITS EXTRAORDINARY PRESERVATION

09:00 Christian Meyer: TRACKS FROM A CRETACEOUS ALLUVIAL PLAIN (AROIFILLA FORMATION, BOLIVIA)

09:15 Matteo Belvedere: THE FIRST UNEQUIVOCAL EVIDENCE OF GREGARIOUS BEHAVIOUR IN BABY SAUROPODS (LATE JURASSIC, NW SWITZERLAND)

09:30 Novella Razzolini: UNIQUE EVIDENCE FOR A BIMODAL ORIENTATION PATTERN IN MEGALOSAURID (THEROPODA: DINOSAURIA) TRACKWAYS ON A MIDDLE JURASSIC TIDAL FLAT

09:45 Diego Castanera: THE TETRAPOD TRACK RECORD FROM THE HUẾRTELES FM: AN EXCEPTIONAL WINDOW INTO THE LOWER CRETACEOUS ICHNODIVERSITY OF EUROPE

10:00 Mario Bronzati: SAUROPOD BRAINCASE ANATOMY AS A PRODUCT OF RAPID MORPHOLOGICAL TRANSFORMATION

10:15 Veronica Diez Diaz: A “LOOK BACK” IN PALAEONTOLOGY: RECONSTRUCTING THE TAILS OF NEOSAUROPOD

10:30 - 11:00 coffee break 1 and poster session 2

11:00 Koen Stein: PRELIMINARY RESULTS ON THE BONE OF HADROSAURS FROM THE LATEST CRETACEOUS OF FAR EASTERN RUSSIA.

11:15 Thierry Tortosa: NEW DISCOVERIES OF RHABDODON FROM THE CAMPANIAN OF PROVENCE (SE FRANCE) PART II: A BETTER UNDERSTANDING OF THE INTRASPECIFIC DIVERSITY

11:30 Eric Mulder: A DUTCH 'CRYSTAL PALACE' - MOSASAUR, AND DINOSAUR IMAGERY IN SUBTERRANEAN GALLERIES, SOUTHERN LIMBURG (THE NETHERLANDS)

11.45 Jordi Marce-Nogue: BIOMECHANICAL TRAITS OF CARNIVOROUS WITH FINITE ELEMENT METHODS

12:00 Soledad de Esteban-Trivigno: EVOLUTION OF SHAPE AND BIOMECHANICS IN CARNIVORES: A PHYLOGENETIC APPROACH

12:15 Heinrich Mallison and Matteo Belvedere:PALEONTOLOGICAL MASS 3D DIGITIZING: LEARNING BY DOING THE MFN’S DIGIS 2015 PROJECT ‘LIBERATION FROM THE BONE CELLAR’ 12 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

12:30 - 14:00 lunch break and poster session 2

14:15 Marton Rabi: MARINE TURTLES FROM THE LATE CRETACEOUS OF NE ITALY HIGHLIGHT THE MYSTERIOUS ORIGIN OF THE LEATHERBACK LINEAGE

14:30 Haiyan Tong: VERTICAL NECK RETRACTION IN AN EARLY STEM SIDE- NECKED TURTLE

14:45 Walter Joyce: MORPHOMETRIC AND POLYMORPHIC CHARACTERS AND THE PHYLOGENY OF GEOEMYDID TURTLES

15:00 Christian Foth: THE EVOLUTION OF CRANIAL DISPARITY IN FOSSIL AND RECENT TURTLES

15:15 - 15:45 coffee break 2 and poster session 2

15:45 Ursula Göhlich: REVISITING A CONTROVERSIAL FOSSIL BIRD: THE ALLEGED AUK PETRALCA IS A LOON

16:00 Dennis Voeten: SYNCHOTRON MICROTOMOGRAPHY REVEALS ADDITIONAL ELEMENTS IN THE NINTH SPECIMEN OF ARCHAEOPTERYX

16:15 Catalina Pimiento: THE PLIO-PLEISTOCENE MARINE

16:30 João Muchagata: SEXUALl DISPLAY AND ROSTRAL VARIATION IN EXTINCT BEAKED WHALE, GLOBICETUS HIBERUS

16:45 Ella Hoch: THE CONTRIBUTION OF PALAEONTOLOGY TO THE HISTORY OF THE ARABIAN CONTINENTAL PLATE (keynote)

17:15: Jeff Liston: LEVIATHAN RISING: THE BLEEDING EDGE OF RESEARCH CHASED OUT OF THE DEPTHS BY THE SHARK OF PUBLIC SECTOR DEADLINES

17:30 - 18:00 break and poster session 2

Teylers Museum Cafe

18:00 - 20:00 Annual EAVP Dinner

13 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday July 9

Main Hall

Plenary

08:30 Paul Barrett: THE SCIENCE OF ‘SOPHIE’ THE STEGOSAURUS: BUILDING A RESEARCH PROGRAMME AROUND A NEW ACQUISITION (keynote)

09:00 Suzie Maidment: LATITUDINAL BIODIVERSITY GRADIENTS IN DEEP TIME: A CASE STUDY USING THE DINOSAURS OF THE MORRISON FORMATION, WESTERN USA

09:15 Richard Butler: THE PERMO-TRIASSIC DIVERSIFICATION OF ARCHOSAUROMORPHS (keynote)

Symposium ‘Hominins’ hosted by Hanneke Meijer

09:45 John de Vos: introduction: DUBOIS: TRINIL, TEYLERS, TEGELEN

10:00 David Lordkipanidze: THE FIRST HOMO OUT OF (keynote)

10:30 - 11:00 coffee break 1 and poster session 3

11:00 Luc Amkreutz: DEFINING NEW BOUNDARIES? PALAEOLITHIC RESEARCH IN THE NETHERLANDS AND ADJACENT DOGGERLAND (keynote)

11:30 Adrian Lister: ‘NATIVITY’ IN THE BONE BEDS OF BETHLEHEM

11.45 Ahmet Aytek: A GENERAL OVERVIEW OF PLEISTOCENE HOMININ REMAINS FROM TURKEY

12:00 Berna Alpagut: 32 YEARS OF EXCAVATION IN PAŞALAR (TURKEY): THE RICHEST HOMINOID LOCALITY OF THE WORLD

12:15 Jordi Marcé-Nogué: BIOMECHANICS AS A PREDICTOR OF ENAMEL THICKNESS IN EXTINCT PRIMATES

12:30 - 14:00 lunch break and poster session 3

14:00 Mark Sier: THE FIRST MODERN HUMANS OUTSIDE AFRICA (keynote)

14:30 José Joordens: REVISITING TRINIL: GETTING TO KNOW HOMO ERECTUS (keynote)

15:00 roundtable discussion

15:15 - 15:45 coffee break 2 and poster session 3

14 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Plenary

15:45 David Unwin: THE REPRODUCTIVE BIOLOGY OF PTEROSAURS

16:00 Aude Cincotta: STUDY OF PRESERVED INTEGUMENTS FROM THE CRANIUM OF A TAPEJARID PTEROSAUR

16:15 Miguel Moreno-Azanza: HIGH CONCENTRATIONS OF TURTLE EGGSHELLS (TESTUDOOLITHIDAE) IN AN EOCENE DELTAIC PLAIN (SOBRARBE DELTAIC COMPLEX, NORTHERN SPAIN)

16:30 Massimo Delfino: THE RISE AND FALL OF THE EUROPEAN CROCODYLIANS (keynote)

Small Hall

Symposium ‘Vertebrates of the Triassic Germanic basin and their Allies’ hosted by Dennis Voeten and Melanie During

08:30 Martin Sander: THE EVOLUTION OF A UNIQUE BODY PLAN: NEW EVIDENCE FROM A TRIASSIC PLESIOSAUR AND LONG BONE HISTOLOGY (keynote)

09:00 Anna Krahl: MUSCLE RECONSTRUCTION AND FINITE ELEMENT ANALYSES OF HUMERI IN PLESIOSAURS AND SEA TURTLES BY DISSECTION OF THE LATTER

09:15 James Neenan: CRANIAL ANATOMY OF A NEW SPECIMEN OF YUNGUISAURUS LIAE AND TOOTH REPLACEMENT IN PISTOSAUROIDEA (DIAPSIDA, SAUROPTERYGIA)

09:30 Jelle Heijne: UNUSUAL TAPHONOMY OF THE MARINE BODY FOSSILS FROM THE LOWER OF WINTERSWIJK, THE NETHERLANDS

09:45 Melanie During: GEOCHEMISTRY OF SEDIMENTARY SEQUENCES FROM THE ANISIAN OF WINTERSWIJK WITH RELEVANCE TO UNDERSTANDING THE PALEO-ENVIRONMENTAL LIVING CONDITIONS OF SAUROPTERYGIANS

10:00 Lukasz Czepinski: PRELIMINARY REPORT ON RE-DISCOVERY OF THE FORGOTTEN TRIASSIC LOCALITY FROM SILESIA (POLAND) WITH REMAINS OF TERRESTRIAL ARCHOSAURS

10:15 Marco Marzola: THE REPTILIAN AND FAUNA FROM THE OF THE JAMESON LAND BASIN (EAST ): REVIEW AND UPDATES

10:30 - 11:00 coffee break 1 and poster session 3

Plenary 11:00 Ilja Nieuwland: THE TALE OF THE CROUCHING DINOSAUR. HOW DINOSAURS

15 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

WERE MADE IN EARLY 20TH CENTURY GERMANY

11:15 Kenneth Monsch: PRELIMINARY SURVEY OF THE MACKEREL-LIKE FROM THE OF THE "GRUBE UNTERFELD" (S. GERMANY)

11:30 Norbert Micklich : NEW AND RARE FOSSILS FROM THE LOWER OLIGOCENE OF THE GRUBE UNTERFELD ("FRAUENWEILER") CLAY PIT

11.45 Pim Kaskes: UNRAVELLING THE HISTORY OF A UNIQUE TRICERATOPS GRAVEYARD FROM EASTERN WYOMING, USA

12:00 Lukas Czepinski: ANATOMY AND STATUS OF BREVICERATOPS KOZLOWSKII - PROTOCERATOPSID DINOSAUR FROM THE LATE CRETACEOUS OF GOBI DESERT

12:15 Justyna Slowiak: THE APPENDICULAR SKELETON OF PROTOCERATOPS ANDREWSI – ITS ONTOGENETIC CHANGES AND IMPLICATIONS FOR LOCOMOTION

12:30 - 14:00 LUNCH BREAK AND POSTER SESSION 3

14:00 Kristof Veitschegger: BRAIN SIZE EVOLUTION IN CAVE BEARS (URSUS SPELAEUS)

14:15 Sarah Stinnesbeck : LATE PLEISTOCENE GROUND SLOTHS OF THE YUCATAN PENINSULA, MEXICO

14:30 Susanna Kummel: PREDOMINANCE OF PENTADACTYLY IN SPRAWLERS

14:45 Kristina Eck: FIRST RESULTS OF THE SCIENTIFIC EXCAVATION NETWORK IN THE CLAY PIT "UNTERFELD" (OLIGOCENE) AT RAUENBERG (GERMANY) - ABUNDANCE AND DIVERSITY

15:00 Jack Oyston: USING BIOGEOGRAPHY AND THE FOSSIL RECORD TO STUDY CONVERGENT EVOLUTION IN VERTEBRATES

15:15 - 15:45 coffee break 2 and poster session 3

Symposium ‘North Sea Basin’ hosted by Natasja den Ouden and Jelle Reumer

15:45 Natasja den Ouden and Jelle Reumer: introduction

16:00 Dick Mol: THE NORTH SEA: A TREASURE TROVE FOR PLEISTOCENE VERTEBRATE PALAEONTOLOGY AND ARCHAEOLOGY (keynote)

16:30 John Jagt: AN UPDATE ON MOSASAUR 'LARS' FROM THE TYPE AREA OF THE MAASTRICHTIAN STAGE, SOUTHEAST NETHERLANDS

End of EAVP Haarlem Scientific Conference

16 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Sunday July 10

Field excursions

(see field guides)

17 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Abstracts

Thursday plenary talks

TRACKING ARCHOSAURS IN SWITZERLAND - PAST AND PRESENT C.A. Meyer1

1 Naturhistorisches Museum Basel, Switzerland

*[email protected]

Keywords: Archosaurs, Dinosaurs, Triassic, Late Jurassic, Cretaceous

Before the discovery of Late Triassic prosauropod and theropod tracks in the Swiss National Park in 1961, dinosaur tracks were unheard of in Switzerland.

Close by, new discoveries - between 2006 and up to the present day - revealed more than six different levels with dinosaur tracks in the eastern part of the Swiss Alps. Apart from theropod tracks, tracks of advanced sauropods as well as prosauropod trackways have been discovered. These tracksites are from Ela unit (Late ) and show that prosauropods and advanced sauropods occur in coeval strata.

From the western Swiss Alps, near Lac d'Emosson, the autochthonous Triassic contains surfaces with hundreds of tetrapod footprints. They were attributed to dinosaurian trackmakers, and the age of the track-bearing strata was considered Late or . Recent studies have shown that these track-bearing strata form a megatracksite that contains only three different ichnotaxa attributed to archosaurs (Chirotherium ssp.) and not dinosaurs. Their age can be constrained to the (Anisian – Ladinian).

The first Late Jurassic tracks were found in a quarry near Lommiswil (Jura mountains; Kimmeridgian) in 1989, followed by a series of discoveries in the same stratigraphic unit. Altogether they form part of the first recognized megatracksite in Europe. This lead to the conclusion that these recurrent levels are tied to 3rd sea level fluctuations. From 2001 onwards, the construction of the Highway A 16 in the northwestern part of the Jura mountains produced a continuous stream of discoveries with multiple layers with dinosaurs tracks that are now the largest dataset of documented tracks and trackways anywhere in the world. The last discoveries were made two years ago, when the first track bearing levels formed in freshwater ponds were located (Oxfordian).

The only record comes from the Schrattenkalk Formation (/Albian) of Central Switzerland, where tracks and trackways of iguanodontid dinosaurs have been found, traditionally regarded as a completely submerged shallow shelf environment.

To sum this up, the renaissance in dinosaur ichnology triggered most of these discoveries and today Switzerland has become one of the key areas for the study of Late Jurassic dinosaur tracks worldwide. 18 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

NEW EVIDENCE FROM THE LOWER JURASSIC OF PATAGONIA SHEDS LIGHT ON THE HISTORY OF SOME OF THE EARLIEST KNOWN FROGS A.M. Báez1* and R.O. Gómez1

1Conicet-Iegeba, Departamento de Ciencias Geológicas, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Ciudad Universitaria, 1428 Buenos Aires, Argentina

*[email protected]

Keywords: , Jurassic, Patagonia, Notobatrachus

The early evolution of frogs is still poorly documented in the fossil record; a 50 My gap persists without any fossils between the oldest known salientians and the first frogs. According to previous research, the oldest known from the southern hemisphere was Vieraella herbstii from supposedly Toarcian–Aalenian beds of southern Patagonia, whereas this record was followed by those of Notobatrachus degiustoi and N. reigi, both of putative Callovian–Oxfordian age, also from Patagonia. New discoveries from the Lower Jurassic of Patagonia coupled with a better resolution of the chronological framework of the frog-bearing levels indicate that the diversification of frogs in this region took place earlier than previously thought. In the last decade, paleontological prospection of the lower Cañadón Asfalto Formation in the area of Cerro Cóndor, central Patagonia, has led to the discovery of many Notobatrachus-like frog remains in different localities. These fossiliferous levels transitionally follow the Lonco Trapial deposits related with the volcanism that affected this region during the Early Jurassic. Recent U-Pb radioisotopic dating places at least some of these levels in the middle-late Toarcian, including those bearing N. reigi. It is noteworthy that this age interval is close to the recent Pliensbachian–Toarcian U-Pb dating of the Kayenta Formation beds that yielded Prosalirus bitis, the supposedly oldest known frog. The new high-precision geochronology of the Patagonian volcanism linked to the western breakup, together with paleontological studies in progress, reveals the persistence of the lineage represented by Notobatrachus throughout nearly 20 My in water bodies associated with volcanic environments.

19 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

NEW ICHTHYOSAURS FROM THE EARLY CRETACEOUS OF COLOMBIA, WITH IMPLICATIONS FOR DIVERSITY IN THE PALAEOTROPICS E. Maxwell1, D. Dick1, S. Padilla1 and M.L. Parra1

1Staatliches Museum für Naturkunde, Rosenstein 1, 70191, , Germany; Centro de Investigaciones Paleontológicas (CIP), Villa de Leyva, Boyacá, Colombia

*[email protected]

Keywords: Early Cretaceous, Reptilia, Ichthyosauria, Colombia, Paja Formation

The Early Cretaceous was a dynamic time in Earth history, characterized by multiple phases of rapid climatic change and development of oceanic anoxia. However, our understanding of how these environmental changes affected marine vertebrate diversity dynamics is hindered by relatively poor sampling of marine tetrapods from this time interval, in particular ichthyosaurs. The Paja Formation (Villa de Leyva, Colombia) has the potential to greatly increase our understanding of ichthyosaur diversity in the Early Cretaceous of the palaeotropics. This marine black shale unit was deposited during the Barremian – Aptian interval, and yields a rich invertebrate fauna, fishes, turtles, plesiosaurs, a fragmentary dinosaur, and many ichthyosaurs.

A detailed examination and analysis of the ichthyosaur material suggests at least three valid taxa are present. The first, Platypterygius sachicarum, is currently known only from the holotype skull. Phylogenetic analysis places it within the Platypterygiinae, a subclade of Ophthalmosauridae, the monophyletic group including all other Cretaceous ichthyosaur taxa. The closely related platypterygiine Muiscasaurus catheti is also known only from a single specimen, but has a much more gracile rostrum. A third taxon is characterized by an extremely long, slender rostrum and very sharply pointed teeth of a morphotype not previously noted for Cretaceous ichthyosaurs. Unlike the previous two taxa, multiple individuals are known, including some postcranial material. This taxon also falls within the Ophthalmosauridae, but its exact phylogenetic position remains unresolved. A new specimen collected and prepared in 2015 may represent a fourth taxon, but requires more detailed study.

These results suggest relatively high taxonomic diversity and ecological differentiation manifested in jaw and rostral morphology among ichthyosaurs from the Cretaceous of Colombia. However, some questions remain to be resolved. Namely, all material originates from Arcillolitas abigarradas Member of the Paja Formation, which spans ~7 Ma: a non-trivial amount of time. Very few specimens have sufficient stratigraphic control, resulting in difficulties when making comparisons with coeval from other regions, as well as challenges in reconstructing whether these species truly co-occurred. Future work will emphasize the stratigraphic and palaeoecological context of this unique locality.

20 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

NEW INSIGHTS INTO THE EVOLUTION OF BASAL TURTLES W.G. Joyce1*

1 University of Fribourg, Department of Geoscience, Chemin de Musée 6, 1700 Fribourg, Switzerland

*[email protected]

Keywords: morphology, phylogeny, biogeography, Testudinata, Testudines

Turtles (Testudinata) have a rich fossil record that provides ample data to assess evolutionary hypotheses. Over the course of the last three decades, new fossil material coupled with detailed anatomical and phylogenetic analysis have therefore yielded important new insights into the evolution of the turtle stem lineage. The majority of phylogenetic analyses now agree that turtles (i.e., the diagnosed by the presence of the ) originated in the Late Triassic and that the crown clade originated in the Middle Jurassic. This topology implies that the pleurodiran type jaw closure mechanism evolved from the cryptodiran type closure mechanism, contrary to assumptions used in previous phylogenetic analyses. 3D modelling confirms that basal turtles were not able to retract their heads and necks within the shell using either of the two mechanisms seen among extant turtles, but they were nevertheless able to protect the head and neck by laterally tucking them below the shell. Taphonomy, histology, morphometrics, and morphological correlates indicate that stem turtles were predominantly terrestrial, which supports the notion that turtles originally evolved in a continental setting and that the aquatic nature of most crown turtles is derived. In parallel with the diversification of crown turtles, at least two of basal turtles, Sichuanchelyidae and Solemydidae, persisted to the Late Cretaceous whereas as third, Meiolaniformes, only went extinct as recently as the Holocene. Biogeographic analysis reveals that Sichuanchelyidae, Solemydidae, and Meiolaniformes were restricted to Asia, Euramerica, and southern Gondwana throughout their evolutionary history, which coincides with the primary distribution of crown turtles. However, given that the interrelationships of these three clades of basal turtles remains unresolved, it remains unclear if this pattern was caused by variance following the breakup of Pangaea or the result of differential extinction.

21 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Fossillegal

INTERNATIONAL LEGISLATION, THE SEMANTICS OF ‘CULTURAL PROPERTY’, AND WHY THEY ARE BAD FOR PALAEONTOLOGY AND PEOPLE J. Martin1*

1School of Museum Studies, University of Leicester, UK

*[email protected]

Keywords: Legislation, Cultural property, Ethics, Fossils, UNESCO

In our small world of palaeontology, it has ended friendships and ruined careers. Important fossils are lost to science, or in limbo. In the wider world, people get shot or imprisoned. Through it, fortunes are made by rich people in the West, while peasant farmers in the South lose the fortunes they never had.‘It’ is the UNESCO Convention ‘on the Means of Prohibiting and Preventing the Illicit Import, Export and Transfer of Ownership of Cultural Property’ of 1970: the origin of all this trouble.I will look back at the ancestry of ‘UNESCO 1970’, to suggest that its ethically-inspired progenitors probably would not have wanted it to turn out the way it did. The wrong turn seems to have been in ambiguity (perhaps intentional, certainly not articulated) about the meaning of ‘cultural property’ in the Convention’s articles. John Merryman, an academic lawyer, discussed ‘thinking about cultural property’ in 1986; he explained that it has two almost-opposite meanings, whose complex bases, in semantics, nationalism and money, I will explore. Stay with me, if you can, as I ruminate on how differently the word ‘culture’ might be understood in the minds of legislators (and politicians) in the Signatory Countries to the 1970 Convention, and speculate about how their interpretations might, from one point of view, be inadvertently (‘culturally’, ‘lost in translation’) mistaken and how, from another point of view, they might coincide neatly with national interests. Maybe ‘UNESCO 1970’ did turn out the way its authors intended. The purpose of the Convention’s Articles was to police international trade in national and personal property (in support of the principles of capitalism, as variously applied in badly, less, or only slightly corrupt regimes – and now, 46 years on, globally). Finally, I will question whether fossils should be in the Convention at all; I mean (except possibly for fossil hominins), whose ‘cultural’ property are they? The subtitle of this talk is really ‘what would I do if I ruled the world?’ But what would you do?

22 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

FOSSILS: NATIONAL MONUMENTS, TRADING OBJECTS, OR BOTH? E. Frey1*

1State Museum of Natural History Karlsruhe, Erbprinzenstraßes 13, 76133 Karlsruhe, Germany

*[email protected]

Keywords: fossils, law, national monument, trading object

The laws that regulate fossil trading get progressively restrictive. Recently, a new law restricting trade of national monuments was put into effect in Germany, with some fossils being listed as national monuments. However, this law had a fundamental problem: palaeontological objects were considered as archaeological objects or cultural monuments, and trading fossils was thus restricted or made impossible. After a shared national protest by private collectors, fossil dealers and palaeontologists, this has now been changed. In many countries, however, palaeontology is considered equal to archaeology, which makes fossil collecting and trading a criminal act and thus impossible. Sometimes, even research loans between scientific institutions are problematic and require huge amounts of paperwork and time.

Practical experience shows that either criminalisation or liberation of fossil trade results in massive disadvantages for the scientific community. Criminalisation fuels the black market and corruption. It boosts prizes for specimens, which mostly vanish into private collections and are then mostly lost for science. Complete liberation results in the exploitation of fossil sites by mass collectors and vandalism. In this scenario, fossils remain cheap as long as they are available, but mostly lack any accompanying scientific data.

Today, there is an international trend to criminalise private fossil collectors and dealers, and to control them with restrictive laws. On the other hand, many palaeontological institutions depend on private collectors and dealers, provided they follow scientific and ethical rules, which is sometimes difficult to control. However, in many countries in the world the laws are so restrictive that even scientific excavations and field studies begin to suffer, especially in multinational projects. Sometimes palaeontologists use legislative restrictions to exclude concurring teams from scientifically important sites. While the trade of endangered living species is legally regulated by an international convention (CITES), the international trade of fossils and minerals is not internationally regulated. An international regulation to this effect should balance both the scientific and trading demands. Is there a way out of this dilemma?

23 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

HANDS OFF, IT’S MY COLLECTION! FOSSILS, MUSEUMS AND THE PROBLEMS OF ACCESS D.M. Unwin1*

1University of Leicester, School of Museum Studies, 19 University Road, Leicester LE1 7LF, United Kingdom

*[email protected]

Keywords: Museum, fossils, access, hoarding

The science of palaeontology is founded on fossils, almost all the critical examples of which (type and figured specimens), and most of the others, are to be found in museums. Data collection, data verification, the construction of data sets and the testing of hypotheses are core processes of palaeontology that depend directly on these museum specimens. Access to specimens, of which there are already tens of millions scattered across several thousand museums, is therefore fundamental to the discipline and one of the most important factors that shape the science and, consequently, our understanding of the history of life. In an ideal world, access would be open to all, at any time, but the reality is that access is highly variable. At one end of the spectrum, enlightened curators and collections managers provide unfettered access to the entire collection and an array of facilities to support visitors. At the other end, access to particular specimens, or even entire collections, may be blocked for years or even decades and continue to have a highly negative impact on the development of ideas. Access may be restricted for practical reasons such as lack of space, or personnel, security concerns, or collections care issues. Not infrequently, however, those responsible for collections have used these and other excuses (‘the specimens are currently being worked on’) to defend the practice of hoarding. While such attitudes may seem justified, ultimately, they are self-defeating. Providing access to fossils not only facilitates palaeontology and adds value to collections, it is also consistent with a growing trend across museums, more generally, to democratise collections, and move away from past practices where they were the preserve of an elite privileged few. Natural history collections are starting to follow this trend. The development of ethical frameworks and the formalisation of access procedures are proving helpful, and digital technologies (e.g. high-resolution photos, CT scans, laser scanning, 3D prototyping) are beginning to play a major role. Much greater effort will be needed, however, if we are to realise the potential of collections that remain locked away because of outdated ideas and practices.

24 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

SHOULD PALAEONTOLOGISTS BEHAVE ETHICALLY? PROTECT THE FOSSILS OR PROTECT THE COMMUNITIES WHO DIG THEM? D.M. Martill1*

1University of Portsmouth, School of Earth and Environmental Sciences, Portsmouth, PO1 3QL, United Kingdom

*[email protected]

Keywords: Fossil collecting, legal frameworks, amateurs, artisan, commercial collectors

Some authorities argue that fossils should only be collected by licensed, professional scientists. This is an extreme view, but an attitude that has been adopted by many nations. Fossil collecting has become illegal for all but a small ‘elite’ of, not always competent, scientists in countries like China, Italy, Brazil and Canada. Germany is now considering adopting a similar approach to earth science conservation. It is misguided.

An alternative approach - the other end of the spectrum, is to have no legislation to ‘protect’ fossils whatsoever. This is almost the case in the United Kingdom. 19th century surgeons obtained cadavers from grave robbers to teach their students and to research their science in defiance of a plethora of laws protecting the dead. I urge palaeontologists to adopt a similar, and what may seem an unethical approach: to simply ignore legislation that claims to protect fossils. Such legislation is elitist, almost always undemocratic, and only serves to reduce the number of fossils that enter the scientific domain: it does not protect fossils; it leaves them in the ground to weather away.

In many countries fossil collecting provides an important source of income for communities struggling to make a living. In desert regions of southern Morocco the Kem Kem (mid Cretaceous) vertebrate fauna would be unknown were it not for a thriving commercial fossil trade. Also in Morocco, the palaeoichthyofauna of the Akrabou Formation (also Cretaceous) would be unknown were it not for the commercial digging of ammonites, where fishes, plesiosaurs and tethysaurs are a frequent bycatch.

When mining and erosion destroy billions of fossils yearly, laws that exclude amateur and commercial collectors appear quite pathetic. Such laws usually add a mountain of bureaucracy to scientific endeavour. They severely stifle the flow of fossils to museums and in some cases they encourage the development of a black market.

25 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

BRAZILIAN PALAEONTOLOGICAL LEGISLATION T. Rodrigues 1*and M.C. Langer 2

1Universidade Federal do Espírito Santo, Centro de Ciencias Humanas e Naturais, Departamento de Ciências Biológicas, Av. Fernando Ferrari, 514, 29075-910, Vitória, ES, Brasil 2Universidade de São Paulo, Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Departamento de Biologia, Av. Bandeirantes, 3900, 14040-901, Ribeirão Preto, SP, Brazil

*[email protected]

Keywords: Brazil, legislation, collection, export, commerce

Brazilian palaeontological legislation includes several laws and decrees that aim to protect fossils found within its territory. The Federal Constitution considers fossils both as goods belonging to the union (Article 20) and part of its cultural heritage (Article 216). As such, it is the government's duty to protect both fossils and fossiliferous sites (Article 23). Palaeontological sites are also specifically protected by a decree-law (4146) from 1942, which states that they belong to the country and that, as such, palaeontological excavations are controlled by the DNPM (Departamento Nacional de Produção Mineral), an authority under the Ministry of Mines and Energy. State and federal museums and universities do not need authorization to collect fossils, but must nevertheless communicate the DNPM of any excavations in course. Palaeontologists who work in private institutions, laymen, or foreigners need DNPM clearance to conduct excavations. In the case of foreigners, there is a decree (98830/1990) that guides the collection of scientific material, including fossils, in Brazil. Among others, it states that collections should be made as joint projects with Brazilian institutions (Article 3). There is also an ordinance (55/1990) from the Ministry of Science, Technology and Innovation (MCTI) on the administrative aspects of that decree, which states that all fossil type-material should be deposited in a Brazilian institution (Article 42). Indeed, exports (temporary or permanent) of fossils that were legally collected is not prohibited, but depends on authorizations from the DNPM and the MCTI. Brazilian law (8176/1991) considers unauthorized collection, commerce, and export crimes against the national heritage that can result in incarceration. Unfortunately, however, hundreds of fossils still leave the country every year unlawfully. A list and short summary of all Brazilian legislation on fossils can be found at the website of the Brazilian Society of Palaeontology, at www.sbpbrasil.org/pt/legislacao-brasileira.

26 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

COLLECTION AND PLACEMENT OF FOSSILS IN COMMERCIAL PALAEONTOLOGY: PRACTICAL AND ETHICAL CONCERNS A. Maltese1*

1Rocky Mountain Dinosaur Resource Center, 201 S. Fairview St., Woodland Park, Colorado 80863 USA

*[email protected]

Keywords: commercial palaeontology, ethics, legal, cooperation

Commercial and academic vertebrate palaeontology have an intertwined and sometimes inseparable relationship dating to the very founding of the science. Though animosity between the sides has grown in recent decades, the fact remains that many times commercial collectors working on private land recover scientifically new or important specimens. Increased education and cooperation has resulted in not only better collection techniques, but also advances in documentation and access. Recent preparation symposia have even invited and welcomed commercial collectors to share new techniques and equipment, an important step in working better together. While a number of commercial collectors vehemently adhere to the idea that they are the only people that can decide the ultimate fate of a specimen, several of the larger companies currently strive to ensure that important specimens are placed in the public trust by some method, whether by sale, donation or trade. This is in keeping with the collection guidelines set out by a major palaeontological society.

Though this is all better than nothing, much more can be done to improve the situation. Those participating in the illicit trade of specimens, that do substandard fieldwork and data collection, that have no regard for the final or long-term disposition of important specimens need to be held accountable. This goes for both commercial and academic palaeontologists. Lofty goals with overall poor execution do nothing to protect important palaeontological resources, and in fact may harm them. Attainable minimum standards in data collection, site access, specimen care and collections access should be developed and adhered to, and better avenues of communication would do more for the fossils and those that work with them than the increasingly exclusionary tones we experience today.

27 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

40 YEARS AS A FOSSIL PROSPECTOR, COLLECTOR AND EXHIBITION MAKER H. Siber1*

1Sauriermuseum Aathal, Switzerland

*[email protected]

Keywords: fossil, collection, private, public, legislation

Even 200 years of fossil collecting for the benefit of public and private collections has not exhausted the supply of fossils by any means. One can even argue that we have barely scratched the surface, and that the best and most important finds are still out there. The primary reason for this is that the technical means for excavating, collecting and preparing fossils have considerably advanced during the last three decades, up to the level that the present act of collecting is - under ideal circumstances - far superior to past collecting efforts. Who has the ambition, the skills, the financial resources and the legal rights to collect fossils properly?

Traditionally, three groups engage themselves in the art and technique of fossil collecting: the professional palaeontologists, the advanced amateurs, and the commercial palaeontologists. All these groups have made significant contributions to classical and modern palaeontology and to palaeontological museum collections worldwide.

Recent developments regarding restrictions to collect fossils primarily target the group of amateurs and so called ‘commercials’ or independent palaeontologists, and have greatly diminished the opportunities for legal fossil collecting. The three groups work best together, in my view, if they pool their resources. Many models of cooperation exist and have been proven beneficial for all parties involved. In addition, the restrictions on fossil collecting in more fossiliferous areas and countries is well meant, but, in my view, essentially counterproductive. Palaeontology requires the effort and input of all three groups: the scientists, the advanced amateur collectors and the ‘commercial’ collectors. Their cooperation is essential and eventually determines the amount and quality of important fossils found and saved for future generations. In my view, it is not necessary that scientifically important fossils are exclusively housed in “public repositories”. The criterion should be accessibility for continued scientific research and not ownership.

28 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

PUBLISH OR PROHIBIT? THE ETHICS OF PUBLISHING ON PRIVATELY-OWNED FOSSILS P.M. Barrett1*

1Department of Earth Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, UK

*[email protected]

Keywords: ethics, private ownership, accessibility, reproducibility, publication

Private collections vary in terms of size, scope and importance, ranging from those of hobbyists with small collections of common, widespread fossils to those of ‘professional’ collectors who accumulate spectacular, and often unique, examples of museum-quality objects. Private ownership of fossils is sometimes controversial, with some countries claiming all fossils as state property while others have a more laissez-faire approach. A complex debate surrounds this issue, which includes considerations of patrimony, scientific significance, public engagement, education and local economics. Among professional palaeontologists, attitudes to private collections have varied through time, between different cultural practices and between individuals, as has the level of engagement between university and museum-based scientists and avid private collectors. Current professional standards, as viewed through the lens of institutional ethical statements and journal editorial policies, generally discourage engagement between scientists and private collectors for several reasons. Firstly, there is a perception that private collectors make key specimens inaccessible for study and that by commercialising the subject they have helped to push many such specimens beyond the purchasing power of academic institutions. Secondly, and more importantly, issues of continuing access to specimens undermines the reproducibility of observations published on material in private hands. Finally, there is a widespread belief that fossils represent common cultural capital that should be accessible to all by being held in the public trust, rather than in private hands. However, there have been several high-profile examples of leading journals breaking these professional standards, by publishing on new, significant material held in private hands. Moreover, various counterarguments have been proposed to emphasise the importance of unlocking the scientific information held in private collections; these arguments also highlight the key role that technology can take in making virtual representations of physical objects more widely available than ever before. Here, I review the arguments on either side of this publication debate, discussing the pros and cons of publishing on privately-owned material, and conclude that, in most cases, the issue of scientific reproducibility should be the primary consideration when making ethical and editorial decisions.

29 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

FOSSIL LEGISLATION - PROTECTION OR DESTRUCTION? R. Albersdörfer1*

1Dinosaurierpark Altmühltal, Dinopark 1, 85095 Denkendorf, Germany

*[email protected]

Keywords: legislation, collection loss, specimen destruction, Archaeopteryx, private collections

Collecting fossils privately and unrestrictedly, possessing specimens and trading with them, has been a fundamental cultural achievement since the time of enlightenment, setting bases for the scientific advancement of generations of palaeontologists. Goethe’s mineralogical and palaeontological collection, for instance, was worth the equivalent of a million euros.

Recently, increasing numbers of countries have tried to protect fossils in general, or at least important specimens, to save these from private ownership and commercial activity, in the hope of securing their continuing accessibility for science. However, what seems reasonable at first glance has become a major force for fossil destruction. Daily, important fossils are destroyed – by industrial activity, erosion, or by the restrictive laws. Only a fraction is saved – in most cases not by institutional field work, but by committed private collectors and serious commercial palaeontologists and dealers. This is not a situation of competition with institutions, for most specimens will be lost anyhow. Each preserved piece, found by whoever, is better off being saved rather than being destroyed by stone crushing machines or erosion.

Most collectors will cooperate with open-minded scientists who respect their efforts and the private ownership of their finds. Moreover, most specimens in public museums actually originate from private or commercial collecting, eventually having found their way to public collections by donation, purchase, or inheritance.

So any law restricting private collecting and unlimited ownership will have unwanted effects.

Collecting will diminish and relentless destruction of millions of fossils by neglect, industrial activity or erosion will be the result.

Private collectors not willing to give up, nor willing to freely donate their energy and the money the hobby consumes, will be driven into illegality. None of their finds will be accessible for science. However, there are alternatives to draconian laws restricting collection and possession. In Bavaria, collecting and ownership has been basically unrestricted (with the exception of some Archaeopteryx specimens). This proves beneficial to all sides: scientists, museums, commercial dealers, and private collectors.

Let’s face the real problem of fossil destruction and not fight ideological wars against those who sacrifice time and energy to achieve what no public entity can ever deliver.

30 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

FOSSILS, PUBLISHING AND OPEN ACCESS N. Stead1*

1PLOS ONE, Carlyle House, Carlyle Road, Cambridge, CB4 3DN, United Kingdom

*[email protected]

Keywords: Publishing, Policy, Open Access, Open Data

In late 2012, concerned with the accessibility to fossils used in research and the legality of their collection, the open access journal, PLOS ONE, launched a set of editorial standards for manuscripts reporting palaeontology research. This policy requires that all authors provide information about where fossils are deposited for future access, details about the permits obtained, and confirmation that the fossils were collected legally. By adopting this editorial policy we aim to promote the reproducibility of palaeontology research and to discourage the illegal collection of fossils.

31 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

PRIVATE COLLECTIONS: PROBLEM OR BONUS FOR SCIENCE? O.W.M. Rauhut1*

1Bayerische Staatssammlung für Paläontologie und Geologie, Richard-Wagner-Str. 10, 80333 München, Germany

*[email protected]

Keywords: private collections, ethics, collaboration, scientific data

Private collections have a long tradition. Most natural history museums started with specimens obtained from private collectors, and acquisition of collections remains an important source of material. In recent decades, privately owned specimens have become controversial, and private collecting is being frowned upon by part of the scientific community. Associated questions are manyfold: Should collecting fossils privately be legal and under which regulations? Should privately owned specimens be considered in scientific studies? What should the relationship between private collectors and professional scientists be?

The first of these questions includes the purely legal aspect that has to be decided upon by lawmakers and varies between countries and sometimes even provinces. A more general question is whether private collecting is considered beneficial or harmful by the scientific community. As this aspect might influence the former - if scientific expertise is seeked in the lawmaking process - scientists should form an opinion. I argue that private collecting is essential to our science, as both staff and funding of public institutions are by far insufficient to explore and exploit fossil resources. Private collection - if properly done - thus helps to discover and preserve our fossil patrimony. Laws that are too restrictive might cause more harm than good. The second question mainly concerns the repeatability of scientific results, which is said to be endangered when privately owned specimens are concerned, so that many scientific journals do not accept scientific studies of such material. However, neither does inclusion in public collections assure their availability for study, nor is this argument valid from a philosophical viewpoint. Science is about discovering and understanding the natural world, and willfully ignoring available scientific data violates this principle. Repeatability is secondary, and is not given in many natural phenomena, the study of which is never questioned. Detailed documentation of the observed data is essential to this question.

Collaboration, not confrontation is thus the key in relationships between private collectors and professional scientists. We should seek to educate private collectors to keep high standards of data collection and repository, and in turn profit from their experience and the data they collect.

32 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

AN INNOCENT ACADEMIC ABROAD?: EASY FOSSILS AND HARD CHOICES, CHINA AS A MICROCOSM OF THE CHALLENGES OF 21ST CENTURY PALAEONTOLOGY J.J. Liston1*

1Yunnan University,Yunnan Key Laboratory for Palaeobiology, Kunming 650091, Yunnan Province, People's Republic of China

*[email protected]

Keywords: fossil legislation, ethics, China, impact factor, site protection

Fossil protection legislation exists for a variety of reasons, most obviously to restrict the irresponsible collection of fossil material, preserving natural science heritage from the unregulated ravages of museums, commercial dealers, private collectors, tourists and academics. Ongoing problems with the smuggling of fossils from China, Brazil and Mongolia have recently focused attention on this issue through a number of high profile media stories, though it affects many other nations. Perhaps surprisingly, the issues and problems surrounding this issue are fairly standard across a variety of territories, regardless of the political systems involved. Not simply restricted to the illegal fossil trade, this smuggling also extends to material that ends up as the subject of scientific study and publication in otherwise reputable journals.

Legislation has prohibited the export of fossils from Brazil since 1942, and unexcavated fossils (as cultural items) have been regarded as property of the Mongolian state since 1924. China similarly regarded fossils as generic unexcavated relics belonging to the state since 1930, but from 1982 has explicitly had a complex administration in place for vertebrate fossils (recently expanded to encompass invertebrate, palaeobotanical and trace fossils), which has to be engaged with in order for material to be excavated or leave China legally for study and eventual return. Unfortunately, although such paperwork appears to have had some impact on the illegal fossil trade (demonstrated by the decline in quantity of Chinese fossil material available at the Tucson Fair), it has also had two less desirable consequences for academic research: encouraging academic theft (to avoid the paperwork) and discouraging international collaboration by legitimate domestic research departments (for the same reason). As a consequence, in 2015 the Society of Vertebrate Paleontology revised its ethics statement to make clear that failure to acquire appropriate permits for either excavating or exporting material would be violating international law, leading to expulsion from the Society.

This presentation will consist of a case study from my time in China, to demonstrate how the allure of highly seductive fossils combines with our current dependence on high impact factor publications, to compel moral compromises in exchange for career survival.

33 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Towards a Biogeographic Synthesis for the European Early Pleistocene based on the Mammal Record

NEW DATA ON THE EARLY PLEISTOCENE VERTEBRATE ASSEMBLAGES FROM COPĂCENI (DACIAN BASIN, SOUTHERN ROMANIA) Ş. Vasile1*, M. Venczel2 and A. Petculescu3

1University of Bucharest, Faculty of Geology and Geophysics, Department of Geology, Laboratory of Paleontology, 1 Nicolae Bălcescu Avenue, 010041 Bucharest, Romania. 2Ţării Crişurilor Museum, 1-3 Dacia Avenue, 410464 Oradea, Romania 3Romanian Academy, Emil Racoviţă Institute of Speleology, Department of Geospeleology and Paleontology, 13-15 Calea 13 Septembrie, 050711 Bucharest, Romania.

* [email protected]

Keywords: Lower Pleistocene, Anura, Caudata, Squamata, Rodentia

Although only recently discovered and briefly studied, the Lower Pleistocene fluvial beds from Copăceni yielded one of the most diverse Pleistocene vertebrate assemblages described from Romania so far. Not only is the faunal list rich in number of taxa, but it also includes representatives of various vertebrate groups: several large herbivores (Mammuthus meridionalis, Stephanorhinus ?etruscus, Leptobos vallisarni, Praemegaceros pliotarandoides), a single carnivore (possibly hyaenid) , several rodents (Castoridae indet., Allactaga sp., Mimomys savini, Lagurodon arankae, Allophaiomys pliocaenicus), one insectivore (Desmana ?radulescui), several fishes (teeth similar in morphology to those of Recent Carassius, Squalius, Tinca, Rutilus, Scardinius, Esox and Silurus), a couple of anurans (Rana sp., Bufo sp.), one caudate (Triturus sp.), and one snake (Natrix tessellata).

Among the microvertebrates, the fishes and micromammals were the only groups represented by well-preserved and diagnostically reliable material (i.e. teeth), whereas the amphibians and snake material was fragmentary and consisted mostly in limb bones or vertebrae, hindering a more precise taxonomical assessment. Large amounts of sediment was screen-washed, in order to obtain better preserved amphibian and squamate remains from Copăceni, hoping to improve the knowledge on their taxonomic affinities.

This proved to be a successful endeavor, resulting in the discovery of better preserved and more diagnostic material that adds to the microvertebrate faunal list: the anurans Pelobates sp., Pelophylax sp. (both based on partial ilia), the salamandrid Lissotriton sp. (based on vertebral and dental material), the colubrid Coronella sp. (based on a partial ), the lacertid Lacerta sp. (based on dental and vertebral material), as well as an indeterminate murid (a fragmentary molar). Other previously reported taxa, such as Rana sp., identified on the basis of a fragmentary , were confirmed by the discovery of more diagnostic material (i.e. a partial ilium). All new discovered amphibian and squamate taxa represent first occurrences from the Pleistocene of the Dacian Basin.

The preliminary taphonomical analysis supports a parautochtonous accumulation of the large vertebrate remains, suggesting that the large terrestrial vertebrates lived nearby the river that hosted most of the herpetofauna. 34 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

TAXONOMICAL, TAPHONOMICAL AND PALEOECOLOGICAL ASPECTS OF A LATE EARLY PLEISTOCENE VERTEBRATE FAUNA FROM THE SOMSSICH HILL 2 SITE (SOUTH HUNGARY) P. Pazonyi1*, L. Mészáros2, Z. Szentesi3, M. Gasparik3, A. Virág1, K. Gere2, R. Mészáros2, D. Botka2, B. Braun2 and L. Striczky2

1MTA-MTM-ELTE Research Group for Paleontology, POB 137, 1431 Budapest, Hungary 2Eötvös Loránd University, Department of Palaeontology, Pázmány Péter sétány 1/C, 1117 Budapest, Hungary 3Hungarian Natural History Museum, Department of Palaeontology and Geology, Ludovika tér 2, 1083 Budapest, Hungary

*[email protected]

Keywords: small vertebrates, Early–Middle Pleistocene transition, Central Europe

The Somssich Hill 2 site is a 10 m deep karstic cavity situated in the Villány Mts (South Hungary), which yielded one of the richest late Early Pleistocene vertebrate assemblages of Central Europe. The age of the locality is around 950 ka based on ESR dating, but an approximately 200 kyr age difference were suggested between the lowermost and uppermost part of the section.

Small mammals (including bats, , voles, lemmings, hamsters, mice, dormice, squirrels, ground squirrels, pikas and hares) are the most abundant faunal elements, but frogs, lizards, snakes and birds are also frequent. A few remains of fish, , turtles and larger mammals (carnivores, most frequently mustelids and canids) were also found. Besides the vertebrates, snails are very common elements of the fauna in the upper part of the section. The fauna represents a mixture of Early Pleistocene relict taxa (including Beremendia, Dryomimus, Borsodia, Allophaiomys) and elements, which are characteristic for the Middle Pleistocene (e.g. modern Microtus species, such as M. nivaloides).

According to the taphonomical analysis, the frogs and snakes most likely fell into the cavity during their lifetime, whereas the small mammal remains most plausibly spent a short time period buried in shallow soil and then they were transported into the pothole by temporary water streams. The fact that the assemblage was accumulated by at least two different and partly selective processes, suggest that a few distortions might be present in the taphocoenosis compared to the biocoenosis once lived in the surrounding area, which must be taken into account in the palaeoecological interpretation of the locality.

Several different and recurring environments (from relatively dry steppe to humid climate with closed vegetation) were detected based on the abundance changes of the faunal elements within the section. However, hygrophilous frogs (such as Bombina variegata, Hyla arborea and Rana temporaria) are indicating that moist habitats were present in the close surroundings of the locality even in the driest intervals. Such mosaic and shifting environment represents well the climate and faunal changes during the Early–Middle Pleistocene transition.

This research was supported by the OTKA K104506 project.

35 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

DMANISI ARITODACTYL ASSEMBLAGE M. Bukhsianidze1*

1Georgian National Museum, 3 Purtseladze Street, 0105 Tbilisi, Georgia

*[email protected]

Keywords: Artiodactyla, Dmanisi, Early Pleistocene

Dmanisi is one of the richest and well dated (ca. 1.8 Ma) sites in the temperate zone of the Western Palaearctic. The faunal assemblage consists of 50 vertebrate taxa apart of Homo. Small mammals, most importantly the presence of typical Late Pliocene water vole Mimomys pliocaenicus, indicate a Late Villanian age and the large mammals correspond to the Late Villafranchian. Absolute and magnetostratigraphic dating bracket the entire fossil record of Dmanisi to 1.83 -1.76 Ma.

Artiodactyls in the Dmanisi fauna are represented by 13 taxa (5 cervids, 1 giraffid and 7 bovids): Pseudodama nestii, Eucladoceros sp., Praemegaceros obscurus, Arvernoceros insolitus, Cervalces gallicus, Palaeotragus priasovicus, Bison (Eobison) georgicus, Gallogoral meneghinii sickenbergii, Capra dalii, Soergelia cf. minor, Praeovibos sp., Pontoceros surprine, Antilopini indet. Among them Dmanisi Gallogoral represents the last and easternmost record of this mountain antelope in Eurasia; Cervalces gallicus record expands the distribution area of this species to the South-East; presence of Praemagaceros, Pontoceros, Soergelia in Dmanisi predates the North Caucasian and other European records by 250-500 thousands of years.

From the point of view of biogeography perhaps the most significant information in the Dmanisi fauna is simultaneous appearance of many new taxa characterizing Late Villafranchian of Europe (e.g.: Bison, Capra, Praeovibos, Soergelia, Pontoceros, Praemegaceros, Pseudodama). This artiodactyl assemblage documents the migration wave of mountain and those exploiting forest edge ecotones from the Inner Asia and Southern Siberia westwards. Incompleteness of fossil record, preservation and dating issues as well as presence of biotic barriers need to be explored as possible reasons of the discrepancies in timing and taxa representations.

36 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

TSIOTRA VRYSSI: A NEW VERTEBRATE LOCALITY FROM THE EARLY PLEISTOCENE OF MYGDONIA BASIN (MACEDONIA, GREECE) G.E. Konidaris1*, D.S. Kostopoulos2, G.D. Koufos2, V. Tourloukis1, and K. Harvati1

1Eberhard Karls University of Tübingen, Palaeoanthropology, Senckenberg Center for Human Evolution and Palaeoenvironment, Rümelinstr. 23, 72070 Tübingen, Germany 2Aristotle University of Thessaloniki, Department of Geology, Laboratory of Geology and Palaeontology, 54124 Thessaloniki, Greece

*[email protected]

Keywords: Villafranchian, Pleistocene, Mygdonia Basin, Greece

The Pleistocene vertebrate localities of Mygdonia Basin (Greece) are known since the end of the 1970s. Numerous fieldwork campaigns, carried out by the University of Thessaloniki, led to the discovery of several fossiliferous sites from which a great amount of fossils has been unearthed and studied. During a survey expedition conducted by the University of Thessaloniki and the University of Tübingen in 2014 a new locality –named Tsiotra Vryssi (TSR)– was discovered. The collected material includes the corvid Corvus pliocaenus (Aves), the hyaenid Pachycrocuta brevirostris, the canid Canis etruscus, the rhinocerotid Stephanorhinus sp., two species of Equus, the bovid Bison sp., the cervid Metacervocerus rhenanus and the giraffid Palaeotragus sp. The locality was further excavated in 2015 and the fauna was significantly enriched by the mammoth Mammuthus meridionalis, the bovid Leptobos sp., Antilopinae indet., and the giant cervid Praemegaceros sp. The presence of Pachycrocuta brevirostris clearly indicates a late Villafranchian age for TSR, while the presence of two Equus species, Praemegaceros sp., and the co-occurrence of Leptobos and Bison, suggest that TSR is chronologically intermediate between Gerakarou-1 and Apollonia-1 (Mygdonia Basin), and therefore it can be preliminary dated to 1.8–1.2 Ma. The TSR fauna will provide additional palaeogeographical and palaeoecological information about a crucial time interval for mammal migrations and turnovers, as well as for the first hominin dispersals from Africa to Europe.

Acknowledgement: This research was supported by the ERC-STG-283503 (“PaGE”).

37 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

KOCAKIR-2: FIRST RECORD OF EARLY PLEISTOCENE CANIS AND HIPPOPOTAMUS FROM BURDUR BASIN (SW TURKEY) F.A. Demirel1*, S. Mayda2**, M.C. Alçiçek3, T.T. Kaya4 and A.I. Aytek5

1Mehmet Akif Ersoy University, Department of Anthropology, 15030, Burdur, Turkey 2Ege University, Faculty of Science, Department of Biology, Bornova, 35100 Izmir, Turkey 3Pamukkale University, Department of Geology, 20070, Denizli, Turkey 4Ege University, Natural History Research and Application Center, Bornova, 35100 Izmir, Turkey 5Mehmet Akif Ersoy University, Department of Anthropology,15030, Burdur, Turkey

*[email protected] **[email protected]

Keywords: Villafranchian, Turkey, Canis, Hippopotamus, Palaeoecology

Kocakır-1 locality (37042’13.1” N, 30019’41.2” E, 1100 m.) located 5 km east of Burdur town (SW Turkey), was discovered by one of us (MCA) in 1999 during a geological survey. The first limited specimens, collected from the fossiliferous layer were described as Hipparion sp. and Giraffidae indet. Recently, Alçiçek et al. revised the fauna by allocating the Hipparion to Equus sp. and Giraffid to Paracamelus gigas. They also described a new specimen as a medium-sized deer (Cervidae gen. indet.). The revised fauna allows the confirmation the age close to that of the European early late Villafranchian (early Pleistocene) for uppermost reaches of the Burdur Basin succession.

During the new project, led by A.D. with the permission granted by the Turkish Ministry of Culture and Tourism, we conducted four short field seasons from 2012 to 2015 to attempt to explore systematically the new fossil potential of the Kocakır region. Consequently, a new fossil-bearing section to the 1.5 km to the southwest of original locality has been found during the field season of 2015, named here as Kocakır-2. Whilst the Kocakır-2 locality is quite richer in species than Kocakır-1, the fossiliferous levels in both localities are stratigraphically comparable.

Here, we will present the tentative results from the preliminary survey of Kocakır-2, producing a fairly diverse fauna reflecting an open habitat comprised of a limited standing body of water. The fauna includes two equids of different size (Equus cf. altidens and Equus sp.), a small rhino (Stephanorhinus cf.etruscus), a giant Camel (Paracamelus gigas), a panthera (the size of P. gombaszoegensis), a wolf-like dog (Canis etruscus), a Hippo (Hippopotamus sp.) and a medium sized Bovid (Bovidae indet.). The well-preserved cranial material of a medium–sized canid and a canine of Hippopotamus represent the first discovery of Villafranchian wolf-like dog and Hippopotamus records respectively. Faunal composition of both Kocakır localities is indicative for the European early late–latest Villafranchian faunas. Therefore, Kocakır-1 and 2 are candidate to be well placed chronologically between Gerakarou- Greece (early late Villafranchian ∼1.9 Ma) and Livakos-Greece (middle late Villafranchian, ∼1.6Ma).

38 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

SYSTEMATIC, BIOSTRATIGRAPHIC AND PALEOBIOGEOGRAPHIC REEVALUATION OF THE MAMMALIAN FAUNAS FROM THE TURKISH VILLAFRANCHIAN LOCALITIES OF KAMISLI, GULYAZI AND Y.SOGUTONU S. Mayda1*, M.C. Alçiçek2, T.T. Kaya3, A. Demirel4, K. Halaçlar3 and D. Kanık4

1Ege University, Faculty of Science, Department of Biology, Bornova, 35100 Izmir, Turkey 2Department of Geology, Pamukkale University, 20070 Denizli, Turkey 3Ege University, Natural History Research and Application Center, Bornova, 35100 Izmir, Turkey 4Mehmet Akif Ersoy University, Faculty of Art & Sciences, Department of Anthropology, Burdur-Turkey

*[email protected]

Keywords: Kamisli, Y.Sogutonu, Gulyazi, Villafranchian, Turkey

Afyon-Gülyazı, Eskişehir-Y.Söğütönü and Amasya-Kamışlı are three Pleistocene localities which were found in late 1960’s by a group of German and Turkish geologists and the result of the first determinations were announced in subsequent papers. The German group had transferred most of the collections to Hannover Geology Department and very few fossils were kept in Turkey, in “MTA” Museum, Ankara. Up to now, hyenids of Kamışlı, Hipparion taxa of Gülyazi and recently elephantids of Y.Söğütönü (NOW, 2016) were studied while the rest have not yet received any attention. This presentation aims to summarize the history of the aforementioned collections and give preliminary updates of the ongoing systematic studies on the original Hannover collections.

The medium-sized Camelid from Gülyazi, which was originally attributed to Paracamelus alexejevi, actually shows great similarities to a younger form; P. alutenesis. Additionally, Y.Söğütönü material could also be attributable to this species. We prefer to allocate Gazella sinensis material of Gülyazi to G. bouvrainae. The rhino and Equus materials of Gülyazı are missing, although an earlier horse, E. stenonis, in the collections was briefly described in an unpublished report. The equid of Kamisli which was listed as E. stenonis shows typical features of E. altidens. An interesting molar from Y.Söğütönü belongs to an advanced Hipparion, here described as Plesiohipparion sp. The canid of Kamışlı which was originally assigned to C. etruscus is now considered C. mosbachensis because of its morphological and metrical resemblance. While the giraffid from Gülyazı shows clear similarities with Sesklo material, we keep the original attribution to Mitilinatherium martinii. Taking into consideration the revised biostratigraphical data, we prefer to correlate the Kamışlı fauna to latest Villafranchian, and Y.Söğütönü and Gulyazı to late Middle Villafranchian, suggesting younger dates for all.

39 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

BIOGEOGRAPHIC ORIGINS OF EARLY PLEISTOCENE MAMMALIAN FAUNA IN THE SOUTHERN LEVANT M. Belmaker1*

1The University of Tulsa, Department of Anthropology, 800 South Tucker Drive, Tulsa, Oklahoma, USA.

*[email protected]

Keywords: Levant, Homo, Ubeidiya, biogeographic corridors

The Southern Levant is a unique biogeographic entity within southwestern Asia. This land bridge, between Africa and Eurasia, emerged during the Miocene as a continuous terrestrial belt that allowed various plants and taxa to migrate through in either direction. The region is known for the eclectic fauna originating from Eastern and Western Palearctic and Ethiopian realms.

It has been suggested that discrete dispersal events took place during the early Pleistocene between Africa and the southern Levant. These pulsed dispersals were driven by climatic and environmental changes that led to the opening and closing of biogeographic barriers between the two regions. Oscillations between aridity and humidity in North African deserts and changes in lake size and location in the Sinai and Negev may have allowed for faunal dispersal from East Africa into the Levant during the humid periods and not during the dry periods.

In particular, it has been suggested that early Homo dispersed into the Levant with a suite of other African taxa during periods of an open biogeographic corridor. However, recent studies have pointed to the continuous human dispersals into the Levant rather than pulsed dispersal events.

This study presents a taxonomic and biogeographic revision of the African taxa in the early Pleistocene of the Southern Levant and studies the spatial temporal pattern of the appearance of African taxa, including early Homo.

Results suggest that several taxa, which were previously considered as African taxa, should be considered endemic or Palearctic in origin. Furthermore, the spatial and temporal distribution of African taxa in the southern Levant is continuous and new African taxa appear in the Southern Levant throughout the entire studied time span. In addition, there is no clear pattern of pulsed dispersals in the region and it appears that mammalian fauna, including Homo, moved freely between sub Sahara Africa and the Southern Levant. Furthermore, there is no correlation between the appearance of Homo populations in the southern Levant and the appearance of African taxa. Thus, the impetuses for dispersals may have been motivated by inter - intra species competition rather than climatic conditions.

40 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

PALEOBIOGEOGRAPHY AND CHRONOLOGY OF STEPHANORHINUS ETRUSCUS (FALCONER, 1858) (MAMMALIA, RHINOCEROTIDAE) IN EURASIA L. Pandolfi1*, E. Cerdeño2, V. Codrea3 and T. Kotsakis1

1University of Roma Tre, Department of Sciences, Largo S.L. Murialdo 1, 00146 Roma, Italy 2Paleontología, IANIGLA, CONICET-Mendoza, Avda. Ruiz leal s/n, 5500 Mendoza, Argentina 3University Babeș-Bolyai, Department of Geology, 1 Kogălniceanu Str., 400084 Cluj-Napoca, Romania

*[email protected]

Keywords: Late Pliocene, Early Pleistocene, paleobiogeography, dispersal, extinction

The occurrence of rhinoceroses during the Plio-Pleistocene has been well documented in Eurasia; several species have taken a considerable biochronologic, palaeoenvironmental and palaeobiogeographic importance since the second half of the 20th century. Among others, the so-called Etruscan rhino, Stephanorhinus etruscus, is one of the most abundantly recorded and better known Eurasian Early Pleistocene rhinoceros. Nevertheless, the first and last appearances of this species, as well as its paleogeographic distribution, are controversial and debated in literature. S. etruscus is documented since the latest Pliocene in Spain (Las Higueruelas, Villaroya), Italy (Montopoli), France (Perrier-Les Etouaires), Slovak Republic (Nová Vieska) and Romania (Iarăș-Cariera Veche). Its presence in the Early Villafranchian assemblage of Villafranca d'Asti (Italy, Late Pliocene) is here considered doubtful because the remains were collected from an undefined stratigraphic level. The rhinocerotid skull from Bethlehem (Palestine; latest Pliocene), assigned to "Dicerorhinus etruscus", displays morphological features (e.g., absence of ossified nasal septum) closer to Dihoplus.

During the Early Pleistocene, S. etruscus occurred in several Spanish, French and Italian localities as well as in The Netherlands (e.g., Tegelen), Germany (e.g., Thiede), Greece (e.g., Aivaliki), Georgia (e.g., Dmanisi), Israel (e.g., Ubeidiya) and maybe Anatolia (Denizli Basin). The last appearance of S. etruscus in Eurasia is debatable. The rhinoceros remains from Pietrafitta (Italy, late Early Pleistocene), previously classified as S. cf. hundsheimensis, are here referred to S. etruscus based on cranial and postcranial morphologies. Etruscan rhino populations survived till the Jaramillo subchrone (around 1.1 Ma) in France (Bois-de-Riquet), Romania (Betfia XII) and Hungary (Osztramos 2 and 8), and close to the Early-Middle Pleistocene transition in Spain (Cueva Victoria, Huéscar 1, Atapuerca TD4, TD6 and TD8) and Italy (Monte delle Piche). Around 1.2 Ma, S. hundsheimensis also occurred in Europe and seems to be coeval with S. etruscus during the latest Early Pleistocene, but the two species did not occur at the same localities. S. hundsheimensis has been recorded in Spain (Vallès-Penedès basin), Italy (Leffe basin level 5), France (Vallonnet), Germany (Untermassfeld), Slovak Republic (Gombasek), Romania (Betfia V, Brașov, Feldioara Cetate and Carieră, Rotbav-D. Țiganilor, Subpiatră) and Greece (Platanochori-1).

41 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

SOME BIOLOGICAL FEATURES OF ELASMOTHERIUM CAUCASICUM (MAMMALIA, RHINOCEROTIDAE) FROM THE LATE EARLY PLEISTOCENE OF EASTERN EUROPE V.S. Baigusheva1, G.I. Timonina1 and V.V. Titov2*

1Azov Museum-reserve, Moskovskaya str, 38/40, 346780, Azov, Russia 2Institute of arid zones of SSC RAS, Southern Scientific centre of RAS, Chekhov str, 41, 344006, Rostov-on-Don, Russia

*[email protected]

Keywords: fossil rhinoceros, teeth ontogenesis, ecology, Early Pleistocene

Elasmotherium species are typical representatives of Early Pleistocene faunas of Eastern Europe. They are not known from contemporaneous localities of Western Europe, which indicates the paleogeographic similarities between Eastern Europe and synchronous associations from Western and Central Asia. Elasmotherium caucasicum Borissiak, 1914 was described by material from Sinyaya Balka, (Taman Peninsula, Russia), where remains of the rhinoceros were found together with numerous bones of Archidiskodon meridionalis tamanensis. This late Early Pleistocene locality is the type locality for the Late Villafranchian – Early Biharian Taman theriocomplex of Eastern Europe. In 2004-2014, the site Sinaya Balka/Bogatyri was studied in collaboration with archaeologists from the IMCH RAS (St. Petersburg). They found tools of the Oldowan type in the fossiliferous layers. There are more than 230 elasmotherii specimens in the collection of the Azov museum-reserve. There are incomplete skull, lower jaws, isolated teeth, as well as postcranial bones. The combined skeleton of E. caucasicum was mounted here. Its shoulder height is 2.4 m, length – 4.2 m. From last year's excavations, 11 individuals of Elasmotheriums of different ages (by mandibles) were found. The ontogenetic changes in lower dentition were determined for adult and subadult individuals.

Deciduous dp4 functioned together with p2-p3 and m1-m2. Tooth m3 at this stage formed, or had just started to erupt. After the full erasing of dp4 it was shifted on p4. At this stage the falling out of p2 and the appearance of m3 happened. The active using of m2 started. At the stage when p4 and m1 were middle erased, the p3 came out. Further, p4 and m1 were primarily worn until the complete disappearance of internal enamel structures. At this age the main mastication load is transferred to m2 and m3. We conclude that adult animals often have no p2 and p3. The overgrown alveolus of deciduous incisor is fixed at 8-11 years old individual. The permanent incisors in Elasmotherium are not known. Dental microwear analysis of E. caucasicum from the Sinaya Balka locality shows a high number of scratches and a relatively high number of pits on M2 enamel. This type of enamel microwear is typical for mixed feeders herbivores. The elasmotherii enamel differs from other rhinoceros by the presence of more numerous scratches.

42 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

LATEST EARLY PLEISTOCENE LARGE MAMMAL ASSEMBLAGES FROM THE IBERIAN PENINSULA AND THE EARLY-MIDDLE PLEISTOCENE TRANSITION J. Madurell-Malapeira1*, D.M. Alba1, V. Vinuesa1, A. Boscaini2, S. Bartolini3, I. Rufí1,4, S. Ros-Montoya5, M.P. Espigares5, P. Palmqvist5, L. Rook3, B. Martínez- Navarro6,7,8 and S. Moyà-Solà1,8

1Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, Carrer de les Columnes s/n, Campus de la UAB, Cerdanyola del Vallès, 08193 Barcelona, Spain; 2 Departamento de Geología y Paleontología, Instituto Argentino de Nivología Glaciología y Ciencias Ambientales (IANIGLA), CCT-CONICET-Mendoza, Avda. Ruiz Leal s/n, Parque Gral. San Martín, 5500 Mendoza, Argentina; 3Dipartimento di Science della Terra, Università degli Studi di Firenze, Via G. La Pira 4, 50121, Firenze, Italy. 4Àrea de Prehistòria, Universitat de Girona, Pl. Ferrater Mora 1, 17071 Girona, Spain; 5Departamento de Ecología y Geología, Facultad de Ciencias, Campus Universitario de Teatinos, 29071, Málaga, Spain;6IPHES, Institut Català de Paleoecologia Humana i Evolució Social, Campus Sescelades, Edifici W3, 43007, Tarragona, Spain; 7Àrea de Prehistòria, Universitat Rovira i Virgili (URV), Avda. Catalunya 35, 43002 Tarragona, Spain; 8ICREA, Barcelona, Spain

* [email protected]

Keywords: Early Pleistocene, Iberian Peninsula, late Villafranchian, Epivillafranchian, Vallpara, díssection.

Since the early 1990s, as a consequence of the first findings of human remains and lithic artifacts from the late Early Pleistocene of Europe, the interest for large mammal assemblages from this time interval has increased dramatically, both from a biochronological and taphonomic point of view. In this regard, the Iberian Peninsula has overall the most comprehensive record of late Early Pleistocene mammals from Europe, including numerous well-dated fossiliferous sites, such as those from the Guadix-Baza and Banyoles-Besalú basins, the Vallparadís section, and the Sierra de Atapuerca. Two distinct faunas (late Villafranchian and Epivillafranchian) are customarily recognized among the large mammal assemblages from the late Early Pleistocene of Iberia, with a boundary between these two biochrons placed at ca. 1.2 Ma.The research carried out during the last decades indicates that a protracted faunal turnover event began at ca. 1.2 Ma, involving the extinction of typically Villafranchian taxa and the arrival of new Galerian immigrants. This faunal renewal started with the first appearance of the cervids Megaloceros savini and Dama vallonnetensis, replacing the earlier Praemegaceros verticornis and Metacervocerus rhenanus, respectively. At the same time, the suids of the Sus scrofa lineage are first recorded, while the latest members of Ursus etruscus are replaced by the earliest representatives of the cave bear lineage; Ursus deningeri. Subsequently, around 1.0 Ma the last representatives of Megantereon whitei and the first representatives of Panthera fossilis are recorded. Finally, after MIS22, the last typically Villafranchian species (i.e., Mammuthus meridionalis, Pachycrocuta brevirostris, Puma pardoides and Lycaon lycaonoides) are recorded during MIS21, roughly coinciding with the earliest records of Crocuta crocuta just before the Bruhnes-Matuyama magnetostratigraphic boundary.In summary, the large mammal record from the Iberian late Early Pleistocene, covering a protracted time span (from 1.6 to 0.8 Ma), evidences the progressive renewal of the large mammal communities since 1.2 Ma to 0.8 Ma, which was probably associated with the change in Earth's climate from orbital obliquity to eccentricity forcing that resulted in the onset of high-intensity glacial cycles and the concomitant paleoenvironmental instability that took place during the Early-Middle Pleistocene Transition. 43 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

BIOGEOGRAPHICAL AND PALAEOECOLOGICAL ASSOCIATIONS OF WESTERN PALAEARCTIC BISON POPULATIONS I. Maniakas1* and D.S. Kostopoulos1

1Aristotle University of Thessaloniki, Faculty of Sciences, School of Geology, GR-541 24 Thessaloniki, Greece

*[email protected]

Keywords: Bison, Pleistocene, ecomorphological adaptations

The current study outlines a comprehensive biogeographical framework of feeding and locomotor adaptations among Bison populations of the Western Palearctic, revealed by inferences from dental wear and postcranial ecomorphological traits. Different populations from the chronological continuum of European Pleistocene can be attributed to distinct size classes and shape trends, representing ecological variants across regional provinces and time intervals for more than 1.5 million years. Body mass and relative limb shape of bisons exhibited significant alterations driven by climatic instability and a series of biological aspects. Through the progressively cooler and more arid climate conditions of the 1.5-0.7 Ma time frame, a rapid increase of body mass and a significant distal limb elongation are evidenced, followed by a more gradual similar trend, which seems to reach its maximum in the heavy-built priscoid forms of the last interglacial. The restoration of a more temperate climate and the return of woodlands, after the last glacial climatic deterioration, led to the general diminution and the limb shortening at the Upper Pleistocene/Holocene transition, as seen in the extant Holarctic bisons. The phenotypic diversity triggered by climatic regionality and landscape-vegetational heterogeneity, is also imprinted in limb structures. Pleistocene bisons were primarily operating in relatively open and dry environments, suited for grazing but they were able to exploit a variety of open and fairly wooded habitats, consuming a broadened vegetational spectrum. The increased arid character of the south-eastern part of Epivillafranchian Europe forced to a more grazing specialized feeding mode and a restricted vegetational spectrum, giving periodically harsh soil conditions. Three distinct bison morphotypes mirror a gradually tripartite biogeographic segregation in the Epivillafranchian Europe: the lightly built and slender limbed primitive forms of the Lower Pleistocene, presented by the Venta Micena bison (Spain), Bison degiulii (Italy) and the bison from Dmanisi (Georgia), in comparison to a more massive bison from Apollonia (Greece) and the large sized Bison menneri (Germany) with its extremely elongated limbs.

44 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

DENTAL MICROWEAR TEXTURE ANALYSIS OF EARLY PLEISTOCENE CERVIDS: A PALEOENVIRONMENTAL APPROACH AT THE EUROPEAN SCALE E. Berlioz1*, D. Kostopoulos2, C. Blondel1 and G. Merceron1

1iPHEP – UMR 7262 (CNRS & University of Poitiers), 6, rue Brunet – Bât. B35 – TSA 51106 – 86073 Poitiers Cedex 9, France 2Faculty of Science – School of Geology, Department of Geology, 54124 Thessaloniki, Greece

*[email protected]

Keywords: Dental Microwear Texture Analysis (DMTA), Cervids, Paleoenvironments, Diet, Pleistocene

Early Pleistocene cervids are abundant and diversified in Europe, with several sympatric taxa (Eucladoceros ctenoides, Croizetoceros ramosus, Metacervoceros rhenanus) displaying different body-sizes and most likely different feeding habits. These deer are widespread during the Early Pleistocene as they occur from Western to Eastern Europe. The fact that these species are present in distant European localities does not necessarily mean, however, that these two geographic areas share the same habitat characteristics. Indeed, some extant herbivore species are known to display high plasticity, allowing them to adapt their feeding behavior to the vegetal resource availability in the habitat rather than to feed on selected vegetal species. The extant Red Deer Cervus elaphus, for example, shows a high dietary plasticity dependent on the local vegetal availability in its habitat that ranges from the cold coniferous forests of Norway to the dry Mediterranean forests of Algeria.

In this study, we will show the dietary variations of the European Early Pleistocene cervids through time and space, during the 2-1 Million years interval by comparing Western and Eastern European areas, thanks to the Dental Microwear Texture Analysis (DMTA). DMTA studies the microscopic wear caused by the food mastication and reflects the physical properties of the food items. It has already proven to be very efficient in deciphering subtle dietary differences among herbivores.

We will show notably that Eucladoceros ctenoides from Western Europe is more engaged in grazing, while the eastern representatives of the species are more involved in browsing. These results highlight the dietary plasticity of this extinct taxon and underline differences in vegetal resource availability between the Balkans and Western Europe, to be interpreted in terms of paleoenvironmental differences.

45 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

EARLY PLEISTOCENE SMALL MAMMALS OF EASTERN EUROPE AND WESTERN ASIA: BIOCHRONOLOGY AND BIOGEOGRAPHY A.S. Tesakov1*

1Geological Institute of the Russian Academy of Sciences, Pyzhevsky 7, 119017 Moscow, Russia

* [email protected]

Keywords: Rodentia, Early Pleistocene, Eurasia, biochronology

The detailed Early Pleistocene biostratigraphic sequence of Eastern Europe allowed to define eight biochrons based on concurrent ranges of arvicolids. Late Villanyian (Gelasian) time range overlaps four biochrons (MNR3-MQR11), the Early Biharian (Calabrian) interval matches another four biochrons (MQR10-MQR7). The Late Villanyian small mammalian faunas of the Northern Black Sea region through southern Western Siberia are specific in a generally more abundant rhizodont lagurines (Borsodia) adapted to open steppe-like landscapes as compared to synchronous faunas of Central and Western Europe dominated by mesophilous Mimomys species. The late Villanyian small mammals of western Anatolia show a combination of central European mimomyian assemblage, Asian lagurines, and local Aegean-Anatolian endemic lagurines of the Kalymnomys. Faunas of the Northern Caucasus importantly differ from the steppe faunas and show typical European mesic assemblages with Mimomys gr. pliocaenicus, Pitymimomys, and Clethrionomys. The latest Villanyian composition of arvicolid faunas is surprisingly similar in Northwestern Europe (Tegelen), western Anatolia (Bicakci), and North Caucasus (Psekups). The important datum of the trans-Holarctic migrational arrival of basal rootless Microtini (the genus Allophaiomys) defines the onset of Biharian and is variably dated from an interval immediately preceding the Olduvai Subchron (ca 2.1-2.0 Ma) to a time slice immediately postdating the Olduvai (ca. 1.8-1.7 Ma). The important basal biochron of the Early Biharian (MQR11) is characterized by the co- occurrence of the earliest Allophaiomys deucalion and advanced hypsodont Borsodia. The autochthonous (non-migrational) transformation of rhizodont Borsodia into rootless Prolagurus s.l. defines the the base of MQR10. The Early Bicharian arvicolid faunas of Eastern Europe and Western Asia show parallel adaptive radiations of rootless lagurines (with half of its diversity eliminated by by the end of Early Pleistocene) and the most successful Microtini which by the beginning of Middle Pleistocene elaborated most extant species groups of Microtus s.l. The Early Pleistocene fauna of circumpolar higher latitude boreal zone is documented only in few localities and includes Lemmus, Predicrostonyx, and specialised Microtini.

46 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

ADDRESSING THE 'ALLOPHAIOMYS GAP' IN BRITISH BIOSTRATIGRAPHY: A NEW EARLY PLEISTOCENE FAUNA FROM WESTBURY CAVE, SOMERSET, ENGLAND N.F. Adams1,2*, D.C. Schreve1, I. Candy1 and R.W. Barendregt3

1Centre for Quaternary Research, Department of Geography, Royal Holloway University of London, Egham, Surrey, TW20 0EX, UK 2Department of Earth Sciences, Royal Holloway University of London, Egham, Surrey, TW20 0EX, UK 3Department of Geography, University of Lethbridge, 4401 University Drive, Lethbridge, Alberta, T1K 3M4, Canada

*[email protected]

Keywords: Early Pleistocene, Mammalia, Matuyama reversed chron, Waalian 'interglacial', Westbury Cave

There are relatively few Early Pleistocene fossil localities in Britain and most are restricted to the East Anglian Crag Basin. Confounding this problem, the shallow marine to estuarine Crag Group contains many unconformities and terrestrial mammal fossils are rare and often reworked. A major unconformity in Crag Basin stratigraphy (from ca. 1.7 to 0.7 Ma) is often referred to as the ‘Allophaiomys gap’ due to the absence of the arvicolid Allophaiomys, a widespread genus in continental Europe at this time.

Westbury Cave in Somerset, south-west England, is one key site outside East Anglia that has purportedly yielded Early Pleistocene fossils, although few attempts have been made to date the deposits. Previous excavators reported sparse mammal fossils from the sands and gravels of the Siliceous Member, including Allophaiomys, but no detailed sedimentological descriptions or systematic bulk sampling were undertaken. The site therefore had unexplored potential to reveal the nature of the terrestrial fauna and palaeoenvironment during this unknown interval in Britain.

Excavations in 2014 have revealed new fossils belonging to six mammalian families (Castoridae, Arvicolidae, Hystricidae, Hyaenidae, Cervidae, Bovidae), including genera and species previously unknown from the Early Pleistocene of Britain and north-west Europe. New palaeomagnetic dating indicates deposition of the Siliceous Member during the Matuyama reversed chron (2.58-0.78 Ma), and biostratigraphical analysis suggests refinement to ca. 1.8-1.1 Ma. Moreover, palaeoenvironmental inferences correlate the Siliceous Member with the Waalian ‘interglacial’ (ca. 1.5-1.3 Ma) of western continental Europe. This is the first time fossils of this age have been reported from Britain.

47 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

A NEW ARVICOLID RODENT FROM THE EARLY PLEISTOCENE OF THE BAZA BASIN, SOUTHERN SPAIN R.A. Martin1*, A.S. Tesakov2 and J. Agustí3

1Murray State University, Department of Biological Sciences, 102 Curris Center, Murray KY 42071, USA 2Geological Institute of the Russian Academy of Sciences, Pyzhevsky 7, 119017 Moscow, Russia 3Institut Català de Paleoecologia Humana i Evolució Social, 43007 Tarragona, Spain

*[email protected]

Keywords: Arvicolinae, Rodentia, Early Pleistocene, Spain

A new, large arhizodont arvicolid rodent is described from early Pleistocene deposits of Barranco del Paso and Barranco de los Conejos in the Baza Basin of southern Spain. The molars of the new arvicolid are very hypsodont but with sparse cementum, undifferentiated enamel and a modified pachyknem schemlzmuster. With “Tibericola” vandermeuleni and Mimomys oswaldoreigi, the new arvicolid represents one of the earliest experiments with arhizodonty among European voles. Including Mimomys medasensis at Barranco del Paso, this set of arvicolids refines the composition of a previously described early Pleistocene MmQ1 biozone in Spain.

48 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

EARLY PLEISTOCENE ARVICOLINES FROM ZUURLAND (THE NETHERLANDS): NEW INSIGHTS INTO HOLARCTIC BIOCHRONOLOGY AND BIOGEOGRAPHY T. van Kolfschoten1* and A.S. Tesakov2

1Leiden University, Faculty of Archaeology, 2333 CC Leiden, the Netherlands 2Geological Institute of the Russian Academy of Sciences, Pyzhevsky 7, 119017 Moscow, Russia

*[email protected]

Keywords: Arvicolinae, Rodentia, Early Pleistocene, Holarctic, the Netherlands

The unique Pleistocene biostratigraphic sequence of the Zuurland is unmatched in Northern Europe in the number of directly superposed faunal associations and the quality of micromammal record. The Zuurland arvicoline record is summarised from the multiple boreholes produced and studied by the renowned Dutch amateur geologist Mr Leijn Hordijk (Brielle). The most complete is the Early Pleistocene sequence divided into faunal units 5, 4, 3. Units 5 and 4 represent two successive stages of Tiglian, Late Villanyian, MN17 assemblage with dominant rhizodont mimomyine voles. Especially diverse is the Unit 4 association with Lemmus kowalskii, Clethrionomys kretzoii, Ungaromys dehmi, Phenacomys sp., Borsodia newtoni, Mimomys pliocaenicus, Mimomys tigliensis, Mimomys hordijki, Mimomys reidi, and Pitymimomys pitymyoides. It documents a fauna integrating the classical European Late Villanyian forest and meadow assemblage of M.pliocaenicus, and a higher latitude Holarctic assemblage with Mimomys hordijki, Lemmus kowalskii, and Phenacomys sp., Borsodia newtoni and Mimomys tigliensis, as well as a ground squirrel Spermophilus cf. primigenius, indicate the direct influence of the East European and West Siberian faunal assemblage adapted to more open landscapes. The Zuurland Unit 3 with Allophaiomys deucalion, Mimomys savini, Mimomys ex gr. reidi-pusillus, Lemmus kowalskii, Clethrionomys kretzoii, Craseomys aff. major evidences the early Biharian, early MQ1 fauna that existed right after the major restructuring of the European arvicoline fauna that was possibly driven by the rapidly propagated Holarctic dispersion of Allophaiomys. The fauna of Unit 3 also combines typical Early Biharian Allophaiomys – Mimomys - Clethrionomys association with some high latitude (Lemmus) and eastern boreal (Craseomys) elements.

49 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

FAUNAL DYNAMICS IN SW EUROPE DURING THE LATE EARLY PLEISTOCENE: IMPLICATIONS FOR THE BIOCHRONOLOGICAL ASSESSMENT AND CORRELATION OF MAMMALIAN FAUNAS M.R. Palombo1*

1Sapienza University of Rome, Department of Earth Sciences, P.le A Moro 5, 00185 Roma, Italy - IGAG-CNR, Via Salaria km 29,300, 00015 Monterotondo (Roma), Italy

*[email protected]

Keywords: Biochronology, Pleistocene, Large Mammals, SW Europe.

During the late Early Pleistocene, the climate forcing known as Mid-Pleistocene Revolution (MPR) induced deep, more or less gradual alterations and latitudinal displacements in European terrestrial biomes and exerted great influence on dispersal and dispersion of mammalian species. Large mammals did not generally move in multi-species waves of dispersal, rather each species changed its range depending on the suitability of environmental conditions in respect to its own environmental tolerances and ecological flexibility. Factors driving the remodelling of the range of a taxon, and time and mode of its dispersal and diffusion into SW Europe differed from species to species as from one territory to another, leading to diachronicity/asynchronicity in local first appearances/lowest local stratigraphical occurrences. As a result, correlations and biochronological assessments of local faunal assemblages may be difficult especially when firm chronological constraints are unavailable. Whether the peculiar composition of the late Early Pleistocene fauna (characterized by the persistence of some Villafranchian species and discrete new appearances of some taxa that will have persisted during the Middle Pleistocene) may be indicative of any high rank biochron is discussed. Evidence from SW Europe suggests that the chronological range of the “Epivillafranchian biochron” (whatever its biochronological rank could be) might span from about 1.5 (i.e. Lowest local Stratigraphical Occurrence, LlSO datum, of, among others, Homo, Xenocyon lycaonoides, Canis ex gr. C. mosbachensis, Megantereon whitei, advanced stenonoid horses, Praemegaceros, Bison) and 0.85 Ma (Highest local Stratigraphical Occurrence, HlSO datum, of X. lycaonoides, M. whitei, Puma pardoides), roughly corresponding to a hypothetical X. lycaonoides-M. whitei “range-zone”. Nonetheless, several factors [e.g. confusing taxonomic treatment of some taxa, heterogeneous consistency of the fossil record in space and time (i.e. number and richness of local faunal assemblages) , taphonomical biases, apparent dyachronicity/asynchronicity of LlSO of some species that dispersed toward Western Europe during the post-Olduvai Early Villafranchian] make it difficult any attempt to formalize the “Epivillafranchian biochron". All things considered, several lines of reasoning suggest an informal use of the term Epivillafranchian, pending a complete revision of the Villafranchian ELMA, particularly the number of its subdivisions and their rank.

50 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

THE LATE VILLAFRANCHIAN SITES OF ORCE (SPAIN) AND THE EARLIEST HUMAN RECORD OF WESTERN EUROPE B. Martínez-Navarro1, 2, 3*, S. Ros-Montoya4, M.P. Espigares4, J. Madurell- Malapeira5, T. Medin2, 6, J. Jiménez-Arenas7, J.M. García-aguilar4, A. Guerra- Merchán4, L. Rook8, and P. Palmqvist4

1IPHES, Institut Català de Paleoecologia Humana i Evolució Social, Campus Sescelades, Edifici W3, 43007, Tarragona, Spain; 2Àrea de Prehistòria, Universitat Rovira i Virgili (URV), Avda. Catalunya 35, 43002 Tarragona, Spain; 3ICREA, Barcelona, Spain 4Departamento de Ecología y Geología, Facultad de Ciencias, Campus Universitario de Teatinos, 29071, Málaga, Spain; 5Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, Carrer de les Columnes s/n, Campus de la UAB, Cerdanyola del Vallès, 08193 Barcelona, Spain; 6National Museum of Eritrea, P.O. Box 5284, Asmara, Eritrea; 7Departamento de Prehistoria y Arqueología, Facultad de Filosofía y Letras, Universidad de Granada, Campus Universitario de Cartuja C.P. E-18011 Granada, Spain 8Dipartamento di Science della Terra, Università degli Studi di Firenze, Via G. La Pira 4, 50121, Firenze, Italy.

*[email protected]

Keywords: Early Pleistocene, Spain, late Villafranchian, Guadix-Baza basin, Orce

The Orce region (Guadix-Baza basin, Andalusia, Spain), has a continuous Early Pleistocene stratigraphic record, starting from the Middle Villafranchian site of Fuente Nueva 1, dated 2.2-2.0 Ma. The Middle-Late Villafranchian boundary, marked by the first appearance of the Ovibovini Praeovibos, is well recorded at Barranco de los Conejos. The Late Villafranchan site of Venta Micena, dated to ca. 1.5-1.6 Ma, is recorded in a continuous layer covering more than one square kilometer and filled with skeletal remains of large mammals, with a density of more than 50 fossils per m2 accumulated by the giant, short-faced hyena, Pachycrocuta brevirostris. This site shows interesting faunal similarities with the Georgian paleoanthropological site of Dmanisi, such as the record of the ovibovine Soergelia minor. Aforementioned fauna is marked by the arrival of the African origin, dirk-toothed cat Megantereon whitei, which is also recorded in all of these Late Villafranchan localities and persists until the Epivillafranchian (ca. 1.0 Ma). Furthermore, the site of Venta Micena is marked by one of the earliest records in the European continent of the megaherbivore Hippopotamus antiquus, a species that originated in Africa as well. Up to now, no record of human presence has been found in this locality.

Two important archeopaleontological localities are recorded in a stratigraphic position above Venta Micena, Barranco León-5 (ca. 1.4 Ma) and Fuente Nueva-3 (ca. 1.3 Ma). A deciduous tooth of Homo sp. was found at Barranco León, which nowadays represents the earliest human fossil of Western Europe. At both sites, a rich lithic artifacts assemblage is associated with abundant fossil remains of several large mammal species, such as Mammuthus meridionalis, Hippopotamus antiquus, Praemegaceros verticornis, and Bison sp., among others. A faunal break in the large mammals record between Venta Micena and the Orce localities placed above, Fuente Nueva-3 and Barranco León-5, is marked by the arrival of Equus sussenbornensis and Ammotragus europaeus, and the extinction of S. minor.

No evidence of Sus gr. scrofa, the principal biomarker of the Epivillafranchian faunas, which is first recorded in Spain at 1.2 Ma, is found in any of the Early Pleistocene sites of Orce. 51 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

BEFORE AND AFTER EARLIEST HOMO DISPERSAL IN EUROPE: EVIDENCE FROM THE EARLY PLEISTOCENE SITES OF THE ITALIAN PENINSULA R. Sardella1,2*, L. Bellucci2,3, F. Bona4, M. Cherin2,5, D.A. Iurino1,2 and L. Rook6

1Dipartimento di Scienze della Terra, Sapienza Università di Roma, Piazzale A. Moro, 5 00185 Roma, Italy 2PaleoFactory, Sapienza Università di Roma, Piazzale A. Moro, 5 00185 Roma, Italy 3Polo Museale Sapienza, Sapienza Università di Roma, Piazzale Aldo Moro 5, 00185 Roma, Italy 4Dipartimento di Scienze della Terra, Università degli Studi di Milano, via Mangiagalli 34, 20133 Milano, Italy 5Dipartimento di Fisica e Geologia, Università di Perugia, Via A. Pascoli, 06123 Perugia, Italy 6Dipartimento di Scienze della Terra, Università degli Studi di Firenze, Via G. La Pira 4, 50121 Firenze, Italy

*[email protected]

Keywords: biochronology, mammals, Pleistocene, Italy

The timing and pattern of Homo dispersal and earliest occupation of Europe is the focus of an ongoing scientific debate. The recent discoveries in Italy, France and Spain, and the consequent increase of available evidence suggest that humans dispersed into Western Europe during the Early Pleistocene.

In Italy the earliest occurrence of humans has been documented in the site of Pirro Nord (Apulia, South Italy), where lithic artefacts have been found together with a diversified late Villafranchian vertebrate assemblage dated around 1.6-1.3 ma. The Pirro Nord assemblage shares similarities in age and composition with those from Monte Argentario (Tuscany) and Pietrafitta (Umbria), where no evidence of human presence has been documented up to now. Older faunal assemblages (approximately between 2.1 and 1.6 ma) from Italy have been investigated in recent years. Together with the revision of “classical” sites such as Olivola and the Upper Valdarno localities an increasing number of data is now available from new excavations in central Italy at Coste San Giacomo (Latium) and Pantalla (Umbria) where diversified vertebrate faunas have been found. The Italian sites provide information of crucial importance to depict an updated and detailed biochronological framework for the middle and late Villafranchian of Europe and for reconstructing the evolution of the Early Pleistocene terrestrial ecosystems.

52 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

PALEOBIOGEOGRAPHIC AND ECOLOGICAL CONTEXT OF EARLY HUMAN DISPERSALS IN WESTERN EURASIA R. Croitor1,2*

1Maison méditerranéenne des sciences de l'homme, LAMPEA, F-13094 Aix-en-Provence, France; 2Institute of Zoology, Academy of Sciences of Moldova, MD-2028 Kishinau, Moldova

*[email protected]

Keywords: Hominines, paleoecology, scavenging, limiting factors

Early human dispersals from Africa are often regarded in the context of ecological belonging of archaic Homo to the carnivore guild (as a scavenger) and its commensalistic relationship with saber-toothed cats Megantereon and Homotherium. The scavenging hypothesis meets several contradictions: the hominid eco-physiology is ill-adapted for the scavenger ecological niche because of limited sense of smell capacities that makes hominids poor competitors with carnivorans, and the specific physiology of thermoregulation (sweating) that exposes hominid presence to ambushing saber-toothed cats. Scavenger niche capacity depends of ecosystem productivity, which normally is declining from low to high latitudes and thus limits diversity and dispersal of scavengers in boreal latitudes. Therefore, new scavenging opportunities for hominids in higher latitudes of Eurasia sound doubtful. The earliest human fossil findings in Western Eurasia are located south of the Alpine-Himalayan Mountain Belt (AMB), which also acted as a biogeographic border during Pliocene and Pleistocene for several herbivorous and mostly omnivorous mammal systematic groups and species (Theropithecus, Mitilanotherium, Cervus nestii, Ursus thibetanus, etc.). In this case, Homo ex gr. erectus differs from ubiquitous carnivorans. The sharp annual seasonality that characterized Early Pleistocene climate of Western Eurasia in the north of AMB could be a limiting factor for human dispersal. Paradoxically, hominine dispersal advanced to the north in Western Europe after climate deterioration ca. 1.0 Ma. The use of fire could be among the factors that permitted to overcome AMB. The Movius Line, which demonstrates a technological difference between the early prehistoric tool technologies in East and West of the Old World and generally coincides with AMB with exception of its western part, is an indirect evidence of the early Middle Pleistocene hominine dispersal in Western Eurasia. The area of human distribution in Western Europe during early Middle Pleistocene also acted as natural zoogeographic refugia for some Villafranchian holdovers, mostly cervids.

53 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

LARGE MAMMALS AFFECTED BY HOMININS: A BIOGEOGRAPHIC SYNTHESIS OF BUTCHERING FOR THE EUROPEAN EARLY AND MIDDLE PLEISTOCENE K. Pawłowska1*

1Adam Mickiewicz University, Institute of Geology, ul. Maków Polnych 16, 61-606 Poznań, Poland

*[email protected]

Keywords: large mammals, taphonomy, Early Pleistocene, Middle Pleistocene, Europe

In the last few decades, some progress has been made towards a synthesis of the data on the presence of early hominins in Europe and their dispersals across the continent in the Early and Middle Pleistocene. The sites that have been documented present various datasets, including hominin fossils, large and small mammal remains, and archeological artifacts.

In this paper, the main focus will be on sites where clear evidence exists of the processing of large mammals by hominins, in the form of cut marks, percussion marks, and others. In this regard, the taxonomic diversity of the mammals will be considered alongside the type of hominin activity. Processing, even in its simplest forms, could have been a strong influence on adoption of meat eating among early members of the Homo genus, as has been discussed recently.

The distribution of all sites, both well-known localities and those more recently discovered (such as Bełchatów and Sitkówka in Poland), will be compared between the European Early and Middle Pleistocene to determine any patterns. An attempt will be made to discuss the possible factors behind such patterns with reference to paleoanthropological and archaeological records, as well climatic inference.

This presentation presents a biogeographic synthesis of butchering for the European Early and Middle Pleistocene and summarizes our current understanding of food processing by hominins, by scrutinizing the data on large mammals affected by such processes.

54 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Fossillegal poster V1

BETTER TOGETHER: AMATEUR AND PROFESSIONAL PALAEONTOLOGISTS IN THE NETHERLANDS N. den Ouden1, 2* and R. Pouwer1, 3

1Naturalis Biodiversity Center, Postbus 9517, 2300 RA Leiden 2Werkgroep Pleistocene Zoogdieren (WPZ) 3Werkgroep voor Tertaire en Kwartaire Geologie (WTKG)

*[email protected]

Keywords: amateurs, citizen science, collections, collaboration, research

In the Netherlands there is a long tradition of amateur and professional palaeontologists working together in collecting and documenting fossils, making observations and publishing articles on them. Many amateurs are dedicated, accurate and sometimes innovative collectors, bringing together important and often unique private collections. In fact, a majority of palaeontological finds are actually collected by amateur palaeontologists, and considerable parts of museum collections originate from private collections. The large number of accessible open-air sites makes it possible for amateurs to build their own collections, while information supplied by professional palaeontologists ensures the quality of its documentation. The benefits of this system are twofold: the amateurs are able to contribute to scientific research and the professionals gain an extra set of eyes and ears in the field. Some amateurs are even doing research on a professional level themselves.

Many palaeontologists, whether amateur or professional, collaborate in associations such as the Werkgroep Pleistocene Zoogdieren (WPZ, Working Group for Pleistocene Mammals) and the Werkgroep voor Tertiaire en Kwartaire Geologie (WTKG, Working Group for Tertiary and Quaternary Geology). These associations organise regular meetings, field trips and workshops, and publish journals and information sheets. Natural history museums in the Netherlands, including the Naturalis Biodiversity Center, often work closely with such amateur associations. The nature of their efforts varies from writing scientific publications and conducting fieldwork to public outreach programmes, citizen science, the organisation of international conferences and dedicated access to collections.

Although the word amateur still carries a negative connotation for some professional researchers, amateur palaeontologists can and should continue to play a vital role in the scientific community. The impact of future contributions does however rely on professionals recognising the valiant efforts of amateurs and working together with them on an equal basis. Such collaborations not only open up new sources of information but, above all, it can be fun and inspiring to work with enthusiastic people living and working outside of the academic sphere.

55 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Fossillegal poster V2 SITE-SPECIFIC LIMITATIONS ON THE USE OF PALAEONTOLOGICAL RESOURCES C.J. Underwood1*

1Birkbeck College, Department of Earth and Planetary Sciences, WC1E7HX, London

*[email protected]

Keywords: conservation law, erosion, fossil collecting

Fossil sites may be damaged by indiscriminate collecting, with important specimens being lost to science or destroyed. As a result, many countries and regions have laws to protect geological heritage from damage through unregulated collection. In many cases, however, laws may be having the opposite effect. Laws may prevent the extraction and study of fossils, prevent amateur enthusiasts from carrying out their hobby, and prevent outreach and educational collecting. Conversely, such regulations may result in fossils being lost to erosion, weathering and large scale commercial damage (quarrying etc.) prior to collection, and may act to exclude workers from outside certain institutions or countries.

The degree of and type of conservation of fossil sites should vary with the situation of a particular site, and this should dictate collecting guidelines, which themselves should be a basis of legal protection. In sites where fossils are destroyed extremely rapidly as they are exposed, such as within rapidly eroding sea cliffs and active quarries, any form of collecting should be viewed as rescuing the fossils from destruction, and in these circumstances commercial collectors based near the site should be viewed as essential to recovering material. Conversely, in very slowly eroding sites where fossils can remain exposed for long periods of time without undue damage, such as wind ablated desert floors, large fossils should be left in situ until they can be carefully extracted.

Account should also be made for the type of collecting and size of fossils present. Excavation of a large vertebrate can cause considerable damage to a site whilst this is not the case with smaller material. Very small fossils, including microvertebrates, are readily destroyed on exposure, even in very slowly eroding sites. Where these can be recovered by bulk sampling without damaging the site, collection should be encouraged as the potential scientific benefits may vastly outweigh any damage done.

It should also be noted that palaeontology is critical to science outreach and education. School and other educational collecting of common fossils should be considered as extremely important and allowing for this should be considered a priority.

56 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

‘Towards a Biogeographic Synthesis for the European Early Pleistocene based on the Mammal Record’ poster R1

THE PRESENCE OF MAMMUT BORSONI IN THE PLIOCENE AND PLEISTOCENE OF OLTENIA (SW OF ROMANIA) A. Popescu1*

1Museum of Oltenia Craiova, Natural Sciences Department, Popa Sapca Street 8, 200424 Craiova, Romania

*[email protected]

Keywords: Mammut borsoni, Oltenia, Pliocene, Pleistocene

This research documents the distribution of Pliocene/Pleistocene mastodons from the territory of Oltenia, Romania. The region of Oltenia occupies the south-west of Romania and is framed by the river Olt (on the Eastern side), by the river Dunarea in the South and by the Meridional Carpathian arch in the North and West. Up to this moment, from the region of Oltenia, two species of mastodons have been discovered: Mammut borsoni and Anancus arvernensis. They have been discovered in rock formations of either Pliocene or Pleistocene age.

The Oltenia Museum in Craiova, Romania, holds material belonging to both species. Also, other museums and collections across Oltenia, as well as literature have been consulted.

M. borsoni occurs isolated or in association with the other mastodon (A. arvernensis), and only very rarely associated with other species of mammals. Thus far, all the fossil samples of M. borsoni discovered on the territory of Oltenia pertain to molars. No fossils belonging to the postcranial skeleton of this species have been inventoried until now. The presence of Anancus arvernensis is better documented on the territory of Oltenia. It occurs both individually and associated with other species. In its case, besides the molars, there have been discovered parts of the postcranial skeleton (for instance femora, humerus, belonging to the exemplar of A. arvernensis near Stoina, discovered in 1928 and described by C. S. Nicolaescu Plopsor in 1929). This exemplar has been the most complete mastodon skeleton discovered in Romania (that is, until 1974). Only a small number of pieces belonging to these species have been described until now, and there are, in the warehouse of the Oltenia Museum (or in that of other institutions), several pieces which have never been studied. My intention is to shed light on all the occurrences of mastodons on the territory of Oltenia and to update the map of the distribution of these species. For Romania and for Oltenia in particular, the two species of mastodons are important from the palaeographic, the paleo-ecological as well as from the stratigraphic point of view.

57 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

‘Towards a Biogeographic Synthesis for the European Early Pleistocene based on the Mammal Record’ poster R2

EVOLUTION OF MAMMOTH MOLAR ENAMEL AND ITS BIOCHRONOLOGICAL IMPLICATIONS A. Virág1* and Ş. Vasile2

1MTA-MTM-ELTE Research Group for Paleontology, Pázmány Péter sétány 1/c, 1117 Budapest, Hungary 2University of Bucharest, Faculty of Geology and Geophysics, Department of Geology, Laboratory of Paleontology, N. Balcescu Ave. 1, 010041 Bucharest, Romania

*[email protected]

Keywords: mammoth, enamel microstructure, biochronology, Late Pliocene - Pleistocene

Studying enamel microstructure proved to be useful for unravelling phylogenetic connections and to draw conclusions on the alimentary habits of fossil vertebrates. Evolutionary trends relating to this tissue were even used for establishing a relative chronology of sites with fossils that once belonged to the same lineages.

Present study used mammoth molars from Hungary and Romania. Detached enamel samples were cut and polished according to sagittal, horizontal and tangential planes. Then they were sectioned for polarising microscopy, or they were etched and prepared for a SEM study. Measurements were taken on micrographs with ImageJ along a line perpendicular to the enamel-dentine junction.

Enamel prisms have a diameter of 6-12 µm and a keyhole- or ginkgo-leaf-shaped cross section. Based on their arrangement, three layers were separable, which have unique extinction patterns under polarised light. The innermost layer (IL) contains irregularly oriented prisms. In the middle layer (ML), prisms are arranged into zones that rise towards the enamel-cementum junction. The orientation of the prisms is roughly the same within each zone but different in the adjacent ones. The decussation gradually weakens outwards until all prisms become parallel. The inner boundary of the outermost layer (OL) is marked by a sudden flattening in prism orientation.

Results are preliminary, but it seems that the IL of Mammuthus rumanus (MN16b) made up 15%, the ML made up 50-55% and the OL made up 30-35% of the total enamel thickness (ET). The ML of the early representatives of M. meridionalis (former MN17) became 5% thicker at the expense of the OL. Around 1.8 Ma, additional 5% thickening of the ML was compensated by the further thinning of the OL. The ML of M. trogontherii (from 0.8 Ma onwards) became 2-3% thicker, whereas the IL became proportionally thinner. The latter taxon was replaced by M. primigenius around 0.2 Ma with a middle layer that makes up 70-80% of the ET, which means 5- 10% thickening compensated by 5% thinning of the IL and OL separately. More samples are needed in the future in order to accurately assess the intraspecific variability.

This research was supported by the MTA Postdoctoral Research Programme.

58 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

‘Towards a Biogeographic Synthesis for the European Early Pleistocene based on the Mammal Record’ poster R3

PLIOCENE AND EARLY PLEISTOCENE CLIMATIC TRENDS IN ITALY INFERRED FROM COMPOSITIONS OF RHINOCEROSES AND GOMPHOTHERES P. Szabó1,2,3, L. Kocsis4, T. Vennemann3, L. Pandolfi5*, J. Kovács1,6, E. Martinetto7 and A. Demény8

1University of Pécs, Environmental Analytical and Geoanalytical Research Group, Szentágothai Research Centre, Ifjúság útja 20, 7624 Pécs, Hungary 2University of Pécs, Doctoral School of Chemistry, Ifjúság útja 6, 7624 Pécs, Hungary; 3Université de Lausanne, Faculty of Geosciences and Environment, Géopolis, Institute of Earth Surface Dynamics, 1015 Lausanne, Switzerland 4University Brunei Darussalam, Faculty of Science, Geology Group, Jalan Tungku Link Gadong, BE1410, Brunei Darussalam 5University of Roma Tre, Department of Sciences, Largo S.L. Murialdo 1, 00146 Roma, Italy 6University of Pécs, Department of Geology and Meteorology, Ifjúság útja 6, 7624 Pécs, Hungary 7University of Torino, Department of Earth Sciences, Via T. Valperga Caluso 35, 10125 Torino, Italy 8Institute of Geological and Geochemical Research Research Centre for Astronomy and Earth Sciences, Budaörsi útja 45, 1112 Budapest, Hungary

* [email protected]

Keywords: stable isotopes, enamel, climate, Pliocene, Early Pleistocene

Pliocene and Early Pleistocene (from ~5.2 to ~1 Ma) climate in Italy is here tentatively reconstructed using stable carbon and oxygen isotopic compositions of teeth enamel of fossil specimens from Rhinocerotidae and Gomphotheriidae taxa. Carbon isotope ratios were measured from the structural carbonates (δ13C), while oxygen isotope ratios from the structural carbonate (δ18OCO3) and also from the phosphate (δ18OPO4) of the bioapatite. 51 specimens from 21 Italian localities were considered in this work. The fossils belong to five different taxa, Stephanorhinus etruscus (n=21), "Dihoplus" megarhinus (n=9), Stephanorhinus jeanvireti (n=6), Stephanorhinus sp. (n=8) and Anancus arvernensis (n=7). Teeth of adult individuals and probably belonged to different individuals were sampled. All the δ13CCO3 results indicate that the investigated taxa were grazers-browsers in pure C3 vegetation. In central Italy the Early Pliocene was characterized by higher than present-day precipitation (mean annual precipitation – MAP- more than 2000 mm) and temperatures (mean annual temperature – MAT ~18 °C). Both parameters show fluctuations in the Late Pliocene and in the Early Pleistocene with a long term decreasing trend. By the end of the Early Pleistocene the values drop to a present day levels (13 °C and 600 mm). In North Italy the δ13C values show a sharp decrease between MN16a and MN16b mammal biozones that is most likely linked to the Northern Hemisphere Glaciation at 2.7 Ma after the "Mid-Pliocene" Warm Period. Recent analyses on plant macrofossils from same area where MN16b mammal assemblages have been reported pointed out the disappearance of several thermophilous plants already within the latest Pliocene. In the Pleistocene precipitation was lower than in the Pliocene by about 1000 mm. Altogether with some exceptions the new isotope data agree well with paleoclimatic reconstructions based on palynological and other proxies.

59 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

‘Towards a Biogeographic Synthesis for the European Early Pleistocene based on the Mammal Record’ poster R4

PALAEOECOLOGICAL REVIEW OF THE PLEISTOCENE CERVIDS FROM HUNGARY BASED ON DENTAL MICROWEAR ANALYSIS B. Szabó1 and A. Virág2*

1Szent István University, Faculty of Veterinary Science, István street 2, 1078 Budapest, Hungary 2MTA-MTM-ELTE Research Group for Paleontology, Pázmány Péter sétány 1/c, 1117 Budapest, Hungary

*[email protected]

Keywords: microwear, Cervidae, Pleistocene, Hungary

Dental microwear analysis has become an extensively utilized method for gathering information on past environments of localities through the understanding of the diversity of alimentary habits within fossil communities. In order to make reliable comparisons of different geographic areas and time intervals, a widespread group is needed that had an equally wide spatial and temporal range. Cervids are one of the most common taxa to be found among the several mammalian remains that occurred in Hungary during the Pleistocene, making them an excellent group for such palaeoecological investigations. In addition, they are not specialized for the consumption of just a small variety of plants, but they are generalists, thus they can inhabit many environments covered by different types of vegetation.

The aim of the present study was to reconstruct the different environments existed in Hungary during the latest Pliocene and Pleistocene using the microwear pattern visible on cervid molars. The teeth of Alces, Capreolus, Cervus, Megaloceros and Rangifer were used from various sites with an age ranging from 2.7 Ma to 11 ka.

Cleaned enamel areas were moulded with Colténe Affinis Precious Regular Body polyvinyl siloxane impression material and then high-resolution transparent casts were made with EPO-TEK 301. The casts were analysed with light microscopy at 35× magnification. Microwear scars were judged as scratches or pits based on their refractive index and length to width ratio (the boundary was 4:1). Five locations were selected on every sample, if possible. Based on the average number of scratches and pits, the specimens were classified into one of the following categories: browser, grazer, or mixed feeder.

According to our results, cervids from glacial interval sites show a tooth wear pattern with significantly more scratches than pits, suggesting a grazing diet that fits well with the steppe vegetation of these cold and dry time spans. Whereas remains from interglacial sites with assumingly forest or forest steppe vegetation show a wear pattern with more pits that is characteristic for browsers or mixed feeders.

This research was supported by the MTA Postdoctoral Research Programme.

60 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

‘Towards a Biogeographic Synthesis for the European Early Pleistocene based on the Mammal Record’ poster R5

EUROPEAN SMALL MAMMAL FAUNAS DURING THE SECOND PART OF THE EARLY PLEISTOCENE - THE BEGINNING OF THE MIDDLE PLEISTOCENE A.K. Markova1*

1Institute of Geography of RAS

*[email protected]

Keywords: Mammals, Quaternary, Europe

The European small mammal faunas correlated with the end of Early and the first half of the Middle Pleistocene were analyzed (1,07 –0,70 ma).

During the first stage, small mammal faunas correlated with the Jaramillo palaeomagnetic (1,07-0,99 Ma) event include remains of Mimomys savini, М. pusillus, Clethrionomys sokolovi, advanced Allophaiomys, Borsodia fejervaryi, Prolagurus pannonicus, Lagurodon arankae and Eolagurus argyropuloi. These faunas were named as Kairian (=Ostrogozhskian) (Eastern Europe) and as “Colle Curti” phase in Italy.

The second phase presents the faunas with the first representatives of subgenera Stenocranius (S. hintoni) and Microtus (Terricola) sp. These faunas include also Mimomys savini, M. pusillus and Allophaiomys pliocaenicus nutiensis and were described in Eastern Europe as Morozovkian faunas. Their localities are in the Matuama.

The next stage is distinguished by the first appearance of Microtus ex gr. oeconomus (=M. ratticepoides). The core of these faunas includes steppe and yellow lemmings Prolagurus pannonicus and Eolagurus argyropuloi, and Microtus (Stenocranius) hintoni. Furthermore, a few Mimomys savini and Allophaiomys pliocaenicus nutiensis remains were found in the faunas. These faunas correlated with the end of Matuama and were described as Petropavlovkian ones. They are close to Trichera Dolina (TD5-TD6 layers). The first appearance of Microtus arvalinus and Prolagurus posterius indicates the new evolutionary stage of small mammals. These are correlated with the very end of Matuama and close to Early Tiraspolian ones in Eastern Europe and to many Western European faunas.

The beginning of the Brunhes normal magnetic Epoch was characterized by both presence of rooted voles of Mimomys genus and more significant quantity of Microtus species. Analysis of Early to Middle Pleistocene small mammal remains, particularly the unique Arvicolinae material, helps to elucidate the evolution in different phylogenetic lineages.

61 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Thursday plenary poster V3

A NEW ADVANCED SAUROPODIFORM DINOSAUR FROM THE LATEST NORIAN/ OF SWITZERLAND F.M. Holwerda1, O.W.M. Rauhut1,2 and H.F. Furrer3

1SNSB, Bayerische Staatssammlung für Paläontologie und Geologie and Department of Earth and Environmental Sciences, Ludwig-Maximilians-University, Richard-Wagner-Str. 10. 80333 München, Germany 2GeoBioCenterLMU, Ludwig-Maximilians-University, München 3Paläontologisches Institut und Museum, Universität Zürich, Karl Schmid-Strasse 4, 8006 Zürich, Switzerland

*[email protected]

Keywords: Triassic, Norian/Rhaetian, Sauropod(omorphs), Switzerland

Sauropods, the emblematic, large, long-necked dinosaurs that are well-represented in the vertebrate faunas throughout the era, most probably originated from small bipedal/quadrupedal basal sauropodomorphs. However, their early evolution and origin is not well understood, and the transition from more basal forms to sauropods remains poorly studied. Depending on the definition of Sauropoda adopted, phylogenetic studies indicate that this transition must have taken place betweenthe Late Triassic (Norian/Rhaetian) and the late Early Jurassic. Well- preserved material of derived non-sauropodan sauropodomorphs from this time is known from Argentina and South-Africa. Although the Triassic of Europe has yielded a wealth of sauropodomorph material as well, with exception of the enigmatic Camelotia, all Triassic taxa recorded so far are quite basal, non-sauropodiform taxa, representing plateosaurids or more basal forms. Apart from named taxa, such as Plateosaurus, Camelotia and Gresslyosaurus, there is material that has not yet received much attention. The museum of Zurich University holds such material, found at Schleitheim in the Kanton of Schaffhausen, Switzerland, which is tentatively dated to be latest Norian to Rhaetian in age. The material had been previously assigned to Plateosaurus, but was found to differ from this taxon in a recent re- examination. The material several presacral and caudal vertebrae, a partial humerus, ilium, partial femur, and other fragmentary appendicular elements, which are most probably derived from one individual. The material differs from all other known basal sauropodomorphs in the development of the postacetabular blade and the brevis shelf of the ilium and the crista tibiofibularis of the femur, and thus represents a new taxon. An extensive phylogenetic analysis combining two recent matrices with additional new characters, retrieves this specimen as a derived sauropodiform or even a basal sauropod, depending on which definition of sauropoda is used. It is more derived than Antetonitrus, but more basal than Isanosaurus, Pulanesaura and Vulcanodon, placing it within the phylogenetic range of the sauropodomorph- sauropod transition. This not only confirms the presence of more derived sauropodomorphs in the Late Triassic of Europe, but also proves a greater diversity of sauropodomorphs in the Late Triassic of Europe than previously assumed.

62 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Thursday plenary poster V4

CONSERVATION STATUS OF THE PALAEONTOLOGICAL COLLECTION OF LO HUECO F. Marcos-Fernández1* and F. Ortega2

1Universidad Complutense de Madrid, 2 Universidad Nacional de Educación a Distancia

*[email protected]

Keywords: Palaeontology, Conservation, Vertebrates, Preparation

The vertebrate fossils from the "Lo Hueco" fossil site (Late Cretaceous. Fuentes, Cuenca, central Spain) have a generally good preservation status. However, all of them are extremely fragile, having shown a high risk of destabilisation due to the effects of both intrinsic and extrinsic factors as composition materials and the processes that occur during extraction, storage and exhibition without control.

An analysis of the state of conservation of the paleontological collection was carried out with the aim of identifying deterioration agents that need control to stop risks and to ensure the continued existence of fossils in time.

1. Fragility related to materials filling the porosity of the fossils. - Clay: it is the predominant lithology of the surrounded matrix and fills the fossil cavities. The particular composition of these clays is unstable for external factors such as humidity. - Gypsum: deposited as needles, lentils or superficial deposits. These deposits can break the fossil structure and are also unstable to humidity and UV. - Iron oxides: they produce a red stain or, in more severe situations, a red crust, sometime so deep as some millimetres. It is unstable because of humidity that causes a variance on the dilation coefficient. It can produce micro-fractures in between the body and the bone surface, destroying surficial layers. 2. Fragility related to the processes of the extraction and not controlled storage conditions: - Removing the pressure produced by the sediment. This decompression produced destabilisation on the materials and the relation in between each other. - Dry up process produces dehydration and this can change the status of certain materials. The change of volume produces cracks that, in the most serious situations, can transform to dust important parts of the fossil.

These have been recognized as the factors to be controlled in order to avoid: - Lack of cohesion that can produce disintegration, collapse and even pulverisation of the fossils, leaving its external aspect without physical consistency. - Rise of fissures and fractures that can lead into breaks and disjunctions directly related with the compression and dilation coefficients of the materials.

63 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Thursday plenary poster V5

A NEW CHONDRICHTHYAN FAUNA FROM THE LATE JURASSIC (KIMMERIDGIAN) OF THE SWISS JURA WITH A SHARK PICTURE DOMINATED BY HYBODONTS 1, 2 3 4 2 L. Leuzinger *, G. Cuny , E. Popov and J.-P. Billon-Bruyat

1Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja (CRILAR), Entre Ríos y Mendoza s/n, 5301 Anillaco, Argentina, 2Section d’archéologie et paléontologie, Office de la culture, République et Canton du Jura, Hôtel des Halles, 2900 Porrentruy, Switzerland, 3LGLTPE, UMR CNRS ENS 5276, Claude Bernard University Lyon 1, 2, rue Raphaël Dubois, F-69622 Villeurbanne Cedex, France, 4Department of Historical Geology and Paleontology, Saratov State University, 83 Astrakhanskaya Str., 410012 Saratov, Russia

*[email protected]

Keywords: fish, hybodont, neoselachian, chimaeroid, lagoon

Marine chondrichthyan faunas from the Late Jurassic of Europe are often characterized by a high abundance of so-called modern sharks, skates and rays (Neoselachii) and comparatively scarce hybodont sharks (Hybodontiformes). This reflects the evolutionary history of sharks during the Jurassic, with a strong radiation of neoselachians in marine realms and a subsequent decline of so-far dominating hybodonts, gradually confined to brackish or freshwater environments until their extinction by the end of the Cretaceous. Here, we present a new Kimmeridgian chondrichthyan fauna from Porrentruy (canton of Jura, NW Switzerland), mainly based on large bulk samples of isolated teeth, resulting from controlled excavations during the construction of the Transjurane highway A16. Contrary to the general European trend in marine settings, modern sharks (Selachimorpha) are rather scarce and predominantly represented by benthic taxa, such as heterodonts and angelsharks, while the abundance of hybodonts is three times greater. Hybodonts are the dominating sharks of our assemblage and also dominate the global chondrichthyan faunal spectrum together with guitarfishes (Batomorphii, Rhinobatidae) and chimaeroids (Holocephali, ‘Edaphodontidae’). The associated fauna indicates that marine conditions prevailed in the lagoonal environment of Porrentruy, although a previous isotopic study on the hybodont shark Asteracanthus revealed a reduced salinity signal. The Porrentruy carbonate platform seemingly offered favorable conditions for hybodonts to thrive during the Kimmeridgian, while they were already scarce or even absent in contemporaneous, neighbouring localities with marine, lagoonal settings (e.g. Solothurn, Cerin, Solnhofen, Nusplingen). The shallower marine water conditions of our study site and a nearby freshwater influence may have rather favored hybodonts and other euryhaline taxa, such as guitarfishes and some ‘edaphodontid’ chimaeroids (Ischyodus).

64 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Thursday plenary poster V6

FISHES ( AND OSTEICHTHYES) FROM THE MAASTRICHTIAN (LATE CRETACEOUS) OF NORTH- EASTERN IBERIA 1 2 A. Blanco * and M. Szabó

1Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, C/ Escola Industrial 23, 08201, Sabadell, Catalonia, Spain. 2MTA-ELTE Lendület Dinosaur Research Group, Pázmány Péter sétány 1/C, Budapest 1117, Hungary

*[email protected]

Keywords: Chondrichthyes, Neopterygians, Teleosteans, Tremp Formation, Maastrichtian

Chondrichthyan and osteichthyan fishes are commonly recovered from Upper Cretaceous outcrops in European localities. Complete or partially associated fishes are extremely rare, and often their remains are recovered isolated through screen- washing techniques together with other small vertebrates. During the last years, intensive sampling for vertebrate microfossils in the Maastrichtian transitional to continental beds of the Tremp Formation (southern Pyrenees, Catalonia, Spain) has yielded abundant fish remains. These beds are exposed in Serrat del Rostiar-1, Serrat del Pelleu, Camí del Soldat, Els Nerets L’Espinau and Fontllonga-6 sites, and represent different environments including brackish mudflats, coastal wetlands and fluvial settings with some marine influence (Soler-Gijón & López-Martínez, 1998; Riera et al., 2009; Blanco & Bolet, 2014; Díez-Canseco et al., 2014; Fondevilla et al., 2015; Blanco et al., 2016).

The sample of Serrat del Rostiar-1 site includes diagnostic remains of Lepisosteus sp. and Atractosteus sp., isolated teeth of Pycnodontiformes and one basal teleostean taxon. Camí del Soldat site has yielded isolated teeth and scales of Atractosteus sp., Belonostomus sp., Pycnodontiformes, Phyllodontidae and one indeterminate teleostean. The Fontllonga-6 sample includes Igdabatis indicus, Rhombodus ibericus, remains of indeterminate pycnodontiform fishes and Atractosteus sp. Els Nerets and Serrat del Pelleu sites are the less diverse assemblages, including a basal teleostean and an indeterminate lepisosteid, respectively.

In contrast, L’Espinau site has yielded the most diverse ichthyofaunal assemblage, including one indeterminate selachimorph, Atractosteus sp., a probable semionotiform, pycnodontiforms, an indeterminate amiid, possible Caturus sp. and Belonostomus sp. remains, two or three different phyllodontids, two basal teleostean taxa and one indeterminate acanthomorph.

The preliminary results suggest a great diverse ichthyofauna in the Ibero-Armorican Domain from the Early Maastrichtian to the end of the Late Maastrichtian. Rays are only present in the brackish outcrops, whereas the fluvial settings have yielded lepisosteid, pycnodontiform and teleostean remains. L’Espinau site represents a coastal environment inhabited by mixed marine and freshwater taxa. More detailed taxonomic affinities, as well as their palaeoecological and palaeobiogeographical implications will be discussed in a work in progress.

65 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Thursday plenary poster V7

WRITTEN IN BONE: THE ONLY KNOWN QUILL PITS OF EARLY PENGUINS 1 P. Jadwiszczak *

1University of Bialystok, Institute of Biology, Ciolkowskiego 1J, 15-245 Bialystok, Poland

*[email protected]

Keywords: Sphenisciformes, Eocene, Antarctic Peninsula, wing, feathering markings

Avian flight feathers can leave traceable marks on bones, which show the positions of their attachment points. In the case of secondary remiges, such scars are termed ulnar quill knobs or pits. The latter are present, among others, in specimens assignable to small- and medium-sized modern penguins (Sphenisciformes). Recently, they have been found in an early (i.e., Paleogene) penguin as well. Reported here is the only known fossil penguin ulna characterized by the presence of two rows of regularly spaced concave marks that resemble quill pits in its present- day counterparts. Importantly, the closest living relatives to penguins, tubenoses (Procellariiformes), have knobs/papillae. The above specimen comes from the Eocene La Meseta Formation on Seymour Island (West ) and belonged in the wing skeleton of a relatively small-sized stem sphenisciform. Undoubtedly, this finding contributes significantly, although indirectly, to our knowledge on plumage evolution in penguins. In this respect, it constitutes another milestone, following the trailblazing article, published by an international team of researchers in 2010, on a giant penguin with preserved feathers (no quill pits reported) recovered from the Eocene of Peru.

66 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Thursday plenary poster V8

MACROEVOLUTION AND THE ADAPTIVE RADIATION OF

MESOZOIC SQUAMATES J.A. Herrera Flores1* and M.J. Benton1

1University of Bristol, School of Earth Sciences, Wills Memorial Building, Queens Road BS8 1RJ, United Kingdom

*[email protected]

Keywords: geometric morphometrics, rates of evolution, Squamata

The Squamata is the order of that includes lizards, snakes and amphisbaenians. Extant squamates are represented by over 9000 living species distributed worldwide, and they are one of the most successful groups of vertebrates. The early origin of squamates is not clear, because of a gap in its fossil record between the Late Triassic and the Middle Jurassic. The oldest known squamates come from the Late Jurassic, but their remains are rare. In contrast, the fossil record of Cretaceous squamates is quite rich, which suggests that this group had a big radiation at the end of the Mesozoic. In order to study the macroevolution of Mesozoic squamates, we compiled a database of 2D images of lower jaws to perform a geometric morphometrics analysis in R with the package Geomorph. We found different changes in morphospace through the Mesozoic, for example Late Jurassic squamates formed a small cluster, Early Cretaceous taxa increased in morphospace occupancy, but also moved into a different space, while Late Cretaceous squamates showed a massive expansion. When we divided Late Cretaceous squamates by form, lizards occupied the widest morphospace, while amphisbaenians occupied an apparently different morphospace from other forms, but very close to lizards. Mosasauroids formed a tight cluster with high diversity, sharing some morphospace with marine snakes and varanoid lizards. Snakes formed a relatively tight cluster that interestingly overlaps with lizards and mosasauroids near the centroid of the plot. When we divided Late Cretaceous taxa by feeding modes, morphospaces suggest that, although there were several feeding modes, Late Cretaceous taxa were clearly separated into two main feeding modes, insectivorous and carnivorous, which could be a factor that influenced their adaptive radiation at the end of the Mesozoic.

67 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Thursday plenary poster V9

USING BAYESIAN TO INVESTIGATE THE EFFECTS OF THE CRETACEOUS-PALAEOGENE MASS EXTINCTION ON SQUAMATE DIVERSITY 1 1 2 2 2 1 C.G. Klein *, N.R. Longrich , D. Pisani , A. Tanner , J. Vinther and M.A. Wills

1University of Bath, Department of Biology & Biochemistry, Bath BA2 7AX, United Kingdom 2University of Bristol, School of Earth Sciences, Life Sciences Building, Bristol BS8 1TQ, United Kingdom

*[email protected]

Keywords: Squamata, Bayesian inference, phylogenetics, Cretaceous-Palaeogene

The clade Squamata arose about 200 Ma, and has undergone numerous diversification events. Today, it is the second most diverse terrestrial vertebrate order, with over 9400 known extant species. The study of this group has provided partial genetic sequences for more than 4100 member species, and so it has become a model clade for the testing of phylogenetic methods. However, comparatively few attempts have been made to time-calibrate the consequent phylogenetic trees, and the results rarely converge on the same divergence dates.

Here we look at the effects of the Cretaceous-Palaeogene (K-Pg) mass extinction on squamate diversity, as well as the effects of outgroup taxon sampling on estimated divergence dates of deeper nodes within Squamata. A dataset of 110 squamate taxa, coded for twelve nuclear and mitochondrial genes, is used to construct a molecular based timescale for the squamates. The sister clade, Rhynchocephalia, consists of a single living species, Sphenodon punctatus. This has implications for the choice of outgroup taxa, as less closely related taxa have long branch lengths, but simultaneously can stabilise results. Further outgroup sampling may be used to counteract problematic branch lengths, creating a number of possible sampling regimes.

Analyses using a single outgroup taxon reveal node ages up to 30 Ma older or younger than expected values based on previous analyses and the fossil record. Increasing the number of outgroup taxa to ten, by sampling from Mammalia, Crocodilia and Aves, pushes divergence estimates of deeper and shallower nodes closer to the expected divergence dates. As well as snakes, several other clades appear as potential candidates of post-extinction radiations, and therefore are in need of closer study to properly characterise the post-extinction recovery of Squamata.

68 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Thursday plenary poster V10

GALLOTIA STEHLINI (LACERTIDAE) FROM THE PLEISTOCENE OF GRAN CANARIA: NEW DATA FROM MICRO CT-SCANS OF UNPUBLISHED SPECIMENS 1 1,2 A. Bolet * and J. Fortuny

1Institut Català de Paleontologia Miquel Crusafont, ICTA-ICP building, c/ de les columnes, s/n, E-08193 Cerdanyola del Vallès, Spain 2C2RP, CNRS-MNHN-UPMC, 8 rue Buffon, CP38, F-75005 Paris, France

*[email protected]

Keywords: Squamata, lizard, Canary Islands, Gallotinae, Quaternary

Gallotia stehlini is the only species of lacertid living in the canarian island of Gran Canaria (with exception of the recently introduced Gallotia atlantica). The identity of subfossil populations in all islands is still debated because many extinct species have been subsequently synonymised or their rank has changed. The presence of small specimens among material from La Isleta locality (Pleistocene) posed a problem regarding whether they corresponded to a juvenile of G. stehlini or to an adult of a small species now extinct in the island. Accordingly, a proper identification of subfossil material from Gran Canaria was important in its potential to highlight differences between the subfossil and extant populations that could reveal the presence of a different species or subspecies, or in assessing the presence of G. stehlini as a subfossil.

We performed a deep study of subfossil remains of Gallotia from Gran Canaria for the first time in order to investigate the identity of the subfossil populations. The remains studied were collected in the late XIXth century from the classical locality of la Isleta and are housed at Museu de Geologia de Barcelona, but had not been previously reported or investigated. Other specimens from the same locality have been cited and shallowly described, but their identification as G. stehlini had a weak morphological basis. These previous vague descriptions of otherwise well-preserved specimens were a consequence of the presence of a crust covering the articulated bones, what precluded a proper observation of morphological details. We performed a Micro CT-Scan of one of the small previously unpublished specimens in order to provide an accurate account of the osteological characters and compare them with the states in different species of Gallotia. Our results indicate that this specimen unequivocally represents a juvenile of G. stehlini, and no evidence for the presence of a different species or subspecies has been identified. The exceptional preservation of the specimens from La Isleta suggest a high potential for investigating a subfossil population in Gran Canaria (including ontogenetic changes), but may necessarily rely in the acquisition of Micro CT-Scans of the remaining specimens.

69 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Thursday plenary poster V11

A NEW GENUS FOR THE EUROPEAN MIDDLE EOCENE TURTLE ‘AFF. ERYMNOCHELYS’ EREMBERTI, AND STUDY OF TWO NEW CLOSELY RELATED FORMS 1 2 3 A. Pérez-García *, F. de Lapparent de Broin and X. Murelaga

1Grupo de Biología Evolutiva, Facultad de Ciencias, UNED, Paseo de la Senda del Rey 9, 28040 Madrid, Spain. 2Sorbonne Universités – CR2P – MNHN, CNRS, UPMC-Paris 6, Muséum national d’Histoire naturelle, 57 rue Cuvier, CP 38, 75231 Paris cedex 5, France. 3Departamento de Estratigrafía y Paleontología, Facultad de Ciencia y Tecnología, Universidad del País Vasco (UPV/EHU), Apartado 644, 48080 Bilbao, Spain.

*[email protected]

Keywords: Eocene, Europe, , , new taxa

The European ‘aff. Erymnochelys’ eremberti is the only member of the Erymnochelys group of turtles (Pleurodira, Podocnemididae) so far defined in the Laurasiatic record. It is known by a single specimen, from the Lutetian (middle Eocene) of the French locality of Saint-Germain-en-Laye (Yvelines, Île-de-France). This specimen corresponds to a partial skeleton, including the skull and lower jaw, the fragmentary but partially reconstructed dorsal carapace, the relatively complete plastron, and various partial remains of the axial and appendicular skeleton. Its description and detailed study are completed here, a diagnosis for this species being proposed for the first time. It is attributed to a new genus, corresponding to both the first genus of the Erymnochelys group defined outside of Africa and the oldest unambiguous generic determination of a member of this group heretofore attested. It corresponds to a coastal form, all the African representatives being continental taxa. This middle Eocene species is not the only member of the Erymnochelys group identified in the European record, this group also being represented in the Early and Late Eocene. Thus, a Spanish specimen from the late Eocene (Priabonian) of Can Beuloví (Sobremunt, Osona, Catalonia), as well as an almost complete shell from the early Eocene (Ypresian) of the French locality of Soulane (Jonquières, Aude, Languedoc- Roussillon-Midi-Pyrénées), are studied. This French specimen is identified as attributable to a new species, being the oldest defined representative of the Erymnochelys group. The Spanish specimen also corresponds to a new species.

70 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Thursday plenary poster V12

THE OLDEST EUROPEAN TESTUDINIDS: DESCRIPTION OF NEW TAXA AND REVISION OF POORLY-KNOWN FORMS 1 1 2 A. Pérez-García *, F. Ortega and E. Jiménez Fuentes

1Grupo de Biología Evolutiva, Facultad de Ciencias, UNED, Paseo de la Senda del Rey 9, 28040 Madrid, Spain. 2Sala de las Tortugas de la Universidad de Salamanca, Facultad de Ciencias, Plaza de los Caídos s/n, Salamanca, Spain.

*[email protected]

Keywords: Eocene, Europe, , Testudinidae, new taxa

Testudinidae is the only extant clade of (i.e. terrestrial turtles). It is recognized as a very successful group, based on its abundance, diversity, and geographic distribution. It is relatively abundant in the fossil record. In this sense, the knowledge about its Neogene evolutionary history is relatively good. However, its Paleogene record is much more limited.

Testudinids are known in Europe from the early Eocene. Scarce information on the taxonomy and systematics of the European Eocene testudinids was so far available. The oldest testudinids from Europe, recovered in early and middle Eocene levels of several countries, are studied here. Some of them were previously known, being generally attributed to North American genera (i.e. and Achilemys). The hypothesis suggesting that the European Eocene genera are exclusive of this continent is confirmed. Thus, two new genera are proposed. The largest collection of European Paleogene testudinids, from the middle Eocene of Mazaterón (Soria, Spain), has been prepared for this study. A new species from Mazaterón is proposed. The validity of some European Eocene taxa is refuted, the French middle Eocene ‘ castrensis’, the Austrian middle Eocene ‘Cheirogaster steinbacheri’, and the Spanish middle-late Eocene ‘Asturichelys multicostatus’ being proposed as nomina dubia. The diagnosis of several forms is emended. In addition, some of the taxa analyzed here are included for the first time in a phylogenetic hypothesis including, among others, the European medium to large testudinids.

71 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Thursday plenary poster V13

ON THE PRESENCE OF (TESTUDINES, PLEURODIRA) IN THE EUROPEAN CENOZOIC RECORD 1 A. Pérez-García *

1Grupo de Biología Evolutiva, Facultad de Ciencias, UNED, Paseo de la Senda del Rey 9, 28040 Madrid, Spain.

*[email protected]

Keywords: Cenozoic, Europe, Pleurodira, Bothremydidae, new taxon

Pleurodira is a lineage of turtles originated in Gondwana. Several migrations of this group to Europe are known from the Upper Cretaceous, and especially, during the uppermost Cretaceous and the Eocene. In this regard, several representatives of Podocnemididae (Pelomedusoides) are known in the Eocene record of this continent, including the abundant and diverse European taxon Neochelys, as well as various forms of the Erymnochelys group, currently under study (e.g. ‘aff. Erymnochelys’ eremberti). A second Gondwanan lineage of Pelomedusoides also migrated from Africa to Europe: Bothremydidae. This group was very abundant and diverse on the uppermost Cretaceous record of this continent. However, it has not so far been unambiguously identified in the European Cenozoic record.

A new representative of Pleurodira is presented here. It comes from the early Eocene of Southwestern Europe. The new taxon is represented by two . It is attributable to Pelomedusoides. Several characters shared with the members of Bothremydidae are recognized in this taxon. Its detailed study, together with the revision of poorly- known specimens, provides new data to assess whether Bothremydidae was part of the turtle fauna of the European Eocene record.

72 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Thursday plenary poster V14

THE OLDEST MEMBER OF THE CROWN PLEURODIRA IN 1 A. Pérez-García *

1Grupo de Biología Evolutiva, Facultad de Ciencias, UNED, Paseo de la Senda del Rey 9, 28040 Madrid, Spain

*[email protected]

Keywords: Cretaceous, Europe, Pleurodira, Bothremydidae, new taxon

Pleurodira is one of the two major lineages of turtles known from the Early Cretaceous to the present. The first representatives of this clade are Gondwanic forms. In fact, extinct and extant pleurodires were restricted to relatively warm regions, their temperature requirements being much more extreme than those of the Cryptodira. Although Pleurodira is now restricted to tropical and intertropical regions, changes in the global temperatures and in the relative position of the landmasses allow the dispersal of some representatives to Laurasia. Thus, these diachronic migrations were known from the Santonian, being especially relevant between the uppermost Cretaceous and the Eocene. In this sense, both Bothremydidae and Podocnemididae were relatively abundant and diverse in the European record (Bothremydidae in the Late Cretaceous and Podocnemididae in the Paleogene).

A new representative of Pleurodira, corresponding to a member of Bothremydidae, is presented here. The new taxon comes from pre-Santonian levels of Southwestern Europe. It is represented by abundant remains, highlighting a skull, relatively complete and partial shells, abundant plates, and appendicular elements. This discovery shows that the first migration of Pleurodira from Gondwana to Laurasia occurred earlier than hitherto proposed.

73 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Thursday plenary poster V15

COMPARATIVE OBSERVATIONS ON THE AND EVOLUTION OF THE FORAMINA ASSOCIATED WITH THE TRIGEMINAL NERVE SYSTEM IN TURTLES 1 1 S.W. Evers * and R.B.J. Benson

1University of Oxford, Department of Earth Sciences, South Parks Road, OX1 3AN Oxford, United Kingdom

*[email protected]

Keywords: Turtles, homology, neuroanatomy, trigeminal nerve

The trigeminal (cranial V) nerve of reptiles exits the endocranial cavity through an opening located anteriorly on the lateral wall of the braincase — the trigeminal foramen. This foramen is on the laterosphenoid-prootic contact in most archosaurs and lepidosaurs, and in taxa with an extracranial trigeminal ganglion, the foramen is modified to a broad fossa containing separate foramina for the trigeminal rami. Extant turtles, and almost all fossil forms, lack an ossified laterosphenoid. Instead, the parietal extends far ventrally, forming the lateral wall of the braincase anteriorly, and contacting the prootic and bony palate. The trigeminal foramen of turtles is therefore located at the parietal-pterygoid contact. These two bones minimally form the dorsal and ventral margins of the foramen, with additional bones such as the prootic posteriorly and the epipterygoid anteroventrally contacting the foramen in many taxa. Topologically, this is at odds with the situation in other reptiles. These differences raise questions about the morphological transformations involved in the origins of the turtle trigeminal foramen, and warrant an assessment of the homology of the foramen, which might inform the phylogenetic position of turtles among . New observations on fossil turtles, including stem-turtles such as Proganochelys or more derived fossil cryptodires such as etalloni reveal a possible hypothesis for the evolution of the trigeminal nerve foramina in turtles. The trigeminal foramen in turtles transmits the maxillomandibular ramus of the CN V, but not the ophthalmic ramus. In Proganochelys, a medial foramen involving the laterosphenoid, and a laterally placed, dorsally open aperture indicates that an internal trigeminal foramen as present in most other reptiles was secondarily reduced in turtles, and that the trigeminal foramen of turtles is possibly homologous to the maxillomandibular foramen of reptiles with extracranial trigeminal ganglia, such as . These observations form the basis for a revision of phylogenetic characters concerning the trigeminal foramen in analyses including turtles and other saurians.

74 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday plenary poster session V16

UPPER BLESA FORMATION (TERUEL, SPAIN), NEW INSIGHTS ON THE EARLY CRETACEOUS MARINE REPTILE FAUNA FROM THE IBERIAN PENINSULA J. Parrilla-Bel1* and J.I. Canudo1,2

1University of Zaragoza, Grupo Aragosaurus-IUCA, Department of Earth Science, Pedro Cerbuna 12, 50009, Zaragoza, Spain. 2Museo de Ciencias Naturales de la Universidad de Zaragoza, Paraninfo, Plaza de Basilio Paraíso, 4, 50005, Zaragoza, Spain

*[email protected]

Keywords: marine Crocodylomorpha, Barremian, Plesiosauria

The fossil record of Early Cretaceous marine reptiles is scarce in the Iberian Peninsula. Isolated and fragmentary remains of vertebrae and teeth mainly represent this record from Upper Barremian of Teruel and Castellón (Spain). In the last years, a new vertebrate fossils collection has been recovered from the Upper Blesa Formation (lower Barremian), in the Iberian Range in Teruel (Spain).

The upper part of the Blesa Formation was deposited in a coastal system. Lithologically are mainly carbonated with more detrital sediments to the top of the formation. The vertebrate remains come from three limestone levels. The young level it is where the fossil remains are larger. We found some plesiosaur remains (vertebrae and teeth), chelonian plates and cranial and postcranial elements of crocodylomorphs: two maxillary fragments, an ilium and a skull fragment. The skull fragment corresponds with the cranial postorbital region. It has large and elongated supratemporal fenestrae, limited by flat and narrow bone surfaces. It has also preserved big otic apertures. In a previous work, the fossil was tentatively assigned to Metriorhynchidae indet. However, a new detailed study suggests a possible relationship with marine tethysuchians, being the first tethysuchian remain described in the Iberian Peninsula and the oldest specimen known to date at the world.

The presence of plesiosaurs and marine crocodylomorphs in the Blesa Formation establishes this formation as an interesting site to increase the knowledge of these groups in the Barremian where they are scarce worldwide.

75 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday plenary poster session V17

OVERVIEW OF MARINE REPTILES FROM THE TURONIAN OF THE OPOLE AREA, SOUTHWEST POLAND S. Sachs1,2*, R. Niedźwiedzki3, M. Kędzierski4, B.P. Kear5, E. Jagt-Yazykova6 and J.W.M. Jagt7

1Naturkundemuseum Bielefeld, Abteilung Geowissenschaften, Adenauerplatz 2, 33602 Bielefeld, Germany 2 Im Hof 9, 51766 Engelskirchen, Germany 3Wrocław University, Institute of Geological Sciences, pl. M. Borna 9, 50-204 Wrocław, Poland 4 Jagiellonian University, Institute of Geological Sciences, Oleandry 2a, 30-063 Kraków, Poland 5Uppsala University, Museum of Evolution, Norbyvägen 18, 752 36 Uppsala, Sweden 6University of Opole, Laboratory of Palaeobiology, Oleska 22, 45-052 Opole, Poland 7Natuurhistorisch Museum Maastricht, de Bosquetplein 6-7, 6211 KJ Maastricht, the Netherlands *[email protected]

Keywords: Plesiosauria, Mosasauroidea, Upper Cretaceous, central Europe

Turonian strata in the Opole area are well known for their well-preserved invertebrate fossils. Amongst the earliest descriptions is Leonhard’s 1897 monograph “Die Fauna der Kreideformation in Oberschlesien”, in which some isolated tetrapod elements, including mosasauroid and plesiosaurian teeth as well as a fragmentary bone named Plesiosauridarum were listed. Part of Leonhard's material is now curated in the collection of the Department of Paleozoology of University of Wrocław. Further, yet undescribed, specimens were found in the collection of the Ruhrmuseum in Essen. At present several plesiosaurian teeth, one mosasauroid tooth and a paddle element (? a plesiosaurian mesopodial) are available. Leonhard assigned the first-named to Polyptychodon interruptus. These are slender, yet conical with strong apicobasal striations, which justify reference to pliosauromorph plesiosaurians. They differ from typically more massive teeth of Late Cretaceous pliosaurids (Brachauchenius/Megacephalosaurus) and those found in coeval strata of the Bohemian Cretaceous Basin (BCB, Czech Republic) and more closely resemble teeth of polycotylids. Interestingly, a similar tooth has recently been described from Turonian strata of the Saxonian Cretaceous Basin (SCB, Germany). So far, mosasauroids are unknown from the SCB; however, a tethysaurine mosasauroid was described from the BCB. A recurved mosasauroid tooth crown, assigned to Liodon anceps by Leonhard (along with a second one tooth that could not be reloacted) shows well-developed carinae, strong lingual folds and finer ornament adapically and a near-smooth labial side with faint facetting. A comparison with coeval, or slightly younger, material from England is needed to assign this material to a genus, or family. Leonhard’s mysterious Plesiosauridarum could not be relocated, yet on the basis of his description and illustration, it can be identified as a damaged mosasauroid vertebra. In conclusion, a diverse marine reptile fauna comprising pliosauromorph plesiosaurs with more slender teeth than those found in coeval strata of the Czech Republic, but similar to a tooth from the SCB and mosasauroids are found. Notably, neither protostegids, which are important faunal components in the SCB and BCB, nor elasmosaurids, which are likewise present in both basins are represented in the material from the lower (lower middle?) Turonian of the Opole area.

76 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday plenary poster session V18

BRIEF REVIEW OF THE LATE CRETACEOUS MARINE VERTEBRATES FROM THE REPUBLIC OF MOLDOVA T. Obadă1,2 and Ş. Vasile3*

1 Academy of Sciences of Moldova, Institute of Zoology, 1 Academiei Street, MD-2028 Chişinău, Republic of Moldova 2 National Museum of Ethnography and Natural History, 82 Mihail Kogălniceanu Street, MD- 2009 Chişinău, Republic of Moldova 3 University of Bucharest, Faculty of Geology and Geophysics, Department of Geology,Laboratory of Paleontology, 1 Nicolae Bălcescu Avenue, 010041 Bucharest, Romania *[email protected]

Keywords: Late Cretaceous, Pliosauridae, Lamniformes,

Upper Cretaceous deposits crop out on small areas in the Republic of Moldova, mostly along the western bank of the Dniester and its tributaries. Although fairly well studied during the past two centuries, these deposits, generally referred to as Cenomanian in age, yielded scarce vertebrate remains. The most notable discovery in this respect is that of “ichthyosaur remains”, mentioned in the 1950’s by Suhov, including four teeth with a particular ridged outer surface, found near the city of Otaci, at a site called “La Izvoare” Ravine.

One of the “ichthyosaur” teeth was found in the collection of the Museum of Ethnography and Natural History (MENH) from Chişinău, and was available for this study. The shape, size, and especially the particular ridged pattern of the tooth surface are consistent with the dental morphotype generally referred to as Poliptychodon interruptus, a brachauchenian pliosaur reported from quite a few Upper Cretaceous sites of Europe. The same beds from Otaci yielded a partial vertebral centrum, which bears a striking resemblance to the markedly amphicoelous elasmobranchian vertebrae. The central area of the partial centrum also exhibits growth rings, but its edges are broken, which hinders a definitive assignment to sharks.

The age of the conglomeratic bed that yielded the Poliptychodon teeth and the putative shark vertebra is well supported by the invertebrate fauna it contains, namely the Cenomanian-indicative cephalopods Praeactinocamax primus and Mantelliceras mantelli.

Another Upper Cretaceous site, located further south along the Dniester, is Bechir’s Ravine near Soroca, where fragments of shark teeth were found. The general morphology of these teeth, albeit lacking the root, is similar to that seen in the anterior teeth of Cretoxyrhina, a well-known and widespread Cenomanian- Campanian lamniform shark. The Upper Cretaceous deposits from Soroca are also dubbed as Cenomanian, but the presence of the Campanian-indicative Belemnitella mucronata in the collections of the MENH suggests that a revision of the local stratigraphy might be in order.

The discovery of pliosaurid and lamniform teeth suggests that the Upper Cretaceous deposits from the Republic of Moldova might have in store more vertebrate remains, and should therefore be more intensely scrutinized. 77 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday plenary poster session V19

NEW INTERPRETATIONS OF THE HOLOTYPE OF ARGANASUCHUS DUTUITI JALIL AND PEYER 2007 (ARCHOSAURIA, PSEUDOSUCHIA), AND ITS TAXONOMIC IMPACT A. Lecuona1* and N.-E. Jalil1,2

1Sorbonne Universités CR2P, CNRS-MNHN-UPMC, Département Histoire de la Terre, Muséum national d'Histoire naturelle, CP38, 57 rue Cuvier, 75005 Paris, France 2Université Cadi Ayyad, Fac. Sci. Semlalia, Morocco

*[email protected]

Keywords: Argana Basin, phylogeny, diagnosis, Pseudosuchia, “rauisuchids”

The hypodigm of the pseudosuchian “rauisuchid” Arganasuchus dutuiti consists of several cranial and postcranial isolated elements coming from the Timezgadiouine Formation (Carnian, Upper Triassic) of the Argana Basin, Morocco. Its holotype is a putative proximal fragment of right pubis (MNHN.F AZA 904), bearing a diagnostic anteroventral orientation of the acetabular rim parallel to pubic shaft and a large contribution to the acetabulum. In the last decade, several new pseudosuchians and “rauisuchians” have been discovered and important advances on their anatomy and phylogeny were made. A complete revision of Arganasuchus and its phylogenetic relationships are being conducted under the light of the current knowledge. The revision of the holotype allows for the recognition of the pubic peduncle of the ilium fused to the proximal pubis, as suggested but then rejected by the original authors. The contact between the bones is not visible due to a fully sutured contact, a rare condition among pseudosuchians but already reported in North American poposauroids. Consequently, both formerly supposed diagnostic characters cannot be considered as such, the large acetabular region mostly or completely belongs to the ilium and the pubic portion of the acetabular rim is anterodorsally inclined like in other pseudosuchians. However, this reinterpretation highlights other diagnostic characters, such as a considerably deep longitudinal groove in the anterior surface of the pubic peduncle; an anterolateral short groove lateral to the previous one; an anteriorly wide pubic tubercle. Consequently, Arganasuchus dutuiti can still be considered a valid taxon. The inclusion of MNHN.F AZA 904 in a phylogenetic analysis along with Stagonosuchus and six new characters (489 characters, 90 taxa; data matrix derived from a large scope archosauriform analysis) depicted it as a basal Loricata and sister taxon of . The latter group is supported by one of the new characters, presence of a longitudinal groove on the anterior surface of the pubis. Loricata shows a polytomy formed by Prestosuchus, Saurosuchus, Stagonosuchus, Batrachotomus+MNHN.F AZA 904, and Fasolasuchus+more derived loricatans, which is due to the instability of Stagonosuchus. The current revision of Arganasuchus is highly important for a better understanding of the Late Triassic assemblages of Northern Africa.

78 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday plenary poster session V20

REDESCRIPTION OF A TELEOSAURID (CROCODYLOMORPHA; THALATTOSUCHIA) SKULL FROM THE LIASSIC OF HOLZMADEN, GERMANY T.H. Trapman1,2*

1Utrecht University, Faculty of Geosciences, Heidelberglaan 2, 3584 CS Utrecht, The Netherlands 2Naturalis Biodiversity Center, Darwinweg 2, 2333 CR Leiden, The Netherlands

*[email protected]

Keywords: Steneosaurus, Teleosauridae, Thalattosuchia, Holzmaden, Liassic

An excellently preserved teleosaurid (Crocodylomorpha; Thalattosuchia) skull, discovered during the late 19th century in Liassic sediments of the Posidonia Shale Formation, has been in the Utrecht University Depot natural history collection for over a century. It was formerly only described as Teleosaurus sp. (collection number G337-1894). The aim of this study is to provide a detailed description of the skull morphology, and assigning the specimen to a species by comparison of morphological characters and morphometric data with known teleosaurid species from the Early Jurassic of southern Germany. Five potential species are recognized from the Holzmaden area to which this specimen may belong: Steneosaurus gracilirostris, Steneosaurus brevior, Steneosaurus bollensis, Pelagosaurus typus, and Platysuchus multiscrobiculatus.

Comprehensive summary of existing morphometric datasets of these species allowed for a detailed comparison. Based on the proportions, overall shape and size of the skull and its individual elements, it can be stated with a high degree of certainty that G337-1894 belongs to Steneosaurus bollensis, which is the most common species of teleosaurid in the Posidonia Shale Formation (Lias ε) of South Germany. The large total skull length, large proportional size of the rostrum (positive allometric growth), and small relative size of the orbits (negative allometric growth), suggests this individual was in a relatively late ontogenetic stage. A detailed description of ontogenetic development in the genus Steneosaurus however, requires more cranial and post-cranial material and histological data of particularly early ontogenetic stages.

79 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday plenary poster session V21

COMPARATIVE ENDOCRANIAL MORPHOLOGY OF TWO THALATTOSUCHIAN (CROCODYLOMORPHA) SPECIES: STENEOSAURUS BOLLENSIS AND CRICOSAURUS ARAUCANENSIS Y. Herrera1,2*

1Bayerische Staatssammlung für Paläontologie und Geologie, Richard-Wagner-Straße 10, 80333 Munich, Germany 2National University of La Plata, CONICET-División Paleontología Vertebrados, Museo de La Plata, Facultad de Ciencias Naturales y Museo, La Plata, Argentina

*[email protected]

Keywords: Teleosauridae, Metriorhynchoidea, Mesozoic, neuroanatomy

Thalattosuchians are Mesozoic crocodylomorphs mainly known from marine deposits from the Lower Jurassic till the Lower Cretaceous. Thalattosuchia comprises two main lineages, Teleosauridae and Metriorhynchoidea, with the latter encompassing the fully pelagic forms, the Metriorhynchidae clade. Here, I present data on virtual endocasts obtained after CT scanning of two Thalattosuchia specimens: Steneosaurus bollensis (BSPG 1984 I 258) and Cricosaurus araucanensis (MLP 72- IV-7-1), and provide a comparative analysis. Three-dimensional reconstructions of the brain, , carotid canal, and paratympanic sinus system were made. Both endocasts show that the brain is sub-horizontally displaced, with not well-marked pontine and cerebral flexures. A conspicuous tube-like cavity that connects with the dorsal longitudinal venous sinus has also been identified in both taxa. In S. bollensis the carotid foramen pierces the exoccipital with a posterolateral direction, differing from the condition in metriorhynchids, on which the foramen has a strictly posterior direction. In the S. bollensis the carotid canals are incompletely ossified in the middle region, being impossible to differentiate the carotid canals from the pneumatic cavity, whereas in C. araucanensis the canals are completely ossified. The semicircular canals of the inner ear of both specimens are aligned in approximately orthogonal planes in three-dimensional space. The canals in S. bollensis are thinner than the ones of C. araucanensis (narrower in diameter relative to overall labyrinth size). In both taxa the anterior semicircular canal is longer than the posterior, which in turns is similar in size to the lateral canal. Several interconnected diverticular expansions from the pharyngotympanic sinus and median pharyngeal sinus related to the paratympanic sinus system have been identified in S. bollensis. In C. araucanensis only a few of them were identified, likely more related to the low resolution of the CT scan than to an anatomical feature of this specimen. The endocast morphology of Steneosaurus bollensis indicates the presence of several features previously suggested to be exclusive of the Metriorhynchidae clade. These findings highlight the limited knowledge about the neuroanatomy of Thalattosuchia, indicating that it is a topic that still needs to be further explored.

80 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday plenary poster session V22

AN ALLIGATOROID FROM THE PALEOCENE OF NORTH AFRICA AND POST-EXTINCTION DISPERSAL OF ALLIGATORS P. Russell 1* and N.R. Longrich1

1Department of Biology and Biochemistry and Milner Centre for Evolution, University of Bath, Claverton Down BA2 7AY, United Kingdom

*[email protected]

Keywords: Crocodylia, Alligatoroidea, Morocco, Paleocene, biogeography

The Alligatoroidea (alligators and caimans) are known from the Late Cretaceous to the present day. Unlike other members of the Eusuchia, which are saltwater tolerant, the alligators are an entirely freshwater clade, confounding our understanding of the groups dispersal patterns during their history. The Cretaceous record of the group is known only from . It is not until after the K-Pg mass extinction that we see a more widespread dispersal, notably the caimanines into South America, the diplocynodontids in Europe, and alligatorids in Asia. To date, there has been no conclusive alligator remains from Africa, as material has been deemed too fragmentary to be diagnostic. Here we describe a new species of alligator from the Paleocene deposits of Morocco. The specimen exhibits exceptional preservation of the skull, with an associated lower jaw and vertebrae material. Associated deposits in Morocco are typically marine and host an exceptionally diverse fossil fauna, including other crocodylomorph groups (Gavialoidea and Dyrosauridae) in the aftermath of the K-Pg extinction. Freshwater taxa, which include this new species, are comparatively rare. Phylogenetic analysis places this new species within the Diplocynodontidae, amongst the stratigraphically earliest members of the clade. This suggests a dispersal into Africa in the early Paleogene and adds to our understanding of the biogeography of the clade. The appearance of alligators around the world in the Paleogene is suggestive of a post-extinction dispersal, with alligators dispersing to occupy niches left vacant by the K-Pg mass extinction.

81 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday plenary poster session V23

FLYING STRAINERS AND A NEW FILTER-FEEDING PTEROSAUR FROM THE LATE JURASSIC OF WATTENDORF (BAVARIA, GERMANY) E. Frey1*, M. Mäuser2 and H. Tischlinger3

1State Museum of Natural History Karlsruhe, Erbprinzenstraße 13, 76133 Karlsruhe, Germany 2Naturkunde-Museum Bamberg, Fleischstr. 2, 96047 Bamberg 3Tannenweg 16, 85134 Stammham

*[email protected]

Keywords: Monofenestrata, Pterodactyloidea, filter feeder, Late Jurassic, Bavaria

Filter-feeding pterosaurs are recognized by the long and slender teeth, which appear unable to apply bite forces. The size of the minimum food items are determined by the interdental distance and the slenderness of the teeth. Therefore, filter-feeders that could strain plankton out of the water have plenty hair- or needle-shaped teeth. Only seven plankton strainers in three genera have been described insofar: Gnathosaurus subulatus and G. macrurus (Late Jurassic Germany and England), Ctenochasma gracile, C. taqueti and C. roemeri Late Jurassic of Germany, France and England and Pterodaustro guinazui (Early Cretaceous of Argentina). The straining apparatus of Pterodaustro is restricted to the mandible of the strongly curved jaws and suggest a slow scooping movement of the head during feeding due to the flexible bristle-like teeth. In contrast the European strainers possess the same type of teeth in both, the upper and lower jaws suggesting a grabbing action. Here we present a new species of plankton feeding pterosaurs from the Late Jurassic laminated limestone of Wattendorf (Bavaria, Germany) with numerous densely standing needle-shaped, but blunt and apically inflated interdigitating teeth in the slightly curved upper and lower jaws. The rostral terminus of the jaws is expanded and the tooth rows open at the very tip of the rostrum. Additionally the terminus of the lower jaw seems to be deflected. Apparently there was an edentulous opening at the tip of the rostrum, which could have allowed the hitherto unnamed pterosaurs to skim through the water pressing nutrient-rich water into the slightly opened jaws. The water rinsed out laterally through the interdigitating teeth, which held back the food items. According to the constriction caudal to the rostral aperture the food item could have had a maximum diameter of about five millimeters and a minimum diameter of about one millimeter according to the distance of the teeth. A functionally similar feeding apparatus today is only known from balaenid Mysticeti.

82 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday plenary poster session V24

DESCRIPTION AND CORRELATION OF JURASSIC VERTEBRATE-BEARING PROFILES IN THE TURPAN BASIN (XINJIANG, NW-CHINA) D. Falk1* and O. Wings²

1State Museum of Natural History Karlsruhe, Erbprinzenstraße 13, 76133 Karlsruhe, Germany ²Hannover State Museum, Hannover, Willy-Brandt-Allee 5, 30169 Hannover, Germany

*[email protected]

Keywords: excavation, Qigu Formation, stratigraphy, sedimentology, fluvio-lacustrine deposits

From 2007 to 2012, paleontological fieldwork of the Sino-German Paleontological Cooperation Project took place in the Turpan Basin (Xinjiang, China). Here, we investigate the sedimentology of nine vertebrate macrofossil localities in desert areas east of Shanshan city. All sites yield steeply inclined fluvio-lacustrine deposits that are stratigraphically located in the Middle Jurassic Qigu Formation. The largest distance between the sites is approx. 2 km. Almost all dinosaur localities lie within a relatively narrow part of red-colored sediments, approximately 200 to 250 m below the top of the Qigu Fm. Comparison of the geological profiles reveals several fining- up cycles of fine sandstones, siltstones to fossil-bearing mudstone layers with thicknesses of several dm. Internal planar bedding indicates temporary shallow water flows with low velocities, whereas elsewhere lithoclasts, small-scale cross-bedding structures, and obscured, internal small-scale ripple marks indicate higher water velocities. Pedogenesis also is present and calcareous nodules indicate groundwater rise under arid or semiarid climatic conditions. The sediments probably represent crevasse splays on a fluvial floodplain in combination with small ponds of residual water from former floods.

All sites except one have produced dinosaur material, dominated by sauropod bones, but also comprising theropod bones and teeth. Only one ornithischian bone was discovered in the area. Turtles are by far the most abundant vertebrate fossils. A spectacular site is Mesa Chelonia, a turtle taphocoenosis without any dinosaur remains. Of particular importance is the Boneanza site which yielded the articulated holotype of the gigantic mamenchisaurid sauropod Xinjiangtitan shanshanesis. Other fossils from Boneanza include articulated fish remains, isolated theropod teeth, and bivalves. Stratigraphically older is the Shanshan Tracksite, an early Middle Jurassic (Qiketai Formation) theropod tracksite which represents the first major record of dinosaur footprints (Changpeipus) from Xinjiang. Picking of screen-washed concentrate from sediment in the vicinity of the tracksite shows that it is rich in microvertebrates. Finds include teeth of hybodont sharks, semionotid fishes, ?lacertilians, crocodilians, basal ornithopod and thyreophoran dinosaurs, as well as amphitheriid and ?docodont mammals.

The Mesozoic sediments in the Turpan Basin continue to have a high potential for important vertebrate fossil finds.

83 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Early Tetrapods Awaken

SHAPE-SHIFTING ENDOCASTS: 3D MORPHOMETRICS AND TRENDS IN PALAEOZOIC FISH ENDOCASTS

S. Henderson1*, T. Challands2, A. Clement3 and S. Giles4

1University of Edinburgh, School of Biological Science, Ashworth Laboratories, Charlotte Auerbach Road, The King's Buildings, Edinburgh, EH9 3FL, Scotland 2University of Edinburgh, School of Geoscience, Grant Institute, The King's Buildings, James Hutton Road, Edinburgh, EH9 3FE, Scotland 3Flinders University, School of Biological Sciences, Adelaide, South , GPO Box 2100, Australia 4University of Oxford, Department of Earth Sciences, Oxford, OX1 3AN

*[email protected]

Keywords: Palaeozoic Dipnoi, Devonian, geometric morphometrics, endocast

In the last two years there has been a proliferation of digital endocasts derived from Palaeozoic fishes. The potential wealth of phylogenetic data that endocasts contribute to resolving phylogenies is great, however until such characters can be reliably identified morphometrically, the efficacy of the information they carry remains somewhat subjective. Here we present the first quantitative, morphometric analysis of a suite of Palaeozoic fish endocasts comprising specimens from the Dipnoi and basal Actinopterygii. This study is also the first to attempt to quantify palaeoneurological characters of Palaeozoic fish using geometric morphometrics on digital endocasts.

We demonstrate morphological changes of both the endosseous inner-ear labyrinth and the main body of the endocast using 3D geometric morphometrics. Our analyses support the concept of several endocast characters that reliably differentiate between the Dipnoi and Actinopterygii, yet also between dipnoan taxa regarded as primitive and derived. Specifically, in the labyrinth: the size of the utricular recess, the size of the ampullae, and the orientation of the semicircular canals; and, in the endocast: ventral expansion of the telencephalon and the configuration of the olfactory nerves. More derived dipnoans display greater ventral expansion of the telencephalon and larger, more differentiated utricular recesses, with trends in the olfactory nerves less easily defined.

A new detailed anatomical description of the endocast of ‘Chirodipterus’ australis, an Upper Devonian dipnoan, is also given and is shown to exhibit a suite of characters more typical of Lower and Middle Devonian dipnoan taxa. Notably, the small utricular recess is unexpected for a taxon of this age, whereas the ventral expansion of the telencephalon and sessile olfactory bulbs are more typical of contemporary taxa. The presence of such 'primitive' characters in C. australis supports its relatively basal position as demonstrated in the most recent phylogenies of Devonian Dipnoi.

84 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

EARLIEST CARBONIFEROUS DIPNOI: POST-HANGENBERG RECOVERY AND THE DAWN OF A NEW ERA OF LUNGFISH

T.J. Challands1*, T.R. Smithson2, J.A. Clack2, J.E.A. Marshall3 and C. E. Bennett4

1University of Edinburgh, School of Geosciences ,James Hutton Road, Edinburgh, EH9 3FE, Scotland 2University Museum of Zoology, Downing Street, Cambridge, CB2 3EJ, UK 3University of Southampton, National Oceanography Centre, University Road, Southampton, SO17 1BJ, UK 4Department of Geology. University of Leicester, University Road, Leicester, LE1 7RH, UK

*[email protected]

Keywords: Dipnoi, Tournaisian, Carboniferous, Hangenberg, Romer’s Gap

A new specimen of a large skull from lowest Tournaisian deposits (VI palynozone, less than 1 My from the Devonian-Carboniferous boundary) of southern Scotland represents the earliest occurrence of a dipnoan following the Hangenberg in the Famennian. Comprising a near-complete cranial dermal skeleton, the arrangement of the skull roofing bones resembles that of the well-known Carboniferous genus Ctenodus. Alongside other dipnoan remains, including tooth plates, operculi, isolated skull bones and postcranial material, the fossil record of lungfish in 'Romer's Gap' can now be demonstrated to comprise a diverse and morphologically disparate fauna indicating rapid post-extinction recovery and niche partitioning by the middle-upper Tournaisian (CM palynozone). The resemblance of the new specimen to Ctenodus implies that lungfish skull morphology was evolving at a slower rate than dental morphology. The new lowest Tournaisian specimen does not exhibit reduction of skull roofing bones seen in younger Carboniferous and more recent lungfish whereas new Tournaisian lungfish dentitions are highly variable in form. Diversification of lungfish dentition is associated with the contemporaneous appearance of the earliest record of an established brackish to freshwater bivalve fauna, a possible key food source for lungfish. Furthermore, the new material from the lowest Tournaisian is large with estimates of full body size ranging between 20 cm to exceeding 1 meter in length. Tournaisian lungfish were among the largest organisms to inhabit the waters in the post-Hangenberg extinction world and demonstrate a trend in increasing body size following extinction rather than body size stasis or reduction as previously proposed for sarcopterygian taxa at this time.

85 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

PREDICTING THE LIKELY LOCATION OF NEW EARLY TETRAPOD SITES: LESSONS FROM RECENT DISCOVERIES IN ROMER’S GAP T.R. Smithson1*, C.E. Bennett2, J.A. Clack1, S.J. Davies2, T.I. Kearsey3, J.E.A. Marshall4, D. Millward3 and J.E. Sherwin2

1University of Cambridge, University Museum of Zoology, Downing Street, Cambridge, CB2 3EJ, UK.

2University of Leicester, Department of Geology, Leicester, LE1 7RH, UK. 3British Geological Survey, The Lyell Centre, Research Avenue South, Edinburgh EH14 4AP, UK.

4University of Southampton, Ocean and Earth Science, National Oceanography Centre Southampton, Southampton, SO14 3ZH, UK

*[email protected]

Keywords: Carboniferous, palaeosols, sandy-siltstones, overbank-flooding

Prior to Stan Wood’s discoveries at East Kirkton Quarry in 1984, the study of early Carboniferous tetrapods in the UK was based entirely on fossils collected from coal, ironstone and oil shale workings. Recent discoveries in the Ballagan Formation (Tournaisian) of the Scottish Borders have shown that new sites are not restricted to economically important deposits but may occur wherever the Carboniferous is exposed. Twelve horizons containing early tetrapods have been discovered so far in the Tweed Basin in a variety of lithologies, all of which are associated with coastal lakes and marshes. The most common lithology that preserves tetrapods is sandy siltstone. Beds of this composition usually overlie palaeosols and were deposited in seasonal flooding events as cohesive flows. This taphofacies contains the most abundant and rich fossil deposits in the early Carboniferous. It includes a diverse variety of partially articulated large and small tetrapods, associated with fishes, bivalves, ostracods, arthropods and abundant plant remains. The conglomerate lags at the base of fluvial channel sandstones often preserve dissociated tetrapod elements. Fossiliferous, finely laminated siltstones are rare, but these do contain the most completely preserved fossils. During the early Carboniferous, the Tweed Basin was surrounded by emergent land, represented today by the Southern Uplands to the north and the Cheviot Hills to the south. The lower lying areas between were traversed by large rivers and housed permanent and temporary freshwater lakes. Overbank flooding and periodic high rainfall provided sediment deposition in which the fauna and flora was preserved. During times of lowland flooding, proximity to land was critical in providing refuge for the terrestrial fauna. These conditions provided the ideal environment for the evolution of terrestrial ecosystems and in northern Britain were fairly stable over millions of years. Similar conditions must have prevailed throughout the Carboniferous, especially at higher equatorial latitudes away from the coal swamps. Basins adjacent to upland areas in which palaeosols form a significant component of the succession are where we should be looking for new early tetrapod sites.

86 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

MORPHOLOGICAL INNOVATIONS IN THE EARLIEST POST- DEVONIAN TETRAPODS: ADAPTATIONS FOR TERRESTRIALITY? J.A Clack1* and T.R. Smithson1

1University Museum of Zoology, Downing St., Cambridge, CB2 3EJ, UK

*[email protected]

Keywords: Romer's Gap, braincase, jugal, tabular, humerus

Although the earliest Carboniferous Tournaisian stage has been considered a depauperate interval for vertebrates and particularly tetrapods (known as “Romer’s Gap”), our recent work has unearthed a rich and diverse suite of Tournaisian vertebrate fossils from the UK. These include many new tetrapod genera, spread across the phylogenetic and morphological divide between Devonian and Carboniferous forms. While some retain plesiomorphic characters, others demonstrate a number of innovations to morphology that are seen for the first time in the Tournaisian tetrapods: 1) Parasphenoids that cross the ventral cranial fissure and underplate and fuse to the braincase base, in one case also underlying the basipterygoid processes. This may relate to strengthening and consolidating the braincase as it is increasingly subjected to gravity and the forces of muscle contraction during head movements on emerging from the water. 2) An exoccipital separate from the basioccipital in contrast to the conjoined ex- and basioccipital of more primitive tetrapods may relate to development of a more mobile occipital joint, also associated with more flexible head movement. 3) Deeply emarginated jugals may imply enlarged orbit and eye sizes compared to Devonian forms and suggest the increased importance of vision in terrestrial forms. It may also mark a shift from a more dorsolateral to a more lateral field of vision in a deeper rather than a flattened skull shape. 4) Humeri in which the foramen for the brachial artery and median nerve passes from the dorsal to the ventral side of entepicondyle. This is related to 5), humeri that show greater than 25 degree twist, related to increasing stride length, and 6) humeri in which the deltopectoral crest becomes more proximally placed and which show the development of a true shaft. 7) The earliest occurrence of a five-digit manus. Some of these innovations, seen for the first time in small animals, may be directly linked to increasing terrestrialization among post-Devonian taxa.

87 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

AN EXCEPTIONALLY WELL-PRESERVED SKULL OF STANOCEPHOSAURUS (TEMNOSPONDYLI: CAPITOSAURIA) LEADS TO A NEW HYPOTHESIS ON THE OPERATION OF THE TEMNOSPONDYL AUDITORY SYSTEM T. Arbez1* and J.-S. Steyer1

1Muséum national d’Histoire naturelle, Centre de Recherches en Paléobiodiversité et Paléoenvironnements (CR2P), UMR 7207 CNRS-MNHN-UPMC, CP 38, 8 rue Buffon, 75005 Paris, France

*[email protected]

Keywords: amphibian, otic, , temnospondyl, tomography

Temnospondyls form an extinct clade of non-amniotic tetrapods, similar in shape to giant salamanders or crocodiles. Most of them were amphibious and inhabited various palaeoecosystems such as rivers, lakes and swamps.

The auditory performance of temnospondyls is still strongly debated. The temnospondyl stapes is often considered a middle ear bone that is close to the anuran condition and linked with an acoustic function adapted to airborne sound perception. However, this bone is sutured with the in several taxa. This peculiar condition questions the traditional acoustic function attributed to the stapes, which could also play a role in the support of a spiracular canal; a conduct supplying water to the internal cavity.

An exceptionally well-preserved skull of Stanocephalosaurus from the Triassic of Algeria could solve the debate. Its cranial exploration was conducted with the micro- CT scan AST-RX of the MNHN. The resulting 3D reconstruction reveals a highly detailed anatomy of the columellar cavity and stapes. The stapes is linked to a crest of the parasphenoid (a palatal bone) and situated laterally to the otic capsule, inside a columellar cavity connected to the buccal cavity by a small canal. This otic architecture leads to a new hypothesis of the stapedial function as part of a hearing system adapted to underwater sound perception.

This hypothesis explains why peculiar morphological features are shared by several temnospondyls (e.g. the dorso-ventral flattening of the stapes, or the delicate curved flange of the pterygoid bordering the columellar cavity) and involves few morphological changes compared to the common anuran auditory system.

88 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

TEMNOSPONDYLS, LISSAMPHIBIANS AND MIDDLE EARS D. Marjanović1*

1Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Humboldt-Universität, Invalidenstraße 43, 10115 Berlin, Germany

*[email protected]

Keywords: Temnospondyli, , middle ear, tympanum, stapes

The origin of the amphibian crown-group is most often sought among the temnospondyls. One character complex that is particularly often pointed to as evidence for such a relationship is the tympanic ear found in frogs, for which osteological correlates have been proposed in many temnospondyls, particularly the amphibamid .

A stapes in a tympanic ear is expected to be small and light; often just the base is ossified, while the long, thin tympanic process – much thinner than the paroccipital process or the basal plate of the parasphenoid – remains mostly cartilaginous. The distal end of this process is expected to lie in or close to the tympanic center for maximal response to its vibrations. The footplate either forms a hinge with the rim of the fenestra ovalis (as in frogs and crown-group diapsids), or acts as a piston within it (as in eucynodonts, incl. mammals).

Given these criteria, most temnospondyls did not have a tympanum, and the best candidate for having one is not Doleserpeton, but the dissorophid ?Broiliellus reiszi. Except in the latter, temnospondyl stapedes are remarkably well ossified, long (reaching the surface of the skull or nearly so), robust (comparable to clubs, handles, planks or pillars, not “rodlike” as they are often called) and immobile (currently unknown in ?B. reiszi). They even form a deeply interdigitated suture with the parasphenoid in mature eryopiforms (, Stanocephalosaurus, Platyoposaurus, ?) as well as, far rootward of Temnospondyli, in Greererpeton. Their distal ends, with the same exception, participate in the caudal wall of the temporal embayment. In Doleserpeton, the paroccipital process bears paired ridges that have been interpreted as support for a froglike cartilage ring around the tympanum, but may have held the stapes instead.

The distal end of the stapes may have plesiomorphically anchored -opening/ closing muscles as in Polypterus, an extant actinopterygian that breathes air through its spiracles. Hearing high-frequency airborne sound was a late, rare innovation restricted to some (not all) forms with fully terrestrial adults – within Temnospondyli, within Amniota, and also within Lissamphibia, where evidence of a formerly tympanic ear in salamanders, or albanerpetids has not been described.

89 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

COMBINED OSTEOHISTOLOGICAL AND COMPUTED FINITE ELEMENT ANALYSIS OF METOPOSAURUS KRASIEJOWENSIS (AMPHIBIA, TEMNOSPONDYLI) SKULL BIOMECHANICS K. Gruntmejer1,2*, C.O.D. Konietzko-Meier1,2,3, A. Bodzioch1,2, J. Fortuny4,5 and J. Marcé-Nogué6,7

1Opole University, Department of Biosystematics, Laboratory of Palaeobiology, Oleska 22, 45-052 Opole, Poland; 2Opole University, European Centre of Palaeontology, Oleska 48, 45-052 Opole, Poland; 3University of Bonn, Division of Paleontology, Steinmann Institute, Nussallee 8, 53115 Bonn, Germany; 4Institut Català de Paleontologia Miquel Crusafont, C\Escola Industrial 23, 08201 Cerdanyola del Vallés, Spain; 5Muséum National d'Histoire Naturelle, Bâtiment de Paléontologie, CP38, 8 rue Buffon, 75005 Paris, France; 6Universitat Politècnica de Catatalunya – Barcelona Tech, C\Colom 2, 08222 Terrassa, Spain 7 Centrum fur Naturkunde, University of Hamburg, CP: 20146 Hamburg, Germany

*[email protected]

Keywords: Temnospondyli, histology, finite element analysis, microanatomy, biomechanics

Three-dimensional Finite Element Analysis is a powerful tool in understanding ecomorphological patterns because it shows how deformation and stress distribution relates to different biological geometries. Over the last years, FE analysis has often been combined with Geometric Morphometrics. FEA may also be complemented with other techniques, such as osteohistological investigations, although this has thus far not been tested. In this respect, digital models can be directly assessed with histology, as bone microstructure reflects loading regime. We used Finite Element Analysis to deepen the understanding of the feeding ecology of the early tetrapod group Temnospondyli, and in particular of the Late Triassic taxon Metoposaurus krasiejowensis, with the aim to assess the stress pattern of the skull under different biomechanical loads. FEA investigations were conducted under three different biomechanical feeding simulations: bilateral, unilateral and lateral biting. In parallel, 18 thin-sections from the dermal and palatal bones of a well-preserved Metoposaurus skull were examined to collect information about the microanatomical variability of the selected bones. Bone thickness and bone porosity influence the biomechanical stress distribution. The bones lose strength and stiffness with increasing porosity; this can be partly compensated for by increasing the thickness of the structure. The bones from the anterior and posterior parts of skull roof, e.g. the , nasal, lacrimal, , quadratojugal and supratemporal, have the optimal histological properties (high thickness and low porosity) to resist biomechanical forces. The bones from the palatal side, e.g. and parasphenoid, possess the weakest mechanical resistance (low thickness and high porosity). The histological model of stress distribution was compared with the three- dimensional FEA model. Both methodologies reveal that unilateral biting was probably not performed by these animals. Osteohistological results suggest that Metoposaurus krasiejowensis was particularly optimized for lateral biting while FEA revealed that bilateral biting could also be supported by these animals. Regarding lateral biting, the high levels of stress simulated on the posterior parts of the skull palate correlates with the histological variability and microanatomical properties for most of the examined bones. 90 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

HISTOLOGICAL STUDIES AS THE KEY TO POPULATION ANALYSIS – THE EXAMPLE OF METOPOSAURUS KRASIEJOWENSIS FROM KRASIEJOW, SE POLAND E. Teschner1,2 and D. Konietzko-Meier1,2,3

1Opole University, Department of Biosystematics, Laboratory of Palaeobiology, Oleska 22, 45-052 Opole, Poland 2Opole University, European Centre of Palaeontology, Oleska 48, 45-052 Opole, Poland 3University of Bonn, Steinmann Institute, Nussallee 8, 53115 Bonn, Germany

*[email protected]

Keywords: Metoposaurus, bone histology, Krasiejów, Late Triassic

Histological data sheds light on various aspects of an animals` biology. Various histological studies have recently been applied on bones of the temnospondyl amphibian Metoposaurus krasiejowensis. However, the aim of these histological studies was slightly different: one analyzed the ecological adaptation in order to get information about biology and mode of life of this species, others focused on skeletochronological imprints or on skull mechanics of this animal.

Here we assess long bones of M. krasiejowensis including humeri, radii, ulnae, femora, tibiae and fibulae. The material originates from Krasiejów, a locality in southeastern Poland. Among the tested bones, two growth patterns are observed. The majority of bones represents Histotype I, which is characterized by an alteration of thick zones and annuli of the same thickness. Strong remodeling is present throughout the entire cortex with a vascularization that ranges from poor to moderate. Histotype II, visible in six humeri and one femur, is characterized by a highly vascularized and extremely thick zone. The annulus is incipient and in some bones completely absent in the entire section.

We do not support the hypothesis in which the histotypes, showing biologically different features, represent a taxonomic diversity. These results can be interpreted as intraspecific variability on the level of a population diversification. Here we favor a population diversification occurring due to a separation in space and/or time. The histotypes can be interpreted as two representatives of different populations of metoposaurids that were once living in the ponds of Krasiejów. A less likely explanation for the different growth patterns is sexual dimorphism, a hypothesis that is rather difficult to test.

91 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

INDIAN METOPOSAURID AMPHIBIANS: MORPHOMETRY, TAXONOMY AND DISPERSAL S. Chakravorti 1* and D.P. Sengupta1

1Indian Statistical Institute, Geological Studies Unit, 203, Barrackpore Trunk Road, Kolkata, West- Bengal 700108

*[email protected]

Keywords: metoposaurs, morphometry, taxonomy

Metoposauridae is an important Late Triassic temnospondyl family known from non- marine sediments of Europe, North-America, Morocco and Central India. In the early nineties, Adrian Hunt erected “Buettneria perfecta” to include all metoposaurids in which the lacrimals enter the orbital margin, which also included the Indian taxon Metoposaurus maleriensis. The genus “Metoposaurus”, sensu Hunt, was thus restricted to encompass only M. diagnosticus from Germany and M. bakery from North America whose lacrimals are excluded from the orbital margin. Recently, Thomas Sulej reported that some specimens of M. diagnosticus also exhibit lacrimals contributing to the orbital margins, as in Buettneria. Hence, the generic status of the Indian metoposaurids, assigned by Hunt to Buettneria, became questionable. Later, it was noted that the generic name Buettneria was preoccupied, and Koskinonodon represents the senior synonym. To resolve the taxonomic status of the Indian metoposaurids, the present work used several biometric studies applied by Colbert and Imbrie in the past and more recently by Sulej. This study shows that the Indian population exhibits a small morphological overlap with the North American and European populations. For further studies into character variation of the metoposaurids at species level, morphospaces were constructed in the R environment, based on the shapes of the skulls, lacrimals (their relation with the orbits), parietals and orbits. It was discovered that Indian forms occupy a distinct morphospace when a combination of lacrimal and parietal shape was considered. Hence, this combination of characters has been chosen to distinguish the Indian population. Other characters, such as comparatively closely spaced nares, canals without a loop behind the orbits, and tabular width reinforce this separation of the Indian population. Therefore, a multitude of characters now identifies the Indian taxon that travelled to India, far away from the main occurrence of the group in Laurasia during Late Triassic. The new genus and species Panthosaurus maleriensis is proposed here for the Indian metoposaurids, where the generic name means “a wandering beast”.

92 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

TRIASSIC TEMNOSPONDYL AMPHIBIANS OF INDIA: THEIR DIVERSITY, DISPARITY AND NON-ENDEMISM D.P. Sengupta1*

1Indian Statistical Institute, Geological Studies Unit, 203 Barrackpore Trunk Road, Kolkata 700108, India

*[email protected]

Keywords: temnospondyl, India, Triassic

Triassic temnospondyl amphibians in India are represented by different families, and include the Trematosauridae, Brachyopidae, Plagiosauridae, Lapillopsidae, , Benthosuchidae, Indobracyopidae, Capitosauridae (), ?Tupilacosauridae, and Chigutisauridae. This is a unique association encompassing both Laurasian and Gondwanan elements. The early Triassic deposits of India have yielded nine of the eleven families stated above. Only three of the Early Triassic families, the Brachyopidae, Capitosauridae and the Trematosauridae, survived up to Middle Triassic. No Early and Middle Triassic temnospondyl families are known to continue into the Indian Late Triassic. Within the Late Triassic, the Carnian has only produced “Buettneria”- a metoposaur taxon, and the Norian has preserved two chigutisaur taxa. India is the only locality to have preserved both northern metoposaurs and southern chigutisaurs in a single Gondwanan formation. Disparity among the Indian Triassic temnospondyls at the generic level is prominent and it is not geographically controlled. One Early Triassic plagiosaurid genus from India, Capulomala panchetensis, carries a bizarre projection at the posterior part of its mandible. A new species of the genus (Capulomala arcadiaensis) from coeval beds of Australia also possesses this projection. The lone Middle Triassic Indian brachyopid has a peculiar cheek emargination, which is also noticed in Vigilius from North America. One of the Late Triassic chigutisaurs from India, Compsocerops, has six horns, one at each of the two tabulars, and squamosals. Recently, similar horn-bearing chigutisaurs are reported from Late Triassic beds of Brazil. Indian temnospondyls exhibit a large variety of morphotypes and include specific morphologies found in various other geographical domains, which attests to the non-endemic nature of this herpetofauna. During the Triassic, the palaeoposition of India remained almost halfway between Laurasia and Gondwana where the circum-Tethyan line was curved and cryptic. The non-endemism of Indian temnospondyls could be an artifact of this palaeoposition.

93 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

THE AMPHIBIANS STRIKE BACK – THE FIRST TEMNOSPONDYL FROM THE PALAEOZOIC OF ZAMBIA J. Steyer1* and C.A. Sidor2

1 Centre de Recherches en Paléoenvironnements et Paléobiodiversité, UMR 7207 CNRS- MNHN-UPMC, 8 rue Buffon, 75005 Paris, France 2Burke Museum and University of Washington, Seattle, Washington, U.S.A.

*[email protected]

Keywords: Gondwana, , end- mass extinction

Temnospondyls from the are best known from the Permian and Triassic of South Africa. Yet adjacent countries have also yielded interesting amphibians from coeval strata. In Zambia, temnospondyls are represented by the brachyopoid Batrachosaurus and the capitosaurs Cherninia and Stanocephalosaurus from the Triassic (Anisian) Ntawere Formation of the northern Luangwa Basin.

Recent exploration and fieldwork in the southern Mid-Zambezi Basin has led to the discovery of the first temnospondyl material from the Palaeozoic of Zambia. The material consists of two nearly complete skulls found in grey siltstones of the lower Madumabisa Mudstone Formation, which is considered middle Permian () in age, based on the co-occurrence of tapinocephalid dinocephalians. These medium-sized skulls belong to adult individuals and are very similar: both show a triangular general outline with a well-developed snout, relatively large orbits, very posterolateraly extended tabulars, a small internarial opening on the snout, and a large anteropalatal fenestra. They are attributed to a new species of rhinesuchid.

Rhinesuchids are basal stereospondyls previously known from the Karoo-aged rocks in South Africa and Malawi. This new discovery therefore enlarges the geographic distribution of the group. It also increases the diversity of the amphibian fauna in Gondwana prior to the end-Permian mass extinction.

94 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

SWIMMING WITH TEMNOSPONDYLS: A NEW APHANERAMMA (TEMNOSPONDYLI, TREMATOSAURIDAE) IN THE EARLY TRIASSIC OF MADAGASCAR J. Fortuny1,2* and J. Steyer1

1Centre de Recherches sur les Paléoenvironnements et la Paléobiodiversité, UMR 7207 CNRS-MNHN-UPMC, Bâtiment de Paléontologie, CP38, 8 rue Buffon, 75005 Paris, France 2Universitat Autònoma de Barcelona, Institut Català de Paleontologia M. Crusafont, CP: 08193 ICTA-ICP Building, Cerdanyola del Vallés, Barcelona, Spain.

*[email protected]

Keywords: Lonchorhynchinae, , Gondwana, Marine tetrapod recovery

Trematosaurs represent a peculiar and large clade of stereospondyl temnospondyls with a wide geographic distribution. However, they remain poorly known in Africa, where they are mainly recovered from South Africa (Trematosuchus and Microposaurus) and Madagascar (Wantzosaurus). We report a new and subcomplete skull of the longirostrine Aphaneramma from the Lower Triassic of Madagascar. With a skull length of about 40 cm, this new specimen may account for one of the largest known trematosaurians. It also enlarges the distribution of the genus, which was only known in Spitsbergen (A. rostratum) and Pakistan (A. kokeni) so far. A phylogenetic analysis of the trematosaurians suggests that Aphaneramma is closely related to the North American Cosgriffius. Because of its wide geographical distribution, Aphaneramma represents an important taxon for understanding the migration routes used by trematosaurs in general, and lonchorhynchines in particular. In this sense, the Malagasy occurrence of Aphaneramma provides new insights in the global distribution of the marine vertebrate fauna just after the great Permian-Triassic mass extinction.

95 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

MINIATURIZATION AND PAEDOMORPHOSIS IN TEMNOSPONDYL AMPHIBIANS C. Pérez Ben1,2*

1University of Buenos Aires, Department of Geological Sciences, Ciudad Universitaria Pabellón II, 1428 Buenos Aires, Argentina; 2State Museum of Natural History Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany

*[email protected]

Keywords: miniaturization, paedomorphosis, Temnospondyli

Miniaturization is a phylogenetic concept that has been defined as the evolution of extremely small adult size in a lineage. It does not simply imply the decrease of the body size, but also involves structural modifications to maintain functional efficiency. In addition to these modifications, miniature tetrapods frequently present a paedomorphic morphology, which results from the ontogenetic truncation involved in the size reduction.

Miniaturization is regarded as a key factor for the evolution of major clades. In particular, current hypotheses propose that at least two of the living amphibian groups originated from a Permian clade of small-sized dissorophoid temnospondyls. The smallest species of this clade have been described as miniaturized and many of their cranial features (e.g., proportionally larger interpterygoid vacuities) have been interpreted as consequences of miniaturization based on comparisons with miniature extant amphibians.

I tested whether the cranial morphology of the smallest dissorophoids reflects the functional constraints imposed by miniaturization as in extant amphibians. To address this, I compared the adults of the small species qualitatively and morphometrically with juveniles and adults of larger dissorophoids.

The adults of small dissorophoids do not present major skull rearrangements with respect to the ancestral condition and do not depart from the allometric trends of the clade, in contrast to what is expected under miniaturization. On the other hand, they resemble juvenile stages of larger forms in many traits, including those features previously associated with miniaturization, such as poor dermal ornamentation, overall thin dermal bones, rudimentary circumorbital elements and tabulars, reduction or loss of ectopterygoid, and oversized orbits and bony structures related to sensory capsules.

In conclusion, the reduction of body size in dissorophoids does not correlate with the morphological changes expected under the size constraints imposed by miniaturization. Instead, the morphology of the smallest forms is largely consistent with paedomorphosis. Therefore, truncation of the ancestral , and not miniaturization, is the factor that played a major role in the morphological evolution of small dissorophoids.

96 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

DIVERSITY AND BODY SIZE EVOLUTION IN PARAREPTILIA A. Elsler1*, M.J. Benton1, M. Ruta2 and A. Dunhill3

1University of Bristol, School of Earth Sciences, Queens Road, BS8 1RJ Bristol, UK; 2University of Lincoln, School of Life Sciences, Green Lane, Lincoln LN6 7DL, UK; 3University of Leeds, School of Earth and Environment, Leeds LS2 9JT, UK

*[email protected]

Keywords: Parareptilia, macroevolution, diversity, body size, Permo-Triassic mass extinction

Parareptilia is an enigmatic extinct clade of early tetrapods, known from the latest Carboniferous to the latest Triassic, which once was considered to contain modern turtles. Despite being a relatively small clade (114 known species), parareptiles were ecologically and morphologically diverse and geographically widespread. Aquatic forms are known as well, represented by the ‘basal’ Mesosauridae, famously used by Alfred Wegener in support of this theory of continental drift. Unlike other tetrapod clades and despite their ecological diversity, parareptiles have been largely neglected in macroevolutionary analyses.

Here we assess biodiversity and body size evolution of parareptiles through time. A comprehensive database of all known valid species of Parareptilia was assembled, containing data on body size, diet and stratigraphic range. Raw taxic diversity changes frequently during the Palaeozoic. A phylogenetic diversity estimate, based on an informal supertree of Parareptilia, offers an alternative interpretation of the fossil record, where the lineage richness increases more gradually throughout the Palaeozoic. Both raw taxic richness and phylogenetic diversity estimates show that the end-Permian mass extinction event did not have a substantial effect on the overall diversity of Parareptilia. Biodiversity peaks in the and decreases rapidly afterwards leading finally to the extinction of the clade in the latest Triassic.

Femur length was used as a proxy for body size and various likelihood models of continuous character evolution were fitted against the data. Preliminary analyses indicate a best fit of the OU model, followed by a trend model.

This study is part of a larger project that tries to disentangle fundamental macroevolutionary processes during the first 200 million years of tetrapod evolution with a current focus on body size evolution. Once data collection and analyses are complete, a comparison of body size evolution in all known clades of early tetrapods will be possible.

97 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

NON-DIAPSID EUREPTILIAN EVOLUTION: TRACK- TRACKMAKER CORRELATION OF AN IMPORTANT GROUP OF PALAEOZOIC TERRESTRIAL TETRAPODS L. Marchetti1* and S. Voigt1

1Urweltmuseum GEOSKOP / Burg Lichtenberg (Pfalz), Burgstraße 19, D-66871 Thallichtenberg, Germany

*[email protected]

Keywords: Non-diapsid eureptiles, ichnotaxonomy, appendicular , 3D acquisition techniques, geometric morphometrics

Tetrapod ichnology regards the study of footprints and trackways left by quadrupedal animals. The track record often constitutes the only evidence of living behaviour for extinct animals. This is the case of non-diapsid eureptiles, including the groups of "protorothyridids" and captorhinids. The former occupies a very basal position in reptilian evolution and comprises small lizard-like forms that evolved into (1) more robust and larger captorhinids and (2) progressively gracile, lizard-like diapsids. Captorhinids are the longest living, most widespread and most diverse late Palaeozoic group of reptiles. They show the acquisition of advanced features such as multiple rows of teeth associated with herbivory and larger body size. Fossil tracks related to these groups show a remarkable abundance and diversity suggesting significant anatomical evolution of the postcranial skeleton. However, a detailed analysis of the track record and of the limb skeletons of these groups is lacking, as is a reliable correlation between tracks and trackmakers. Therefore, we designed a comprehensive study on the track-trackmaker correlation of non-diapsid eureptiles, which could be important for the understanding of the evolution, biostratigraphy and biogeography of early reptiles. The two-year project is tri-partite: (1) Ichnotaxonomic revision of non-diapsid eureptilian tracks; (2) analysis of non-diapsid eureptilian appendicular skeletons; and (3) synapomorphy-based track-trackmaker correlation. The quantitative and qualitative analysis will benefit from the application of photogrammetry, 3D acquisition of fossil data as well as geometric morphometric methods.

98 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Insular Vertebrate Palaeontology

OVERSEA DISPERSAL TO MADAGASCAR: A GAMBLING MIRACLE? P.P.A. Mazza1*, A. Buccianti1 and A. Savorelli1

1University of Florence, Department of Earth Science, Via La Pira 4, 50121 Florence, Italy

*[email protected]

Keywords: oversea dispersion, natural rafting, insular colonization, land mammals, endemics

How organisms reach oceanic islands is a conundrum that is often easily resolved evoking natural rafting. Large masses of vegetation or dense mats of mosses would be dislodged by violent storms, carried down rivers and buoyed out to sea, forming temporary floating islands. Animals are either caught, or take refuge on them and are carried by favorable currents to sail rapidly across stretches of saltwater, crossing wide bodies of water otherwise insurmountable. The theory involves sporadic, accidental, and highly selective dispersal from a continent to an island. Owing to its intrinsically fortuitous nature, sweepstakes dispersal should generate unpredictable, oligotypical, and incomplete insular faunas compared to mainland communities.

The hypothesis might account for the arrival on islands of several plants, invertebrates, and small poikilotherm animals with low metabolic rates, low food requirements, and low water flux rates, and, therefore, greater ability to withstand oceanic distances. In contrast, it raises many more questions than it answers in the case of terrestrial mammals, which are quite more complicate to drift out to sea. Natural rafting is a plausible theory for the colonization of islands at any distance by land mammals only if an intricate network of variables is analyzed and a number of conditions satisfied. The phenomenon involves at least three sets of variables: 1) biological variables; 2) characteristics of the floating islands; 3) physical variables.

The biological variables implicate the physiological and eco-ethological characteristics needed to 1a) survive the stresses to which a rafter is subjected while crossing the ocean, and 1b) successfully and stably colonize an island. The island must enable travellers to find the minimum conditions for surviving the crossing and making it to the new land in satisfactory state of health. The physical variables include the oceanic and wind currents that can drive floating debris to an oceanic island. The point is: are the chances that all these conditions will be met so high to explain all or at least most of the successful colonizations of the oceanic or oceanic- like islands by land mammals?

99 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

THE INSULAR PARADOX L.W. van den Hoek Ostende1*

1Naturalis Biodiversity Center, P.O Box 9517, 2300 RA, Leiden, The Netherlands

*[email protected]

Keywords: island evolution, life traits, invasive species

In the course of their evolution, species continuously change in order to remain optimally adapted to their surroundings. This goes for mainland and island species alike. In addition, some trends seem the same for insular endemics all over the world, the most famous ones being gigantism and dwarfism in mammals, and flightlessness in birds. The general appearance of these adaptations is a testament that they help species to be perfectly adapted to the island life.

However, despite the inhabitants being adapted to the local environment, island ecosystems are notoriously vulnerable to outside influences. Although introduced species may also develop plagues on the mainland, they never have the disastrous effect on an entire ecosystem that we find on islands. This is a much a generality as the insular adaptations.

The three main drivers behind insular evolution are the absence of mammalian predators, competitive release and limitation of resources. Obviously, the introduction of cats, dogs and even pigs is a major threat to insular species that have lost their main defenses against predators. This effect is, however, strengthened by the adaptations to limited resources.

The main line of defense for most of the rodents is producing sufficient offspring to ensure the continuance of the species. Undoubtedly, this life trait must have led to regular overpopulation on islands upon colonization by a rodent species. The individuals that most likely survived the following starvation are the ones that invested in their own body, rather than in offspring. As a result, insular evolution led to species are typically K-select (i.e., at the slow end of the fast-slow continuum). This is, among other traits, corroborated by the signs of prolonged longevity of insular species as indicated by various authors. Even though they often occupy a wide niche on the island, insular endemics have the life traits of mainland specialists, making them as vulnerable to outside influences. Which leads us to the greatest paradox: insular evolution creates all-round specialists.

100 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

EVOLUTION OF THE SOUTHERN OCEANS WITH A FOCUS ON MADAGASCAR AND ITS RIF-RAF INHABITANTS (ISOLATED BY RIFTING OR RAFTING): TECTONICS AND CLIMATE CHANGE AS DRIVERS OF BIODIVERSITY J. Masters1* and M. de Wit2

1Africa Earth Observatory Network, Department of Zoology and Entomology, University of Fort Hare 2Department of Geological Sciences, Nelson Mandela Metropolitan University, South Africa

*[email protected]

Evolutionists have long sought to understand how Madagascar was populated by its eccentric vertebrate fauna. This ‘Garden of Earthly Delights’ apparently evolved in isolation after Madagascar rifted from Africa 170-140 Ma, and from India 50 Ma later. Vertebrate lineage divergence dates estimated from molecular sequences do not correlate well with the island’s tectonic history, and no molecular analysis of a vertebrate group has supported isolation of extant lineages by rifting, generating ingenious hypotheses of transoceanic dispersal by some form of rafting. Madagascar’s limited mammalian fauna comprises clades from three of the four superorders that emerged in the latest Cretaceous: the Afrotheria, Laurasiatheria and Euarchontoglires. The only superorder not represented on Madagascar is the endemic South American lineage, the Xenarthra, although the closest relatives of Malagasy iguanas, boas and dinosaurs were South American.

The evolution of the Indian Ocean is complex and dates back to 150 - 200 Ma ago. It has involved continuous rifting and rotation of microcontinents, including the early rifting of [India-Madagascar, Antarctica-Australia] from Africa (170-140 Ma); and the separation of India from [Australia-Antarctica] (120 Ma), Madagascar (84 Ma), Seychelles (66 Ma), Australia and Antarctica (43 Ma). Wisps of continental crust were isolated during this process, such as Mauritia that formed a ribbon-like structure between Madagascar and India between 83 and 66 Ma and may have formed a transient land-bridge. There are indications that more ancient micro-continents are buried under the Indian Ocean. Rifting between India and the Seychelles (66 Ma) was contemporaneous with massive volcanism in India and extinction across the Indian Ocean. How did the system recover from this catastrophic climatic perturbation? Where did survivors hide and how did they adapt to new ecosystems? What colonization routes were available for vertebrates during the complex evolution of the southern oceans?

101 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

A HYDROTAPHONOMIC MODEL FOR THE MARE AUX SONGES DODO-LAGERSTÄTTE IN MAURITIUS: MIRED IN TOXIC LOW-VISCOSITY MUD K.F. Rijsdijk1*, P.G.B. De Louw2, J.P.Hume3, J.K. Lubeek4,5, M.N. Jansen5, J. de Vos6, A. Janoo7, L.P.A.M. Claessens8,6, S.J. Kluiving5, A. Gill9, E. J. De Boer9 and H.J.M. Meijer10

1Computational GeoEcology Group, Institute for Biodiversity and Ecosystem Dynamics, University of Amsterdam, P.O. Box 94248, 1090 GE, Amsterdam, The Netherlands 2Department of Soil and Groundwater, Deltares, Postbus 85467, 3508 AL, Utrecht, The Netherlands 3Bird Group, Department ofLife Sciences, Natural History Museum, Akeman Street, Tring Herts, HP23 6AP, UK 4Department of Earth Sciences, Utrecht University, Heidelberglaan 2, 3584 CS Utrecht, The Netherlands 5Institute for Geo- and Bioarcheology, Faculty of Earth and Life sciences, VU University Amsterdam, De Boelelaan 1085, 1081 HV Amsterdam, The Netherlands 6Naturalis Biodiversity Center, Darwinweg 2, 2333 CR Leiden, The Netherlands 7Department of History and Political Science, University of Mauritius, Réduit, Mauritius 8College of the Holy Cross, Department of Biology, 1 College Street, Worcester, Massachusetts 01610, USA 9Palaeoecology & Landscape Ecology Group, Institute for Biodiversity and Ecosystem Dynamics, University of Amsterdam, P.O.Box 94248, 1090 GE Amsterdam, The Netherlands 10The Natural History Collections, University Museum of Bergen, University of Bergen, Postboks 7800, 5007 Bergen, Norway

*[email protected]

Keywords: miring, mass mortality, dodo, lagerstätte, hydrotaphonomy

During an anomalous megadrought episode 4200 years ago (4.2 ka), many thousands of vertebrates died within a century within a small shallow lake at Mare aux Songes in Mauritius. Radiocarbon dating suggests that the mass mortalities occurred within 80 years. Excellent conservation conditions included low energy conditions and near permanent anoxic burial in calcium-rich ground water soon after death. Plausible death scenarios include toxic water conditions related to cyanobacteria blooms and conditions promoting miring. However, the precise death and post-mortem conditions as well as the mechanisms underlying the miring process remain unknown.To assess this we mapped the 3D-orientation of vertebrate bones within the bone layer. All bones of all species are chaotically mixed, with no articulation and only one single case of association. Incompleteness of skeletons exceeds 90%. Bones show no preferred orientation and the mean dip of bones is generally significantly less than 22°. In situ excavation indicates that bone size sorting is absent.We attribute the chaotic orientations of bones to bioturbation by vertebrates wallowing in the exposed mud during drought. Loss of skeletal material is due to hydrotaphonomic processes that involved extreme seasonal fluctuations of groundwater (1 m). Geohydrological data suggest that the lake floor remained water- saturated (a soggy peat soil) as sea water welled up during dry spells. During wet seasons, the lake refilled and pneumatic elements buried in the mud floated away. Wind-driven currents transported bone material to the margins of the lake where the bones weathered away. The general low angle orientations of limbs (<22°) and post- mortem loss of limb material, indicate that miring in a high-viscous mud is unlikely. Instead, short-legged animals became trapped in a low-viscous mud and, lacking footholds within the cohesionless peaty-mud floor, animals drowned amidst a crowd of thirsty animals. Ultimately, death at Mare aux Songes was multi-causal, involving both poisoning and low-viscosity miring affecting mostly non-volant vertebrate species. 102 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

THE FEEDING BIOMECHANICS OF HAAST’S EAGLE (HARPAGORNIS MOOREI) A.H. van Heteren1,2,3*, P. Ross4, J.A. Ledogar1, M.R.G Attard1,5 D. Sustaita6, P. Clausen4, P. Scofield7 and S. Wroe1

1University of New England, School of Environmental and Rural Science, Earth Sciences Building, NSW 2351 Armidale, Australia; 2Universität Bonn, Steinmann Institut Bereich Paläontologie, Nußallee 8, 53115 Bonn, Germany; 3Zoologische Staatssammlung, Sektion Mammalogie, Münchenhausenstraße 21, 81247 München, Germany; 4University of Newcastle, School of Engineering, NSW 2308 Newcastle, Australia; 5University of Sheffield, Department of Animal and Plant Sciences, Alfred Denny Building, S10 2TN Sheffield, United Kingdom; 6Brown University, Department of Ecology and Evolutionary Biology, Box G-W, RI 02912 Providence, United States of America; 7Canterbury Museum, Geological Sciences, Rolleston Avenue, 8013 Christchurch, New Zealand

*[email protected]

Keywords: Insular gigantism, feeding biomechanics, eagle, vulture

Haast’s eagle (Harpagornis moorei) – an extinct predatory bird from New Zealand estimated to weigh nearly 50% greater than the largest modern eagles – is thought to have been capable of taking down large-bodied prey, including moa (~20-200 kg). Here, we use finite element analysis to assess the mechanical performance of Haast’s eagle relative to some modern eagle (Aquila audax, Haliastur spenurus, Hieraatus morphnoides) and vulture (Aegypius monachus, Vultur gryphus) species, during simulations of various killing and feeding behaviours. We applied loads simulating a simple bite using scaled muscle forces estimated from modern buzzards (Buteo buteo), as well as one lateral shaking behaviour, and two pulling (posterior, dorsoventral) behaviors. We found that all eagle species, including Haast´s eagle, exhibit relatively stiff crania during biting. However, with respect to posterior and dorsoventral pulling, Haast’s eagle shows affinities with the vultures by exhibiting reduced stiffness. These results suggest that Haast’s eagle may have engaged in a combination of hunting as well as scavenging behaviours that involved increased tearing and pulling of the tissues of large carrion. In the latter case, adaptations facilitating such behaviours may have been at the cost of having a relatively weaker cranium. Future studies on the biomechanics of talon gripping in Haast’s eagle will further elucidate hunting/feeding behaviour in this species.

103 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

HUMAN IMPACT CHANGES TOOTH WEAR OF MALAGASY BLACK RATS D.E. Winkler1*, T.H. Andrianasolo2,3, L.Andriamandimbiarisoa4, J.U. Ganzhorn2, S. J. Rakotondranary2, T.M. Kaiser1 and E.Schulz-Kornas1,5

1Center of Natural History (CeNak), University of Hamburg, Hamburg, Germany; 2Biocenter Grindel and Zoological Institute, University of Hamburg, Hamburg, Germany; 3Département Biologie Animale, Université d’Antananarivo, Antananarivo, Madagascar; 4QIT Madagascar Minerals, Fort Dauphin, Madagascar; 5Max Planck Weizmann Center for Integrative Archaeology and Anthropology, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany

*[email protected]

Keywords: surface texture, tooth wear, diet, Madagascar, island evolution

Dietary characteristics and environmental variables are important selective factors directing ecological diversification in rodents. On Madagascar, the introductions and spread of the commensal black rat (Rattus rattus) can be seen as example cases to study dietary niche occupation and dietary adaptation in an insular environment. We investigate how tooth wear as a measure of dietary adaptation of black rats differs between four distinct habitats (village, manioc field, spiny forest, rainforest) with different dietary resources. The rat teeth are an example case for tooth function adapted to a wide variety of food material properties and can be seen as multifunctional tools. We use the 3D surface texture analysis (3DST, using 30 surface roughness parameters according to ISO 25178) as a measure of dietary abrasiveness. 3DST is applied on the occlusal surface of the upper first molar of 37 black rat specimens. The rainforest sample displays less rough and less voluminous surface textures compared to the village samples as indicated by smaller values for height parameters (Sa, Sp, Sq), inverse areal material ratio (Smc) and volume parameters (Vm, Vmc, Vmp, Vv and Vvc). We therefore rank sampling areas from highest to lowest abrasiveness (village>manioc fields/spiny forest>rainforest). The rats from villages and rainforest differ to such an extent that one could have interpreted them to belong to different species. The pronounced difference between rats from human habitations compared to rats from associated fields or natural vegetation are interpreted to clearly indicate shifts in dietary niche occupation in relation to human impact. It indicates a high degree of variability in terms of ingesta abrasiveness as well as highly flexible tooth functionality.

104 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

MULTI PROXY DIETARY TRAIT RECONSTRUCTION IN PLEISTOCENE HIPPOPOTAMIDAE FROM THE MEDITERRANEAN ISLANDS (CETARTIODACTYLA, MAMMALIA) E.Bethune1*, T.M. Kaiser1 and D.E. Winkler1

1University of Hamburg, Centre of Natural History, Martin-Luther-King-Platz 6, Hamburg, Germany

*[email protected]

Keywords: dietary reconstruction, Hippopotamidae, island evolution, Pleistocene

Geographically isolated insular species constantly have to cope with energetic restrictions and often evolve adaptations that are distinct from their mainland ancestors. During the Pleistocene, several Mediterranean islands were inhabited by now extinct Hippopotamidae: Hippopotamus pentlandi from Sicily, Hippopotamus melitensis from Malta, Hippopotamus creutzburgi from Crete and Phanourios minor from Cyprus. They underwent diverse changes in locomotion, dentition and body size, indicating differences in their ecological niches. In this study, the palaeoecology of the dwarfed hippopotami is reconstructed by using two dental dietary proxies. Firstly, Dental Topography Analysis using Geographic Information Systems was applied to measure the slope of the occlusal surface, which provides large scale information on adaptive dietary traits. Secondly, 3D surface texture analysis (3DST) using 30 parameters according to ISO 25178-2 was employed for measuring the roughness of the enamel surface, which correlates with dietary abrasiveness. Mean slope values gradually increase from the larger hippopotami, H. pentlandi and H. melitensis, over the medium sized, H. creutzburgi, to the smallest, P. minor. Low mean slopes in the larger species reveal lower occlusal reliefs, representing rather horizontal jaw kinematics and complex enamel patterns as indicative of grazing species in ruminants. The highest mean slope values found in the smallest and lophodont P. minor are characteristic for a larger occlusal relief featuring compression-basins for extracting cell content. The relatively low and homogenous enamel surface textures in H. creutzburgi and H. pentlandi indicate dietary specialisation similar to the extant Hippopotamus amphibius, which feeds on fresh grasses near the shores, free of accumulated dust. In contrast, the rougher and more variable enamel surface textures in H. melitensis and P. minor suggest more generalistic dietary traits and/or dietary shifts linked to a climate change, resulting in an alteration of environmental and vegetal conditions from the Pleistocene to the Holocene.

105 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

LIGHTS AND SHADOWS IN THE EVOLUTIONARY PATTERNS OF INSULAR ELEPHANTINI M.R. Palombo1*

1Sapienza University of Rome, Department of Earth Sciences, P.le Aldo Moro 5, 00185 Roma, Italy

*[email protected]

Keywords: Mammuthus, Palaeoloxodon, Pleistocene, islands, evolutionary patterns

Dwarf elephants represent some of the most outstanding examples of the “island rule” (i.e., the graded phyletic body-size change undergone by vertebrates, mainly mammals, in isolated environments), and show some apomorphies suggesting homoplasy among endemic taxa that originated from different mainland species. Fossil elephants are especially appropriate subjects for analysing body-size changes in isolated environments because of the frequency with which they colonised islands, giving rise to several endemic taxa (more or less reduced in size in comparison with their own continental ancestor) even on a single island. Therefore, it is possible to compare different patterns of speciation and body-size modification, assessing the influence of various biotic and abiotic factors (e.g., distance to the mainland, topography of island and area, climate, , structure of insular communities) in driving the evolutionary process of insular species. This research focuses on dwarf Elephantini and aims to validate the generality of the observed evolutionary patterns, propose causal explanations, investigate on the parallel appearance of key morphological features correlated to insular dwarfism in different species/lineages, and to underline unaddressed questions. Results obtained provide evidence that: 1) body size variation in insular Elephantini is not directly dependent on the island surface area; 2) the structure of the insular fauna and ecological interactions with competitors are among the most influencing factors; 3) evolutionary pattern and body size shifts may be indirectly affected by isolation (constraining type and number mainland colonizers) and island physiography/environment (sometimes triggering adaptive radiation processes); 4) body size shifts mainly depend on adaptive strategy for a more efficient energy use under the special conditions of insular ecosystems, and to a lesser degree on the time in isolation. All in all evolutionary patterns of insular Elephantini suggest that in isolated environments, the shift in size of non- carnivorous species largely depends on the structure of the community and on the nature of vacant species. Various questions, however, remain unaddressed, particularly the challenging issue of separating anagenetic from adaptive radiation processes in those islands where phylogenetically close populations are recorded by a variety of elephants differing in size/morphotypes.

106 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

NEW UNEXPECTED BREAKTHROUGHS FROM THE MIOCENE ENDEMIC FAUNA OF THE APULIA PLATFORM (ITALY): A FAR MORE COMPLEX SCENARIO A. Savorelli1*, F. Masini2, P.P.A. Mazza1 and S. Colombero3

1University of Florence, Department of Earth Science, Via La Pira 4, 50121 Florence, Italy; 2University of Palermo, Department of Earth and Marine Sciences (DISTEM), via Archirafi 22, 90123 Palermo, Italy; 3University of Turin, Department of Earth Science, Via Valperga Caluso 35, 10125 Turin, Italy

*[email protected]

Keywords: Scontrone (Abruzzo), Gargano (Apulia), late Miocene, Insular endemics, Italy

Fossil remains of Deinogalerix from Scontrone (Abruzzo) and of "Apodemus" from Gargano (Apulia), but also other Gargano specimens recovered during 2005 – 2009 excavations, offer new insights into the late Miocene faunal communities of the so- called Abruzzo-Apulian palaeobioprovince.

Two new, different-sized species of Deinogalerix from Scontrone have dental proportions different from those of all the other species of Deinogalerix, which seem to rule out a direct ancestry of the Gargano clade. This discovery reveals that Deinogalerix had a longer history than previously supposed. It also shows that this erinaceid was monophyletic and radiated into a set of strongly endemized species.

The Gargano “Apodemus” has always been considered a non-endemic MN13–14 species because of the very high frequency of the t7 morphotypes. Nonetheless, a combination of primitive and advanced features separates it from Apodemus. This murid therefore forms a distinct evolutionary lineage that reached exclusive morphological traits in its most advanced representatives.

A newly discovered murine species shares a number of synapomorphies with Mikrotia, coupled with primitive features in common with several murid genera from the mainland. It therefore represents a candidate for the most likely ancestor of Mikrotia. Preliminary analyses indicate the continental genus Hansdebruijnia as the most suitable relative of this murid.

A new strongly endemized cricetid genus was found. It shows a peculiar combination of features that distinguish it from all the known mainland species. Comparisons distinguish it from cricetids younger than MN9 and indicate a possible stemming from a “Democricetodon-like” or “Cricetodon-like” Miocene mainland ancestor. These affinities suggest an age of dispersal spanning the MN1-9 interval.

The new overall evidence from Deinogalerix and from the new cricetid implies that the Apulia Platform faunas have a much longer history than previously supposed and polyphasic origin. The new murid and "Apodemus" also indicate that the "Terre Rosse" fissure fillings are not older than MN13 and that some of the ancestors of the fauna arrived from eastern areas.

107 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

THE GIANT BIRD GARGANTUAVIS: A CASE OF INSULAR EVOLUTION IN LATE CRETACEOUS EUROPE? E. Buffetaut1,2* and D. Angst3

1CNRS (UMR 8538), Laboratoire de Géologie de l'Ecole Normale Supérieure, 24 rue Lhomond, 75231 Paris Cedex 05, France; 2Palaeontological Research and Education Centre, Mahasarakham University, Maha Sarakham, Thailand; 3Palaeobiology Research Group, Biological Sciences Department, University of Cape Town,Private Bag X3, Rhodes Gift, 7701 South Africa.

*[email protected]

Keywords: Gargantuavis, Aves, Europe, Late Cretaceous

The giant flightless bird Gargantuavis philoinos is known from relatively scanty remains from a few localities in southern France and northern Spain, which in the Late Cretaceous were part of the Ibero-Armorican island. It has not been reported from other islands of the Late Cretaceous European archipelago. Its bone microstructure, indicating protracted cyclical growth, is reminiscent of that of various insular vertebrates, including non-avian dinosaurs, birds and mammals. The evolution of large flightless birds (e.g. moas in New Zealand, elephant birds in Madagascar etc.) on islands is a well known phenomenon, and Gargantuavis may be considered as a result of such a process on the Ibero-Armorican island. However, most insular giant flightless birds evolved in ecosystems that were devoid of large terrestrial predators. This does not apply to Gargantuavis, as it shared its habitat with various carnivorous theropod dinosaurs, including abelisaurids and dromaeosaurids. The place of the apparently graviportal and therefore relatively slow-moving Gargantuavis within such an ecosystem is unclear, all the more so that its diet is unknown. To gain a better understanding of the evolution of Gargantuavis in the peculiar insular environment of Late Cretaceous Europe, more information about its biology is needed. This can be achieved both by the discovery of additional fossil material and by investigations using approaches such as isotopic geochemistry, which can provide information about crucial aspects of the biology of this giant bird, such as its food.

108 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

LATEST CRETACEOUS MICROVERTEBRATE DIVERSITY OF THE TRANSYLVANIAN LANDMASS V.A. Codrea1, M. Venczel2*, A. Solomon1 and C. Fărcaș1

1Department of Geology, Faculty of Biology-Geology, Babeş-Bolyai University, 1 Kogălniceanu Str., 400084 Cluj-Napoca, Romania; 2Department of Natural History, Ţării Crișurilor Museum, Dacia Avenue 1-3, 410464 Oradea, Romania

*[email protected]

Keywords: Adaptive radiation, Lissamphibia, Maastrichtian, Multituberculates, Squamata

Uppermost Cretaceous (Maastrichtian) continental deposits from various Transylvanian sedimentary basins (Rusca Montană, Hațeg, Transylvanian) of western Romania bear a diverse assemblage of fossil vertebrates, including dwarfed dinosaurs, pterosaurs, turtles, crocodilians, birds, multituberculate mammals, fish, lissamphibians and squamates. New material from a number of microvertebrate localities allowed to document at least one albanerpetontid (Albanerpeton sp.), six different anurans (Alytidae: Paralatonia transylvanica, cf. Bakonybatrachus sp., cf. Eodiscoglossus sp., and a new species of Alytidae; Bombinatoridae: Hatzegobatrachus grigorescui; Pelobatidae indet.), six different families of lizards (geckonids, paramacellodids, scincids, borioteiids, teiids and anguimorphs), two different species of madtsoiid snakes (Nidophis insularis and Herensugea sp.) and three different taxa of endemic multituberculate mammals (Kogaionon ungureanui, Barbatodon transylvanicus and B.oardaensis). The assemblages indicate that part of this complex fauna originated from the cratonic Europe, North-America and also from Africa. Some groups, when reaching the Transylvanian Landmass, underwent an adaptive radiation and occupied different ecological niches. Pairs of closely related taxa (e.g. the large Barbatodon transylvanicus and the small B.oardaensis, gigantic and small sized teiid lizards and fossorial - semifossorial madtsoiid snakes) may have resulted as an adaptation to limited food supplies amid semiarid climate seasons and insular conditions. The existence of such pairs of sister taxa indicates a long-term persistence of the ”Hațeg Island” ecosystems extending to several million years. Most groups of this peculiar assemblage disappeared at the end of the latest Cretaceous, but a smaller part survived successfully the K/Pg extinction event (e.g. the alytid frogs and multituberculate mammals).

This research was supported by grant PN-II-PCE-2011-3-0381 of the Romanian National Council of Scientific Research - CNCS.

109 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

THE LATE JURASSIC MAMMAL ASSEMBLAGE FROM THE LANGENBERG QUARRY (LOWER SAXONY, GERMANY) O. Wings1*, T. Martin² and A.H. Schwermann²

1State Museum of Hannover, Hannover, Willy-Brandt-Allee 5, 30169 Hannover, Germany ²Steinmann-Institute, University of Bonn, Nussallee 8, 53115 Bonn, Germany

*[email protected]

Keywords: Kimmeridgian, Multituberculata, Dryolestida, Docodonta, Triconodonta

Despite its richness in Jurassic strata with a plethora of vertebrate fossils and more than 200 years of palaeontological research, no Jurassic mammals have been found in Central Europe until recently. During the 2014 fieldwork of the Europasaurus- Project, the first isolated mammal teeth were discovered at the Langenberg Quarry near Goslar in northern Germany.

This quarry is a classic outcrop of late Oxfordian to late Kimmeridgian marine strata which are palaeogeographically located in the Lower Saxony Basin. In the Jurassic, the area was surrounded by palaeo-islands which provided ample terrestrial habitats. Remains of terrestrial vertebrates and plants were embedded in the shallow marine basin. The Langenberg Quarry is best known as the type locality of the dwarf basal macronarian sauropod dinosaur Europasaurus, but also yielded numerous other terrestrial taxa.

Among the microvertebrate remains recovered by screen-washing at the Langenberg is a variety of mammal teeth. Multituberculata are represented by several isolated molariform teeth. One molar represents a new taxon with affinities to Proalbionbaatar from contemporaneous deposits of the Guimarota mine (Portugal). Docodonts as typical elements of Jurassic mammal faunas are represented by indeterminate fragments of molars. They occur regularly, at low diversity, in the Jurassic of Western Europe and North America and are much more diverse in Central and East Asia. Dryolestids as stem therians are a dominant element of Jurassic mammalian faunas of the western hemisphere but are very rare in Asia. The three molars that have been found at the Langenberg show a close relationship with dryolestids from the English Purbeck. The Langenberg docodonts and dryolestids are the easternmost European representatives of these groups. Triconodonta are represented by a eutriconodont molariform with a distinct cingulum. Unlike in the more basal amphilestids, all three main cusps of the Langenberg eutriconodont are of almost the same height.

The Langenberg findings contribute to our understanding of Jurassic European insular ecosystems and close a significant gap in our knowledge of early mammal evolution. Particularly noteworthy is the great diversity of mammal remains in this marine lagerstätte: despite the rather limited amount of screen-washed material, four major Jurassic mammalian groups have been recorded.

110 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday plenary talks

DIVERSITY OF MEXICAN CERATOPSIDS H.E. Rivera-Sylva1*, E. Frey2, R. Guzmán-Gutiérrez3 and P. Dodson4,5

1Departamento de Paleontología, Museo del Desierto, Carlos Abedrop Dávila 3745, 25015, Saltillo, Coahuila, México. 2Staatliches Museum für Naturkunde Karlsruhe, Geowissenschaftliche Abteilung, Erbprinzenstrasse 13, 76133 Karlsruhe, Germany. 3Dirección del Medio Ambiente, Municipio de El Llano, Aguascalientes, Ags., México. 4Department of Earth and Environmental Science, University of Pennsylvania, PA, 19104, USA, 5School of Veterinary Medicine, University of Pennsylvania, PA, 19104, USA.

*[email protected]

Keywords: Ceratopsidae, centrosaurine, chasmosaurine, diversity, Mexico

In the past decade, numerous new taxa of ceratopsian ornithischians have been described from Mexico. Apparently, ceratopsians experienced a regional diversification in this area. Here, we provide a critical review on Mexican ceratopsians and present a new centrosaurine ceratopsid from the Aguja Formation at the municipality of Ocampo, Coahuila. Because the squamosal of this taxon is the only bone with diagnostic features, we restrict the comparison with the other taxa to that bone. We also discuss possible reasons for the rapid specification of Mexican ceratopsians. To date Ceratopsia are represented by three species: (1) a new centrosaurine ceratopsian from strata of the Aguja Formation at Ocampo, Coahuila; (2) Coahuilaceratops magnacuerna, represented by material from the Cerro del Pueblo Formation from Coahuila; and (3) Agujaceratops mariscalensis, represented by an isolated and incomplete squamosal from the Aguja Formation of Coahuila. The two latter taxa belong to Chasmosaurinae. Biogeographically, ceratopsids appear to have had small species ranges, with north-south endemisms within the Western Interior Basin during the Late Campanian. The diversity of endemic Mexican ceratopsids suggests a hitherto unknown barrier that separated the Mexican area from North Laramidia during the Late Cretaceous and prevented a faunal interchange. Although the available material is currently limited, there is the potential to discover more and better preserved specimens in the near future. This will enhance our knowledge on the radiation in dependence of the palaeobiogeographic situation of ceratopsians that inhabited Laramidia during the Campanian.

111 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

THE DUTCH TYRANNOSAURUS REX SPECIMEN AND ITS EXTRAORDINARY PRESERVATION P. Kaskes1*, A.S. Schulp2,1, P.L. Larson3, J. Smit1, K.F. Kuiper1, H.A. Abels4, M.A. Prins1 and C.J. Beets1

1Vrije Universiteit Amsterdam, Department of Earth Sciences, De Boelelaan 1085, 1081 HV Amsterdam, the Netherlands 2Naturalis Biodiversity Center, Darwinweg 2, 2333 CR Leiden, the Netherlands 3Black Hills Institute of Geological Research, 117 Main St., SD 57745 Hill City, USA 4Delft University of Technology, Geoscience and Engineering Department, Stevinweg 1, 2628 CN Delft, the Netherlands

* [email protected]

Keywords: Tyrannosaurus rex, Theropoda, Hell Creek Formation, Maastrichtian, taphonomy

In September 2013, Naturalis Biodiversity Center (Leiden, the Netherlands) and the Black Hills Institute (Hill City, USA) excavated a new specimen of the theropod dinosaur Tyrannosaurus rex. The specimen comes from a fluvial sandstone deposit of the Upper Maastrichtian Hell Creek Formation, southwest of Jordan, eastern Montana, USA. It is a relatively complete and exceptionally well-preserved individual containing an articulated skull, left scapula and coracoid, furcula, much of the axial skeleton, an extraordinarily complete cage, and right leg. Provisional bone count indicates that well over 50% of the skeleton has been recovered, which grants this discovery a position in the top five of most complete Tyrannosaurus rex specimens ever found. The robust morphotype suggests the skeleton represents a female individual. The specimen displays multiple healed bone pathologies. Preliminary histological analysis suggests a very mature animal.

In September 2014, geological fieldwork was conducted by the Vrije Universiteit (Amsterdam, the Netherlands) and Naturalis to study its taphonomy and sedimentary context. The skull was found at the deepest part of the section whereas the pelvis and leg were located higher in the sequence and 10 to 8 m away from the skull, respectively. Rip-up clasts and climbing ripples observed at the excavation site point towards high palaeocurrent strengths. Additional grain-size and thermogravimetric analysis show that the unique three-dimensional preservation of this Tyrannosaurus rex is linked to a rapid burial under a 3.20 m thick blanket of fine, poorly consolidated, well sorted sand with a high carbonate content. We hypothesize that a single, massive flood event buried this carnivorous dinosaur shortly after its death. Furthermore, the sandstone was not compacted, thereby preventing bone deformation and maintaining the original three-dimensional morphology. Lastly, the high carbonate content, represented by angular, detrital dolomite grains, protected the skeleton against subsequent leaching. These characteristics of the sand body entombing the Tyrannosaurus rex provided the perfect recipe for its extraordinary preservation.

From September 2016 onwards, the specimen, currently registered in Leiden as RGM 792.000, will feature prominently in a new dinosaur exhibition of the Naturalis Biodiversity Center.

112 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

DINOSAUR TRACKS FROM A CRETACEOUS ALLUVIAL PLAIN (AROIFILLA FORMATION, BOLIVIA) Ch. A. Meyer1*, B. Thüring1, S. Thüring2 and D. Marty1

1Naturhistorisches Museum Basel, Switzerland, 2Naturmuseum Solothurn, Switzerland

*[email protected]

Keywords: Dinosaur tracks, Cretaceous, Bolivia, Alluvial plain deposit

For a couple of years Bolivia has been known to yield some of the largest dinosaur tracksites in the world. Apart from the mostly unstudied Toro Toro site (Toro Toro Formation; ?Coniacian – Santonian), the Cal Orck’O site (El Molino Formation; Maastrichtian) is the most famous locality in the Central Altiplano of Bolivia.

We report here on a small site from the Aroifilla Formation (Coniacian) close to Potolo in the Central Andes. The surface was accidentally uncovered by local people while blasting rocks for construction. Because the site will be covered by an irrigation dam in the future the local authorities invited us to document the surface before the finaldestruction.

The track bearing surface is steeply inclined and approximately 30 m2 are accessible without danger. 58 tridactyl footprints and 11 oval shaped footprints have been observed. The tridactyls show a strongly inward curved middle digit (mean FL 20 cm) and are similar to tracks reported from the Humaca site (Chaunaca Formation; Santonian - Campanian). The oval shaped footprints have been left by sauropods, but due to their preservation as underprints, no details can be seen. The trackbed contains invertebrate burrows that can be assigned to the ichnotaxon Steinichnus. These negative epichnia are the product of cricket mole larvae in the uppermost part of the substrate.

The section consists of medium- to fine-grained reddish sandstones with trough cross bedding. Rippled sandstones are thin-bedded (cm- to dm-scale), coarse to fine-grained, and contain abundant current and wave ripple marks, small-scale cross stratification, mudcracks, and mudstone rip-up clasts. These features point to a deposition on an alluvial plain of a shallow braided river system. Desiccation fractures indicate paleosol development in an arid climate.

113 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

THE FIRST UNEQUIVOCAL EVIDENCE OF GREGARIOUS BEHAVIOUR IN BABY SAUROPODS (LATE JURASSIC, NW SWITZERLAND) M. Belvedere1*, D. Marty1, K. Stevens2, S. Ernst1, N. L. Razzolini3, G. Paratte1, M. Cattin1, C. Lovis1 and C.A. Meyer4

1Office de la Culture, Paléontologie A16, Hôtel des Halles, P.O. Box 64, CH-2900 Porrentruy 2, Switzerland 2Department of Computer and Information Science, University of Oregon, OR 97403, USA 3Institut Català de Paleontologia Miquel Crusafont (ICP), Mesozoic Research Group, Carrer Escola Industrial, 23, 08201 Sabadell, Spain 4Naturhistorisches Museum Basel, Augustinergasse 2, 4001 Basel, Switzerland

*[email protected]

Keywords: vertebrate ichnology, baby sauropod, gregarism, Kimmeridgian, Switzerland

Excavations along Highway A16 (NW Switzerland) in Kimmeridgian tidal-flat deposits of the Jura carbonate platform revealed more than 650 sauropod and tridactyl (mostly theropod) trackways, from very large (PL>1 m for sauropod, >50 cm for theropods) to tiny footprints (PL<10 cm for both sauropod and tridactyl trackmakers). Since the small tracks may have been left either by small adults or by young individuals of a larger species, ‘baby’ dinosaur tracks are not easy to identify as such in the fossil record. We present a group of 8 subparallel tiny sauropod trackways, of equal preservation, in very close association to each other, suggesting that the animals were moving together. The tracks are very small (mean PL: 11.6 cm; PW: 7.8 cm; ML: 4.4 cm, MW: 7.6 cm; small heteropody) and well-preserved (dI-IV, dI-II claws, manus ungual). A slightly larger trackway (PL: 22.9 cm) is in close vicinity to the smaller ones on the same level. Additionally, 17 small sauropod trackways (PL between 30.5-46.1 cm) have a similar configuration and track morphology (digits and claws of pes, manus ungual), but proportionally shorter digits. These slightly larger trackways are subparallel to the group of tiny trackways, and all but four proceed in the opposite direction from the tiny trackways. We attribute the tiny tracks to very young sauropods of the same species as had left the small tracks, based on their morphological similarities (where the observed proportional differences of the digits compared to overall track length reflect allometric growth). The presence of very large sauropod (PL>1 m) and huge theropod (PL>50 cm) tracks on a level only about 10 cm higher supports our conclusion that the very small tracks were created by juveniles. Because most of the larger sauropod trackways are not heading in the same direction, at a first glance this ichnoassemblage of tiny tracks may provide evidence for separated gregarious behaviour, although the question of if and how ‘baby’ sauropods were protected against large predators in an open tidal-flat environment should not be disregarded.

114 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

UNIQUE EVIDENCE FOR A BIMODAL ORIENTATION PATTERN IN MEGALOSAURID (THEROPODA: DINOSAURIA) TRACKWAYS ON A MIDDLE JURASSIC TIDAL FLAT N. L. Razzolini1*, O. Oms2, D. Castanera3, B.Vila1,4, V. F. dos Santos5 and A. Galobart1,4

1Mesozoic Research Group, Institut Català de Paleontologia 'Miquel Crusafont' (ICP), C/ Escola Industrial 23, E-08201 Sabadell, Catalonia, Spain 2Universitat Autonoma de Barcelona, Facultat de Ciències (Geologia), 08193, Bellaterra (Spain) 3Bayerische Staatssammlung für Paläontologie und Geologie and GeoBioCenter, Ludwig- Maximilians-Universität, Richard-Wagner-Str. 10, 80333 Munich, Germany. 4Museu de la Conca Dellà, carrer del Museu, 4, 25650 Isona, Lleida. 5Museu Nacional de História Natural e da Ciência – Universidade de Lisboa, Rua da Escola Politécnica, 58, 1250−102 Lisboa, Portugal

*[email protected]

Keywords: Bathonian, megalosaurid tracks, tidal flat, bimodal arrangement, paleoethology

A new tracksite in the Vale de Meios quarry (Serra de Aire Formation, Bathonian), at the Lusitanian Basin of Portugal, preserves more than 700 theropod tracks (track length range 35 cm-80 cm) organized in at least 80 long unidirectional trackways. The Vale de Meios tracks are among the largest theropod tracks ever reported and have been assigned to Megalosauripus isp. They were produced by large individuals of the Megalosauridae family, the dominant tetanuran clade for this age in Europe as shown by the osteological record from Europe. The tracksite is preserved in an inter- tidal flat located at the edge of a lagoon. Directional orientation pattern analyses of the trackways, revealed that various individuals crossed the tidal flat most likely in accordance to tide cycles, and directing toward the barrier during low tide periods. Such clear bimodal orientation arrangement (forth and back), interpreted as single or small aggregates of large theropods individually moving toward on the shoreline, is highly uncommon as it is the high concentration of large theropod footprints. It is indeed a key and unique reference to understand the composition of the Middle Jurassic theropod fauna, being both the ichnological and osteological record for this age extremely scattered. Furthermore, it provides paleoethological observations on the basis of the unique trackway orientation and paleoenvironmental reconstruction of the tracking surface.

115 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

THE TETRAPOD TRACK RECORD FROM THE HUÉRTELES FM.: AN EXCEPTIONAL WINDOW INTO THE LOWER CRETACEOUS ICHNODIVERSITY OF EUROPE D. Castanera1* and J. I. Canudo2

1Bayerische Staatssammlung für Paläontologie und Geologie and GeoBioCenter, Ludwig- Maximilians-Universität, Richard-Wagner-Str. 10, 80333 Munich, Germany; 2Aragosaurus–IUCA, Paleontología, Facultad de Ciencias, Universidad de Zaragoza, 50009, Zaragoza, Spain.

*[email protected]; [email protected]

Keywords: Berriasian, dinosaurs, pterosaurs, crocodylomorphs, turtles

The Huérteles formation, located in the Cameros basin (North Spain), is well-known in the scientific ichnological literature due to the abundance of Berriasian tetrapod tracksites. The Huérteles formation was deposited in a tide-influenced fluvial-deltaic setting and has yielded and outstanding sample of mainly dinosaur but also other tetrapod tracks such as pterosaur, crocodylomorph, and turtle tracks. The main groups of dinosaurs (theropods, ornithopods, sauropods and stegosaurs) have been identified. Theropod tracks are the most abundant and at least two different ichnotaxa have been differentiated (Kalohipus bretunensis and Iberosauripus isp). The presence and validity of other ichnotaxa (Megalosauripus isp. and Archaeornithipus meijidei) is still debated. Ornithopods (Iguanodontipus burreyi), sauropods (Parabrontopodus-like and Brontopodus-like) and stegosaurs (Deltapodus isp.) complete the dinosaur track record. Besides, pterosaur tracks are quite abundant with many tracksites showing manus-only tracks although complete trackways with manus and pes sequences have been identified. Thus, several ichnospecies assigned to Pteraichnus have been described. Pteraichnus palacieisaenzi is the largest (footprint length about 15 cm) pterosaur ichnotaxa. Other 6 small sized ichnospecies have been described although further work is needed in order to understand the validity of all of them considering the possibility that some of them could be extramorphological or ontological variations of the others. Crocodylomorphs tracks (Crocodylopodus meijidei and cf. Crocodylopodus) and turtle tracks (Emydhipus cameroi and other unamed traces) have been also described. When compared with other European formations from the Tithonian- Berriasian interval (e.g: Purbeck Limestone Group, Bückeberg Fm., Villar del Arzobispo Fm.) the tetrapod track record from the Huérteles Fm. shows the highest ichnodiversity of all of them. Some of the described ichnotaxa from the Huérteles Fm. are shared with the different formations (e.g: Iguanodontipus burreyi, Deltapodus or Iberosauripus) suggesting some similarities between the ichnoassemblages.

116 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

SAUROPOD BRAINCASE ANATOMY AS A PRODUCT OF RAPID MORPHOLOGICAL TRANSFORMATION M. Bronzati1,2*, R. Benson3 and O. Rauhut1,2

1Bayerische Staatssammlung für Paläontologie und Geologie, Richard-Wagner-Str. 10, 80333, Munich, Germany. 2Ludwig-Maximilians-Universität München, Munich, Germany. 3Department of Earth Sciences, University of Oxford, OX1 3AN, Oxford, UK.

*[email protected]

Keywords: Sauropodomorpha, Claddis, Paleobiology

The body plan of sauropod dinosaurs, with an elongated neck and gigantic body size, differs substantially from the small, bipedal, gracile early sauropodomorphs of the Carnian (Late Triassic). This highlights the astonishing morphological disparity of sauropodomorphs. Recent studies and fossil discoveries have shown that features of the morphology of the vertebrae, limbs, girdles, and dentition, which were previously assumed as unique for sauropods, appeared in a mosaic fashion in the Sauropodomorpha lineage, among non-sauropodan sauropodomorphs (“prosauropods”). We investigated the timing of evolutionary modifications in the skull roof and braincase anatomy of Sauropodomorpha, prompted by the reanalysis of a Middle Jurassic (Bathonian) sauropodomorph braincase from England. This specimen is tentatively referred to the genus Cetiosaurus, and our phylogenetic analysis confirmed its sauropod affinities based on the presence of sauropod synapomorphies (e.g. reduction of the temporal region of the skull, basisphenoid shorter than basioccipital, lack of the vagal foramen). We examined rates of character state evolution using our revision and expansion of sauropodomorph braincase characters used in phylogenetic analyses. Principal coordinates analyses implemented in the R package Claddis indicate that sauropod braincases occupy a distinct region of morphospace than “prosauropods” and dinosaur/archosaur outgroups. Furthermore, statistical comparison of evolutionary rates indicates that these major anatomical modifications occurred in a short time interval, indicating rapid morphological transformation. The braincase houses soft tissues such as the brain, cranial nerves, and inner ear, important for the palaeobiology of the animal. Some studies have pointed out a direct correlation between changes in soft tissues anatomy and the assumed ecology of the animals. In the case of sauropods, a reduced flocculus has been previously linked to the quadrupedal stance. However, the most likely scenario is that a reduced flocculus indicates a less refined control of head and eye movement, which can be linked to the adoption of a fully herbivore diet. Also, sauropods have shorter vestibule in comparison to theropods, indicating differences in hearing sensitivity. In this context, our results favour the hypotheses of drastic changes concerning the paleobiology of sauropods in relation to their “prosauropod” relatives.

117 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

A “LOOK BACK” IN PALAEONTOLOGY: RECONSTRUCTING THE TAILS OF NEOSAUROPOD DINOSAURS V. Díez Díaz1*, H. Mallison1 and D. Schwarz1

1Museum für Naturkunde - Leibniz Institute for Evolution and Biodiversity Science, Invalidenstrasse 43, 10115 Berlin, Germany

*[email protected]

Keywords: Sauropoda, tail reconstruction, biomechanics, 3D model

Biomechanical studies of sauropods are often centered on the mobility of the postcranial skeleton, like the locomotor capabilities of the limbs, or the flexibility and range of motion of the axial skeleton. Neck posture and stabilization, and their association with feeding range, have been studied in Sauropoda for a long time, developing and using new techniques and methods. In contrast, the biomechanics of sauropod tails have been given little consideration, although the variations in length and bone shapes suggest variation in the biomechanical and behavioural functions, potentially with significant influence on our understanding of sauropod biology.

Using CAD (computer-aided design) analyses we are developing a research project – founded by the Alexander von Humboldt-Foundation – to study the biomechanics of the tail of several neosauropod taxa within Macronaria and Diplodocoidea. We are assessing the posture and range of motion thanks to virtual 3D skeletal mounts realized in McNeel Associates Rhinoceros 5.0. NURBS Modeling. The muscle moment arms and lines of action during motion are being assessed in Musculographics SIMM 7.0. Simplified 3D volumetric models based on the 3D muscle volumes from the Rhinoceros models will be imported into MSC ADAMS 2010 for multibody dynamic modelling to assess inertia moments, as well as gravitational effects on tail joints.

One of the principal aims of this project is to establish a biomechanical framework for tail variability in different sauropod groups, and link it to biological and ecological aspects of these animals, as well as explain why the tails developed these types of morphologies (i.e. “whip-lash” or clubbed). This project will greatly improve the comprehension of the tail anatomy of Sauropoda, and in general terms to the full biological and biomechanical knowledge of the body of these giant dinosaurs.

118 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

PRELIMINARY RESULTS ON THE BONE HISTOLOGY OF HADROSAURS FROM THE LATEST CRETACEOUS OF FAR EASTERN RUSSIA. K.H.W. Stein1,2, Y. Bolotski3, I. Bolotski3, P. Claeys1 and P. Godefroit2

1Earth System Science – AMGC, Vrije Universiteit Brussel, Pleinlaan 2, 1050 Brussels, Belgium 2Royal Belgian Institute of Natural Sciences, Directorate 'Earth and History of Life', Rue Vautier, 29, 1000 Brussels, Belgium 3Russian Academy of Sciences, Far Eastern Branch, Blagoveschensk, Russian Federation

*[email protected]

Keywords: hadrosaur, Maastrichtian, bone histology, Far Eastern Russia

Hadrosaurs such as Olorotitan and Edmontosaurus, were among the most abundant and successful Cretaceous dinosaurs. They spread to all continents and some reached body masses rivalling those of the giant sauropods. However, some hadrosaur groups disappeared from the North American continent 5 to 6 million years before the extinction event, whereas the same groups thrived in Asia until the event. The Maastrichtian hadrosaur communities from Far Eastern Russia are a prime example to study the biology of late surviving lambeosaurine hadrosaurs. The bonebeds from which these taxa were excavated show a high species diversity and a high abundance of various skeletal elements, most notably long bones. We sampled long bones from different size classes of the lambeosaurines Olorotitan and Amurosaurus. Both taxa show highly vascularized cortical bone consisting mainly of a woven-parallel fibered complex with plexiform to laminar vascularization, indicative of high growth rates. Only few lines of arrested growth (LAG) could be observed in even the biggest specimens (femur length ~1 m). It remains unclear if this indicates an uninterrupted juvenile growth phase, or if mass increase was extremely high in the first years, but the high average growth rate is in accordance with high growth rates reported for North American hadrosaurs (e.g. Maiasaura). We also sampled the tibia of a large tyrannosaurid theropod (corresponding femur length ~ 0.92 m) which was recovered from the same layers as Olorotitan, however, its growth is in sharp contrast with that of the hadrosaurs. The thin section taken from the shaft of the proximal tibia shows a woven-parallel fibred complex with plexiform to laminar vascularization, however, it also shows numerous (>15) LAGs, indicating a much slower average growth and higher chance of survivorship than for hadrosaurs. These data add to a more global perspective on diversity and ecology of Latest Cretaceous dinosaur faunas.

119 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

NEW DISCOVERIES OF RHABDODON FROM THE CAMPANIAN OF PROVENCE (SE FRANCE) PART II: A BETTER UNDERSTANDING OF THE INTRASPECIFIC DIVERSITY T. Tortosa1* and Y. Dutour2

1Réserve Naturelle de Sainte-Victoire (RNSV), Conseil Départemental des Bouches- du-Rhône, Hôtel du Département, Direction de l’Environnement, 52 avenue Saint Just, 13256 Marseille, France. 2Muséum d'Histoire naturelle d'Aix-en-Provence (MHNA), Parc Saint-Mitre, 166 Avenue Jean Monnet, 13090 Aix-en-Provence, France.

*[email protected]

Keywords: Dinosaur, Rhabdodon, Upper Campanian, Provence, Dimorphism

The present study is the continuation of previous work on the ornithopod dinosaur Rhabdodon from Provence, presented at the 13th Annual Meeting of EAVP (Opole, Poland). Here we note the discovery of new partial skeletons which confirm the evidence of the high specific diversity of this genus and new information on intraspecific dimorphism.

At least four fragmentary skeletons of small specimens have been excavated by the RNSV team, jointly with the MHNA, in the historical locality of Grands-Creux, near Roques-Hautes, a famous place for the unearthing of the hundreds of dinosaur eggs. These discoveries provide additional unknown elements of the anatomy of Rhabdodon. They include cervical, dorsal and caudal vertebrae series, a forelimb (with a partial hand), the pelvis and a complete leg (with a pes). The most striking features is the difference of morphology observed on three ischia with the same size and coming from the same horizon. These discoveries provide a new light concerning the sexual dimorphism of Rhabdodon. This observation allows a reinterpretation of previous discoveries from Vitrolles-Couperigne locality (Pincemaille-Quillevéré, 2002; and unpublished works).

Based on the characteristics of the pelvis and the femur, the specimens from Grands-Creux seem to be closer of the isolated elements found on the historical locality of Fox-Amphoux (Var). Unfortunately, the material can not be directly attributed to Rhabdodon priscus from the La Nerthe Tunnel because of the lack of common elements.

These new discoveries confirm the complex combination of features between specimens from Provence: La Nerthe Tunnel (Bouches-du-Rhône), Trets (Bouches- du-Rhône), Vitrolles-Couperigne (Bouches-du-Rhône), Fox-Amphoux (Var), Pourrières (Var), the Saint-Chinian area (Hérault); and establish the basis for the recognizing of the different types of dimorphism (interspecific, ontogenetic and, now, sexual) in order to better define the genus Rhabdodon and better estimate the European palaeobiodiversity in the Ibero-Armorican island during the Late Cretaceous.

120 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

BIOMECHANICAL TRAITS OF CARNIVOROUS JAWS WITH FINITE ELEMENT METHODS J. Marcé-Nogué1* , S. De Esteban-Trivigno2,3, C. Escrig4, L. Gil4 and T.M. Kaiser1

1Centrum für Naturkunde, University of Hamburg, Martin-Luther-King-Platz 3, 20146 Hamburg, Germany. 2Institut Català de Paleontologia, Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Spain 3Transmitting Science, Piera, Spain 4Universitat Politècnica de Catalunya, c/colom 11, 08222, Terrassa, Spain

*[email protected]

Keywords: biomechanics, carnivorous, Finite Element Analysis, lower jaw

In recent years important advances have been accomplished in comparative functional analysis of vertebrate structures. That involved the introduction of virtual models that allowed testing of functional hypotheses using Finite Element Analysis (FEA) and its combination with Geometric Morphometrics (GMM) and mathematical, statistical and engineering approaches.

Classical biomechanics were used to estimate muscular and biting forces as a load response in the models using dimensionless parameters whereas FEA generated stress distribution patterns allowing a comparative approach of chewing biomechanics under certain loads. Herein, FEA results were compared quantitatively from different models of extant and extinct carnivorous jaws in order to establish a robust methodology in a comparative context. Recommendations are stated to be taken into account when building the models (scaling methods to remove the effect of the size, material properties, orientation, etc.) as well as pathways of comparison in obtaining isolated points or whole model.

Plane models of extant and extinct carnivorous including dietary traits such as insectivory, herbivory, omnivory, carnivory were established using ANSYS FEA Package v.16 for Windows 7 (64-bit system) in order to obtain the von Mises stress distribution of the planar models and compare them. Recent procedures to compare quantitative FEA results from different models were used combining the Stress distribution maps and the values for the whole model. That includes exploratory methods such as box-plots and Principal Component Analysis. Phylogenetic signal was tested by means of Bloomberg’s K. Patterns of stress and load response can thus be understood as adaptive to biomechanical diet response.

The results obtained suggest that the proposed methodologies provide powerful indicators that are suitable to apply in comparative approaches as well as for pattern detection as a proxy system of niche specificities. The new methods proposed are shown to be extremely useful when exploring the effect shape has on strength and stiffness of bone structures, such as the lower jaws of the carnivorous employed as a model here. This is especially for extinct species, which these methodologies can help to understand better its feeding behaviour.

121 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

EVOLUTION OF JAW SHAPE AND BIOMECHANICS IN CARNIVORES: A PHYLOGENETIC APPROACH S. De Esteban-Trivigno1*, J. Marcé-Nogué2, A. Boscaini3 and J. Cantalapiedra4

1Institut Català de Paleontologia M.C., Z Building, c/ de les Columnes, s/n. Campus de la UAB. 08193 Cerdanyola del Vallès, Barcelona, Spain. 2Centrüm für Naturkunde University of Hamburg, Martin-Luther-King-Platz 3, 20146 Hamburg, Germany. 3Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales, Av. Ruiz Leal s/n Parque General San Martín, 5500 Mendoza, Argentina. 4Museum für Naturkunde, Leibniz-Institut für Evolutions und Biodiversitätsforschung, Invalidenstraße 43, 10115 Berlin, Deutschland.

*[email protected]

Keywords: Carnivora, Finite Element Analysis, Geometric Morphometrics, Macroevolution, Phylogenetic Signal

Carnivorans are a highly diversified group that shows adaptations to many different ecological niches. Despite the relationship between jaw shape and biomechanics has been thoroughly studied in the past, no study has analysed both characteristics in an evolutionary framework using Finite Elements Analysis (FEA) together with Geometric Morphometrics.

In this work we explored the relationship between the mandibular shape (analysed by Geometric Morphometrics methods) and biomechanics (quantified with Finite Element Analysis) of almost 30 species belonging to 10 families of Carnivora in an evolutionarycontext. For FEA a novel approach was used, generating 100 variables representing equal ranges of stress values. Two cases were analysed, the first one analysing the stress values when the force was applied in the canine (Set 1), and the second one the force simulated in a more posterior place of the tooth row (Set 2).

A Principal Component Analysis (PCA) of the shape data showed low integration and many directions of change, with only PC2 partially related with diet. Shape data had phylogenetic signal. On the contrary, most FEA variables were not related to the phylogeny. There was a certain degree of covariation between shape and stress levels. For Set 1, this covariation turns out to be non-significant when phylogeny is taken into account. However, for Set 2 this relationship, although smaller, is maintained when considering the phylogeny.

Taking all the evidence into account, our results suggested that phylogeny played an important role in carnivoran jaw shape evolution. On the other hand, biomechanical variables were more correlated with diet and more independent of phylogeny. As FEA is affected by shape, the conclusion is that not all the shape variability was related with function, and other factors are probably involved. Combining information from both sets shed light into specific adaptations and point out the importance of using different biomechanical hypothesis to biologically analyse FE results.

Our results are preliminary and describe broad scale evolutionary patterns, as just a small fraction of Carnivora is represented here. Adding more species to the analysis will prove whether the observed patterns apply at smaller scales (e.g. within tribes and genera). 122 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

PALEONTOLOGICAL MASS 3D DIGITIZING: LEARNING BY DOING THE MFN’S DIGIS 2015 PROJECT ‘LIBERATION FROM THE BONE CELLAR’ M. Belvedere1,2*^, H. Mallison1^ and B. Schurian1^

1Museum für Naturkunde Berlin - Leibniz Institute for Evolution and Biodiversity Science, Invalidenstrasse 43, 10115, Berlin, Germany 2Office de la Culture, Paléontologie A16 Hôtel des Halles, P.O. Box 64, CH-2900 Porrentruy 2, Switzerland ^ shared first authors

*[email protected]

Keywords: digitization, photogrammetry, collection management

Collecting digital data is becoming increasingly important both for research and collection management aspects. Many methods exist, delivering models of different qualities and with differing properties: X-ray based methods (CT, microCT) deliver outside shape and internal structure. Surface scanning (photogrammetry, laser scanning) delivers high resolution shape and colour data. Resulting files have been used in research, but mass digitizing of entire collections without a direct, immediate research use is still rarely performed. In 2015, the State of Berlin, via its digiS program, funded the 3D-digitizing of the large Tendaguru dinosaur bones in the MfN Berlin’s collection.

Our project was a deliberate attempt at speedy low-cost mass digitizing of cumbersomely large and heavy specimens under suboptimal work conditions within the Bone Cellar itself, because it is rarely possible to transport large numbers of heavy specimens to a photography lab. Therefore, we used photogrammetry with a hand-held Canon 70D DSLRs equipped with 10-18 mm wide-angle lenses and LED ring lights. With this equipment we were able to take photographs at close distance with even, shadow-free light, but also overview images showing entire bones and large (0.5 m) scale bars.

In order to support the bones, we constructed small tables matching bone size, which allowed freedom of movement despite the narrow corridors of the Bone Cellar, and use cut-to-fit pieces of Ethafoam as support. In order to allow model calculation in one chunk per bone, two tables with differently coloured support material and background were used to shoot two sets of images of each specimen. For bones too large to be transported between tables, we covered the background to alter it completely between photo sets.

We digitized ~400 bones, with a total of about 100.000 photos taken (~ 650 GB). The average error of the models generally was lower than 0.2 mm, respective to a 0.5 m reference scale bar, i.e. < 0.4%. We also accomplished a low failure rate: less than 20 bones had to be reshot to obtain suitable models.

This success can serve as a model for other digitizing efforts, both in collections or during research visits.

123 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

USING BIOGEOGRAPHY AND THE FOSSIL RECORD TO STUDY CONVERGENT EVOLUTION IN VERTEBRATES J. W. Oyston1*, M. Hughes2, P. J. Wagner3 and M. A. Wills1

1Department of Biology and Biochemistry, University of Bath, Bath BA2 7AY, United Kingdom 2Department of Zoology, The Natural History Museum, London SW7 5BD, United Kingdom 3Department of Paleobiology, National Museum of Natural History, Smithsonian Insitution, Washington D.C. 20013-7012

*Correspondence Email: [email protected]

Keywords: Convergence, Phylogeny, Biogeography, Disparity, Morphospace

Convergent evolution is widespread across the tree of life, indicating that the evolution of even complex traits may be far more predictable that traditionally supposed. The fossil record of vertebrates provides an unparalleled resource with which to study convergent evolution, yielding many striking examples including multiple origins of powered flight, hydrodynamic bodyforms and the keratinised rostrum. High levels of convergence in the fossil record suggests the potential range of morphological form is shaped by developmental and ecological constraint. Many vertebrate clades reach maximum disparity early in their evolutionary history and the rate of evolution of new character states decreases up the phylogenies of most clades, consistent with a restricted morphospace and widespread convergence on the macroevolutionary scale. This has profound consequences for inferring evolutionary relationships using morphology. Indeed, molecular phylogenies with strong support have overturned our traditional understanding of relationships based on morphology numerous times, proving that morphological trees can frequently be strongly misled by convergent evolution. In order to have greater confidence that our views on the relationships of extinct vertebrates are not being obfuscated by convergent evolution, we need to test our phylogenies using other independent sources of data that may indicate relatedness. When dealing with wholly extinct groups where it is impossible to test evolutionary hypotheses using a molecular framework, we have two main sources of data with which to test our phylogenies. The first and more commonly used method uses fossil ranges to test how stratigraphically consistent a given topology is. However, many instances of convergence do not occur in isolation but in groups known as convergent radiations. In the case of mammals it would appear that while traditional relationships based on morphology have been strongly influenced by convergence, the robustly supported major clades recovered in molecular studies for the most part map onto endemic regions. If diversity is primarily the result of successive radiations into biogeographic regions, adapting to similar ecological niches and evolving a similar suite of traits each time then biogeographic data might prove valuable in identifying likely instances of convergence in many groups of fossil vertebrates.

124 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

MARINE TURTLES FROM THE LATE CRETACEOUS OF NE ITALY HIGHLIGHT THE MYSTERIOUS ORIGIN OF THE LEATHERBACK LINEAGE M. Rabi1*, M. Delfino1, A. Villa1 and G. Carnevale1

1University of Torino, Department of Earth Sciences, Via Valperga Caluso 35, 10125 Torino, Italy

*[email protected]

Keywords: , , Cretaceous, phylogeny

Leatherbacks constitute a bizarre clade of marine turtles today represented by a single species, Dermochelys coriacea. Besides being among the largest living reptiles, a series of peculiar features make it a particularly unique vertebrate. These include a greatly reduced, “embryonic” skeleton, the development of a secondary bony carapace, and highly elevated growth and metabolic rates reminiscent to that of marine mammals. Cold water tolerance enables D. coriacea to attain a cosmopolitan distribution and to lead a truly pelagic lifestyle with capability of extremely deep dives for their almost exclusive prey of jellyfishes. Due to their highly specific anatomy and lifestyle, however, the fossil record of leatherbacks is very poor. Even more frustrating is that the earliest representatives (Eocene) already show most of the peculiarities of the living leatherback and therefore provide few insights regarding the early evolution of the lineage.

The subject of the present research is the extinct Mid-Cretacous Protosphargis veronensis from the Scaglia Rossa limestone of Veneto Province, NE Italy which shows remarkable similarities in the morphology of the primary shell to leatherback turtles. New material suggest that its close relative was hoffmanni from the type-Maastrichtian; a species exhibiting further unique similarities in the skull and growth strategy with leatherbacks.

The two alternatives are that either P. veronensis together with A. hoffmanni are primitive representatives of the leatherback lineage or instead, we face an extreme case of parallel evolution. This issue is linked to another fundamental hiatus in our understanding of marine turtle evolution: it is yet to be clarified whether the extinct Cretaceous clade Protostegidae represents an entirely independent marine invasion from crown-sea turtles. Such an independent evolution would imply that Cenozoic hard-shelled sea turtles (stem + crown ) developed remarkable parallels with protostegids (following the Late Cretaceous extinction of the latter) and likewise did the clade of P. veronensis – A. hoffmanni with leatherbacks. However, our phylogenetic analysis only supports this scenario when the characters pertaining to marine adaptation (i.e. the potential subjects of parallel evolution) are omitted from the dataset. Consequently, the origin of leatherback turtles is more of a mystery than ever before.

125 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

VERTICAL NECK RETRACTION IN AN EARLY STEM SIDE- NECKED TURTLE H. Tong1*, J. Anquetin2,3 and J. Claude4

1Mahasarakham University, Palaeontological Research and Education Centre, Kantarawichai, Mahasarakham 44150, Thailand 2JURASSICA Museum, Route de Fontenais 21, 2900 Porrentruy, Switzerland 3University of Fribourg, Department of Geosciences, Chemin du Musée 6, 1700 Fribourg, Switzerland 4Université de Montpellier, Institut des Sciences de l’Evolution de Montpellier, UMR 5554 CNRS/UM2/IRD, 2, Place Eugène Bataillon, cc64, 34095 Montpellier Cedex 5, France

*[email protected]

Keywords: Testudines, stem Pleurodira, Platychelys oberndorferi, , neck retraction

For protective reasons, turtles developed complex cervical structures allowing neck and head retraction within the shell. Pleurodires (side-necked turtles) bend their neck sideways and tuck their head under the carapace anterior to the pectoral girdle. Cryptodires (hidden-necked turtles) bend their neck in a vertical plan and withdraw their neck and head within the shell between the shoulder girdles. Consequently, the morphology of the cervical vertebrae in these two groups is different. Platychelys oberndorferi is an early stem Pleurodiran turtle known from the Late Jurassic of Germany and Switzerland. The shell of Platychelys shows several pleurodiran characters notably the sutural contact between the pelvis and the shell. Two cervical vertebrae of P. oberndorferi from Solothurn, Switzerland, housed in the Natural History Museum of Basel (NMB So.596) are identified as the cervicals 6 and 8. The centrum of the cervical 6 is biconcave and that of the cervical 8 is biconvex. Ginglymoidy is present on the posterior central articulation of the cervical 6 and anterior articulation of the cervical 8. The wider than high articulation of the centrum, the ginglymoidy, the widely spaced pre and postzygapophyses and the weakly developed transverse processes which are anteriorly located are very different from the cervicals of pleurodiran turtles, but closely resemble those of cryptodires. This cervical morphology is incompatible with the sideways folding of the neck seen in pleurodires, but indicates a vertical bending of the neck as in cryptodires. However phylogenetic analyses, including the updated neck characters, continue to support the pleurodiran affinities of P. oberndorferi. Recent studies on the evolution of cervical vertebrae revealed that disparity of vertebral shape was high at the dawn of turtle evolution, and declined after the modern groups evolved, reflecting a stabilization of vertebral shape associated with the two modes of neck retraction. Platychelys is a good example of convergence in the neck evolution of turtles and reveals that a cryptodiran-like mode of neck retraction first evolved in Late Jurassic stem pleurodires.

126 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

MORPHOMETRIC AND POLYMORPHIC CHARACTERS AND THE PHYLOGENY OF GEOEMYDID TURTLES W.G. Joyce1*, E. Ascarrunz1 and R. Garbin1

1University of Fribourg, Department of Geoscience, Chemin de Musée 6, 1700 Fribourg, Switzerland

*[email protected]

Keywords: morphometrics, polymorphism, phylogenetic reconstruction, , Testudines

The shell of turtles is a complex morphological system consisting of numerous dermal bones that are overlain by a non-corresponding set of epidermal scutes. Given the great number of elements and the large amount of possible variation in their topographic arrangement, the turtle shell has proven to be a valuable source of character information for the identification of species and phylogenetic reconstruction. At the same time, however, the turtle shell shows enormous amounts of intra-specific variation and exhibits high levels of homoplasy. These difficulties are particularly prevalent among turtles of the clade Geoemydidae, one of the primary clades of (pond and terrestrial turtles). However, given that the fossil record of geoemydids mostly consists of shells, being able to extract phylogenetic informative characters from this structure is of the greatest imperative to understanding the evolution of the group.

We here utilize a two-branched approach in search of phylogenetically informative shell characters among geoemydids. For the first branch we collected three- dimensional data for 76 type I landmarks among >130 testudinoid carapaces using a microscribe. For the second branch we scored 72 discrete morphological characters for 307 specimens, of which 70 characters were observed to be polymorphic. Although sampling is not overlapping between the two approaches, both datasets sample over 70 percent of extant geoemydid diversity and select outgroups. The landmark data are continuous characters by nature, which we analyzed as such using clustering, parsimony, and likelihood methods. The discrete character dataset was analyzed using combinations of coding schemes and phylogenetic inference methods. The results from both branches were evaluated by comparison to an updated molecular phylogeny. Additionally, we used cross-validation to assess the precision of backbone-constrained analyses of the discrete character dataset. Although preliminary results confirm the difficult nature of shell characters for the clade, both approaches provide meaningful insights.

127 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

THE EVOLUTION OF CRANIAL DISPARITY IN FOSSIL AND RECENT TURTLES C. Foth1* and W.G. Joyce1

1University of Fribourg, Department of Geosciences

*[email protected]

Keywords: Turtles, Skull evolution, Geometric morphometrics, Disparity through time

Turtles (Testudinata) are a diverse group of reptiles with a rich fossil record that extends back to the Late Permian. While the origin and phylogenetic interrelationships of turtles have become clearer in recent years, macroevolutionary processes, including changes in morphological diversity (= disparity) through time, are still poorly understood. We here investigate the cranial disparity of turtles, by applying landmark-based, two-dimensional geometric morphometrics to 172 representatives of the turtle lineage and their hypothetical ancestors grouped into 20 time bins ranging from the Permian to the Recent. Cranial shapes were captured in three different anatomical views, superimposed, and subjected to principal component analyses. The principal components containing the most significant shape variation were identified via the broken-stick method and subsequently mapped onto a time-calibrated supertree to estimate ancestral skull shapes. After binning terminal and ancestral taxa into equal time periods, the sum of variance was calculated on the basis of the principal components for each bin. Disparity was found to be greatest in lateral view and smallest in ventral view. In all three anatomical views, three different phases can be detected. In the first phase, disparity increases gradually from the Permian to the Palaeogene with only a minor decline at the K/T boundary. Although global warming was previously suggested to cause an increase in diversity, and by extension in disparity, we find the fragmentation of Pangaea during the Mesozoic to be a more plausible factor for this development. After its maximum, disparity declines strongly towards the Miocene, only to recover partially towards the Recent. This collapse in disparity is probably a result of habitat destruction due to global drying, combined with the homogenization of global turtle faunas through increased transcontinental dispersal in the Tertiary. The disparity minimum in the Miocene, marking the beginning of the third phase, is probably an artifact of the methods employed herein. Interestingly, turtles show no significant shift of their position within morphospace through time. However, when comparing the two major lineages, disparity of pan-cryptodires is generally higher than that of pan- pleurodires, while both groups occupy significantly different areas in morphospace for each time bin.

128 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

REVISITING A CONTROVERSIAL FOSSIL BIRD: THE ALLEGED AUK PETRALCA IS A LOON U.B. Göhlich1* and G. Mayr2

1Natural History Museum Vienna, Department of Geology and Paleontology, Burgring 7, A- 1010 Vienna, Austria 2Forschungsinstitut Senckenberg, Ornithological Section, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany

*[email protected]

Keywords: Petralca austriaca, loon, Gaviiformes, Early Miocene, marine fossil birds

The fossil bird Petralca austriaca was described by Mlíkovsky in 1987 as a fossil species of auks (Alcidae), within a new subfamily Petralcinae. The holotype and only known specimen is a poorly preserved partial skeleton consisting of wing bones in two slabs. The fossil was discovered in marine Early Miocene deposits in the Austrian locality Pucking. P. austriaca was announced as the oldest record of Alcidae in the Cenozoic of Europe.

However, the assignment of P. austriaca to the Alcidae has been recently doubted and it was proposed that the species is instead a loon (Gaviidae). But taxonomic discussion was hampered by the rather superficial original description of P. austriaca, which was in part due to the incompleteness and poor preservation of the partial skeleton.

A recent re-preparation of the fossil yielded new data on the skeletal morphology of Petralca, which allow more detailed comparisons with auks and loons. We show that the taxon can be clearly distinguished from auks in various skeletal features and can be confidently identified as a loon.

Among gaviiformes, Petralca appears to be more closely related to the genus Gavia than to the fossil taxa Colymboides and Colymbiculus. Unusually thick bone walls of the limb bones indicate well-developed diving capabilities for Petralca austriaca.

129 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

SYNCHOTRON MICROTOMOGRAPHY REVEALS ADDITIONAL ELEMENTS IN THE NINTH SPECIMEN OF ARCHAEOPTERYX D.F.A.E. Voeten1,2*, M. Röper3, S. Bureš1 and P. Tafforeau2

1Palacký University, Faculty of Science, Department of Zoology and Laboratory of Ornithology, 17. listopadu 50, 771 46 Olomouc, Czech Republic 2European Synchrotron Radiation Facility, 71 Avenue des Martyrs, 38000 Grenoble, France 3Bürgermeister-Müller-Museum, Bahnhofstrasse 8, 91807 Solnhofen, Germany

*[email protected]

Keywords: Archaeopteryx, birds, Late Jurassic, tomography, synchrotron

The ninth or Bürgermeister-Müller specimen of Archaeopteryx was discovered and first described in 2004. It consists of a partially disarticulated right wing representing one of the larger individuals known to date. Contrary to other Archaeopteryx specimens, its skeletal elements generally experienced limited compressional deformation after burial and therefore conservatively retained their original three- dimensional morphology. We have subjected the main slab that preserves nearly all of the osseous material to propagation phase contrast X-ray synchrotron microtomography (PPC-SR µCT) on beamlines BM05 and ID19 at the ESRF in Grenoble in order to visualize morphological aspects of the wing skeleton that remain embedded in the limestone matrix. This approach revealed that the bones visible on the surface are preserved largely intact and led to the discovery of four additional elements thus far undescribed for this specimen. Distal to the radius, the radiale was resolved. It is a rounded tetrahedral element capped by a flat, anteriorly projecting triangular process. The somewhat reniform distal carpal III was recognized as well. It is preserved locked between the semilunate carpal (distal carpals I+II), metacarpal II and metacarpal III. The bulbous proximal portion of carpal III communicates with the posterodistal aspect of the semilunate carpal. Its anterolateral margin is in close contact with the posteroproximal edge of metacarpal II. The smaller, flattened distal part of carpal III borders the proximal end of metacarpal III. Postmortem hyperflexion and subsequent disarticulation rotated manual ungual I, which is slightly shorter yet more robust than manual ungual II, to cover the distal half of manual phalanx I. The corresponding keratinous claw became dislodged from the ungual. Nearly the complete right anterior limb skeleton is now accounted for, with notable exception of an ulnare that might not have been present in vivo or has been taphonomically separated from the association. Our study provides valuable new information on the three-dimensional geometry and interrelations of the anterior limb bones of Archaeopteryx, particularly those in the wrist.

130 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

THE PLIO-PLEISTOCENE MARINE MEGAFAUNA EXTINCTION C. Pimiento1,2*, J.N. Griffin3, C.F. Clements4, S. Varela5, M. Uhen6, J. Velez- Juarbe7 and C. Jaramillo2

1University of Zurich, Paleontological Institute and Museum, CH-8006, Switzerland. 2Smithsonian Tropical Research Institute, Box 2072, Balboa, Panama. 3Swansea University, Department of Biosciences, Wallace Building, Singleton Park, Swansea, SA2 8PP, UK. 4University of Zurich, Department of Evolutionary Biology and Environmental Studies, Zurich, CH-8057, Switzerland. 5Museum für Naturkunde. Leibniz Institute for Evolution and Biodiversity Science. Invalidenstrasse 43, 10115 Berlin, Germany. 6George Mason University, AOES Geology, MSN 6E2, Fairfax, VA 22030, U.S.A. 7Department of Mammalogy, Natural History Museum of Los Angeles County, Los Angeles, California 90007, U.S.A.

*[email protected]

Keywords: climate change, ecological shifts, functional diversity, habitat loss, vertebrates

The onset of the Pleistocene epoch around 2.5 million years ago was characterized by increased climatic variability and sea-level oscillations. Here, based on a new analysis of the fossil record, we identify this time as a previously unrecognized extinction pulse among marine megafauna. Over 30% of Pliocene species were extirpated, including representatives of mammals, seabirds, turtles, and sharks. Reconstructions showed that falling sea-level reduced continental shelf area by 29% during this period, identifying loss of productive coastal habitats as a probable extinction driver. Model analyses revealed signals of selectivity, with species vulnerability to extinction partially explained by traits of thermoregulation and guild. To gauge the ecosystem consequences of this event, we further evaluated its impact on functional diversity. Eight unique functional entities disappeared, along with 17% of functional space. Origination of new species, including the modern killer whale and polar bear, created new functional entities and contributed to a functional shift of 20%, but only partially compensated for the space lost, leaving modern assemblages functionally depauperate compared to their Pliocene analogues. The extinction event reported here was the last major extinction that marine megafauna experienced, and demonstrates that these species were more vulnerable to climatic changes in the recent geological past than previously thought, and that they are probably still in the recovery phase today.

131 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

SEXUAL DISPLAY AND ROSTRAL VARIATION IN EXTINCT BEAKED WHALE, GLOBICETUS HIBERUS J. Muchagata1, 2* and O. Mateus1, 3

1Faculdade de Ciências e Tecnologia/Universidade Nova de Lisboa, 2829-516 Caparica, Portugal; 2Departamento de Geociências, Universidade de Évora, Largo dos Colegiais 2, 7000 Évora, Portugal; 3Museu da Lourinhã, Rua João Luis de Moura, 95 2530-158 Lourinhã, Portugal

*[email protected]

Keywords: Ziphiid cetaceans, beaked-whales, sexual dimorphism, secondary sexual organ, echolocation

Iberian extinct ziphiid, Globicetus hiberus, bears a peculiar large bony sphere in the rostrum, the Mesorostral Process of the Premaxillae or MPP. The MPP varies in size and shape of growth in the six specimens studied and seems to have an allometrically growth in one subgroup, but not in the other, suggesting subgroups correspond to males and females (sexual dimorphism). Even more, some rostral structures, such as the medial pad of the premaxillae seem to be associated with the specimens with lower and leaner MPP’s and ossification of the mesorostral canal by the vomer can also be of value in differentiating sex. Beaked whales are deep-diving, echolocation-user odontocetes and able to perceive bones as distinctive echoic images with their sonar; therefore the MPP may work as a secondary sexual organ (“antlers inside” hypothesis by Gol´din, 2014), a mute display structure acting as an “acoustic flag” to be perceived through echolocation by other individuals, giving information about the shape and size of the MPP.

132 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

THE CONTRIBUTION OF PALAEONTOLOGY TO THE HISTORY OF THE ARABIAN CONTINENTAL PLATE E. Hoch1*

1Museum Sønderjylland, Gram Museum of Palaeontology, Lergravsvej 2, DK-6510 Gram, Denmark

*[email protected]

Keywords: Arabian palaeontology, fossil Sirenia, Qatar, Tethys, Ardea bennuides

Rifting of Afro-Arabia and opening of the Red Sea and Gulf of Aden from late in the Miocene split off the Arabian continental plate. Differences in terrains made the Romans divide Arabia into Arabia Petraea, Arabia Deserta and Arabia Felix (felix meaning fecund, fertile) delimited by Sinus Arabicus (the Red Sea), Indicum Mare and Sinus Persicus. Geology teaches thatnorthward drift of Gondwana narrowed the nearly circumterrestrial ocean Tethys; and during its drift the Arabian plate collided with Eurasia, and the northeastern margin of Arabia became faulted and folded up into the Zagros Mountains before subduction below Asia. Intraplate lateral compression probably contributed to forming the NW-SE trending syncline that contains the Tigris Euphrates plain and the Persian(/Arabian) Gulf. The Gulf is shallow (below 100 m, excepting the Hormuz Strait). Its large animals include Cetacea, Dugong dugon and Rhincodon typus (whalesharks). Fossils uncovered in dry western Abu Dhabi represent Crocodilia, Ardeidae, Deinotheriidae, Equidae, Hippopotamidae and many others found in fluvial sediments of the late Miocene Baynunah Formation. Most of the animals were of genuine AfroArabian provenance; Hipparion (Equidae) and others had close ancestors on the northern continents and on the Indian subcontinent. Their presence supports other evidence of a Near or Middle East 'Gomphotherium landbridge' that first severed Tethys. The Baynunah fauna and flora indicate lush and well-watered landscapes. Such conditions may have persisted into modern times interrupted by incursions of sea-water over low- lying lands. During the last Pleistocene glaciation the Gulf basin was a valley of rivers and lakes. It became flooded during the Holocene sea-level rise. The Qatar Arch, a basement high, have exposed Eocene limestones that are locally superimposed by the Dam Formation of early-middle Miocene calcareous / gypsiferous / silty deposits. Sirenian fossils are discovered in southwestern Qatar in middle/late Eocene or Dam Formation sediments. The area and a dugong population off Qatar's west coast are protected under Qatari law. Sirenia evolved in Palaeogene Tethys from amphibious quadrupeds related to elephants. The Qatar sirenian fossils reflect Arabian Tethys shelf conditions. The dugongs are 'newcomers' arrived with the Holocene flooding of the Gulf basin. A huge, extinct heron, Ardea bennuides Hoch 1979, recorded from sub-recent deposits on Umm an-Nar (Abu Dhabi) and Failaka (Kuwait) exemplifies the remarkable diversity in Gulf Ardeidae.

133 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

LEVIATHAN RISING: THE BLEEDING EDGE OF RESEARCH CHASED OUT OF THE DEPTHS BY THE SHARK OF PUBLIC SECTOR DEADLINES J. Liston1,2*

1National Museums Scotland, Edinburgh, SCOTLAND; 2Peterborough Museum & Art Gallery, Peterborough, England

*[email protected]

Keywords: Jurassic, pachycormid, Leedsichthys, public sector display, paraloid B72, volunteers

Over field seasons 2002 and 2003, the most complete specimen of the large suspension-feeding fish Leedsichthys was excavated from the Star Pit, just outside Whittlesey, one of the last clay brick pits that Alfred Leeds collected from. Consisting of over 2,300 parts, the specimen became known as ‘Ariston’ because it went on and on, rather like the old utility advertisement. Over 100 days of excavation involved over 3,100 staff hours to extract this prodigious quantity of material, achieving the record of the longest dig for a single vertebrate specimen in Europe.

In the ensuing years, over 11,750 hours of preparation time was conducted by volunteers at Peterborough Museum and Art Gallery, led by Alan Dawn. In July 2015, the Esmée Fairbairn Foundation awarded a £66K grant to the museum through the Museums Association’s Collections Fund, to provide a ‘virtual display’ of the specimen. Although such approaches can often be seen as low-effort substitutes to displaying the real object, undermining actual displays by encouraging potential visitors to migrate to the internet instead, this was a solution to a very real practical problem: the bones of this fish, while occasionally large are always extremely fragile, and would require an exceptionally long and complex armature to display them – an armature for which no gallery in the museum was large enough.

In addition, virtual display allowed the possibility of incorporating far more associated information with the specimen than would be permitted according to museum orthodoxy.

In order to achieve this, a significant amount of backdated curation was undertaken, as well as translating the original 26 x 12 metre area of plastic maps into a navigable digital form, and identifying the bones and arranging them within the first ever skeletal reconstruction of this animal - the required research often being completed only days before required delivery for the final product. Thus the resulting website tells the story from the first discovery through the 100 days of excavation, the site map translates into the bones that form the skeleton, which fleshes to an animated reconstruction of the 9 metre long ‘Ariston’ specimen of Leedsichthys from Whittlesey.

134 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

SPLASHING AND WALKING WITH EARLY TETRAPODS: EARLY PERMIAN ICHNOLOGICAL RECORD FROM THE CATALAN PYRENEAN BASIN E. Mujal1*, J. Fortuny2,3, O. Oms1 and A. Bolet3

1Departament de Geologia, Universitat Autònoma de Barcelona (UAB), E-08193 Bellaterra, Spain; 2C2RP, CNRS-MNHN-UPMC, 8 rue Buffon, CP38, F-75005 Paris, France; 3Institut Català de Paleontologia Miquel Crusafont, ICTA-ICP building, c/ de les columnes, s/n, E-08193 Cerdanyola del Vallès, Spain

*[email protected]

Keywords: Vertebrate ichnology, swimming scratches, anmniote traces

Tetrapod footprints are the most abundant vertebrate record of the Late Paleozoic and therefore crucial in the understanding of early tetrapod locomotion, habitats, and evolution. The Early Permian ichnological record is known worldwide and particularly well represented. It is often encountered in red-bed deposits resulting from the aridization of Pangea. In this period, the Pyrenees (NE Iberian Peninsula) existed in the equatorial region of the supercontinent as sedimentary basins infilled by volcanosedimentary deposits.

In the recent years, we analyzed several outcrops bearing abundant tetrapod trace fossils in the Catalan Pyrenees. As a result, we were able to identify a wide diversity of ichnomorphotypes attributed to temnospondyls, possible seymouriamorphs, possible diadectomorphs, araeoscelids, bolosaurids, captorhinids and . Of particular interest for the track maker’s behavior, we discovered three different forms of swimming traces, corresponding to the ichnogenus Characichnos and associated with small- and large-sized temnospondyls (Batrachichnus salamandroides and sp., respectively), and possible seymouriamorphs (cf. Amphisauropus). These remains represent an excellent tool to deepen the understanding of early tetrapodlocomotion.

The swimming traces correspond to digit tip prints dragged over the surface, i.e., scratches. The ichnites occur in groups of two to four digit scratches, ranging in length from 5 to 60 mm. Scratches are mostly curved or slightly sinuous (sigmoidal), but sometimes also straight. These traits, together with the expulsion rims commonly observed in the posterior part, record the limb movement of the trackmakers. Moreover, several surfaces present transitions from walking to swimming traces. Trackways composed of scratches and rounded digit tip impressions display irregular patterns indicating a relatively buoyant gait of the trackmaker. In all, these early tetrapods were capable of both terrestrial and subaqueous locomotion, which permitted a wide environmental distribution and colonization of the continental realm.

135 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Insular Vertebrate Palaeontology poster V2

PRESENCE OF BARBATODON OARDAENSIS (MAMMALIA, MULTITUBERCULATA) IN THE UPPER CRETACEOUS OF THE RUSCA MONTANĂ BASIN (SOUTHWESTERN TRANSYLVANIA, ROMANIA) V.A. Codrea1*, A.l. Solomon1, M. Venczel2, C. Fărcaș1 and T. Smith3

1University Babeș-Bolyai, Department of Geology, 1 Kogălniceanu Str., 400084 Cluj-Napoca, Romania 2Ţării Crişurilor Museum, Department of Natural History, Dacia Av. 1-3, 410464, Oradea, Romania 3Royal Belgian Institute of Natural Sciences, Directorate Earth and History of Life, Rue Vautier 29, B-1000, Brussels, Belgium

*[email protected]

Keywords: Multituberculates, kogaionids, Maastrichtian, Occidental Carpathians, Romania

Multituberculates – often called the “Mesozoic rodents” – occurred early in the European fossil record and evolved from Middle Jurassic to Palaeogene on the old continent. Since the Late Jurassic, multituberculates were abundant in Europe, the diversity of families increasing in the Early Cretaceous. In the Upper Cretaceous of Europe, these mammals are scarce, being recorded exclusively from the terrestrial deposits of the Hațeg and Transylvanian basins (Romania). Moreover, the multituberculates known from this area belong to a unique family - Kogaionidae, documented by two genera (Kogaionon – with the single species, K. ungureanui - and Barbatodon – with two species, B. transylvanicum and B. oardaensis). The group succeeded in crossing the K/T boundary. However, they were replaced around the Late Palaeocene by the world-widely distributed North American Neoplagiaulidae, the last family occurring in Europe before the extinction of the multituberculates from the old continent in Eocene. Here, we report multituberculate teeth originating from the Maastrichtian fluviatile sediments of the Rusca Montană Basin (Occidental Carpathians). This is the westernmost occurrence of these Cretaceous mammals in Romania. These teeth are assigned to Barbatodon oardaensis, the smallest Cretaceous kogaionid, and represent the first record of this species outside the Metaliferi sedimentary area (southwestern Transylvania). Based on the known discoveries since the mid-1980’s, the distribution of Maastrichtian kogaionids can now be summarized as follows: i. small-sized kogaionids were present in the Metaliferi area of the Transylvanian Basin, in the Densuș-Ciula Formation of the northern side of the Hațeg Basin as well as in the Rusca Montană; ii. small and medium-sized kogaionids were recorded from the Râul Mare sites in the Sânpetru Formation of the Hațeg Basin; iii. the largest kogaionids, K. ungureanui and B. transylvanicus, were reported from the well known Sânpetru and Pui localities (Sânpetru Formation), also in the Hațeg Basin .

This research was supported by grant PN-II-PCE-2011-3-0381 of the National Council of Scientific Research - CNCS (to V.C., A.S., M.V., C.F.) and Projects MO/36/001-004 of the Belgian Science Policy Office (to T.S.).

136 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Insular Vertebrate Palaeontology poster V3

RECONSTRUCTING INTERNAL ANATOMY OF BONE LAYER IN MAURITIAN DODO-LAGERSTÄTTE: 3D-BONE MAPPING USING GEOGRAPHIC INFORMATION SYSTEMS J.K. Lubeek1,2*, M.N. Jansen2, S.J. Kluiving2, P.G.B. De Louw3, H.J.M. Meijer4, J.P. Hume5, J. De Vos6, A. Janoo7, L.P.A.M. Claessens8,6, R. Floore9 and K.F. Rijsdijk10

1Department of Earth Sciences, Utrecht University, Heidelberglaan 2, 3584 CS Utrecht, The 2 Netherlands; Institute for Geo- and Bioarcheology, Faculty of Earth and Life sciences, VU University Amsterdam, De Boelelaan 1085, 1081 HV Amsterdam, The Netherlands; 3Department of Soil and Groundwater, Deltares, Postbus 85467, 3508 AL, Utrecht, The Netherlands; 4The Natural History Collections, University Museum of Bergen, University of Bergen, Realfagbygget, Allègt. 41, Postboks 7800, 5007 Bergen, Norway; 5Bird Group, Department of Life Sciences, Natural History Museum, Akeman Street, Tring Herts, HP23 6AP, UK; 6Naturalis Biodiversity Center, Darwinweg 2, 2333 CR Leiden, The Netherlands; 7Department of History and Political Science, University of Mauritius, R_eduit, Mauritius; 8College of the Holy Cross, Department of Biology, 1 College Street, Worcester, Massachusetts 01610, USA; 9Hollandia Archeologen, Tuinstraat 27A, 1544RS Zaandijk, The Netherlands; 10Computational GeoEcology Group, Institute for Biodiversity and Ecosystem Dynamics, University of Amsterdam, P.O. Box 94248, 1090 GE, Amsterdam,The Netherlands *[email protected]

Keywords: 3D-bonebed analysis, GIS, dodo, lagerstätte, taphonomy

We present a novel workflow for the analysis of bonebeds in 3D involving Geographic Information Systems (GIS), geometrical corrections and in situ sampling by means of a rotary scoop. In order to assess and reconstruct death and post- mortem scenarios that gave rise to the bone layer of 4,200 yr old Mare aux Songes Dodo (Raphus cucullatus)-Lagerstätte in Mauritius, spatial bone distribution, 3D- orientation and completeness of vertebrate bones and tree stems were mapped, visualised and analysed by means of GIS and assessed geostatistically. In 2009, a dry excavation pit was realised in the marsh. The pit was located within the reconstructed palaeo-centre of the lake, representing conditions of low-energy lacustrine accumulation. Bone and wood orientations were measured by compass and their 3D-positions were mapped. In 2010, the Dodo-polder flooded, but dry excavation and 3D-measurements were continued in 2011 by means of a specially designed rotational scoop, measuring 190 cm by 140 cm, the so-called “Floore-arc”. During lifting of the Floore-arc the originally horizontally deposited layers were tilted and folded. In this current study the deformation and implemented structural geometric corrections were assessed in order to reconstruct original in situ compass orientation measurements. XYZ-Positions of 423 bone and wood elements were mapped and described during excavation. Objects were assigned centroids to generate a 3D-model in GIS. We found that all bones of all species are chaotically mixed, with no articulation and only one single case of poor association. Based on MNI estimates and total bone finds per species, skeleton incompleteness is estimated >90% for all species. Bones show no statistically preferred orientation and the mean dip of bones are generally significantly less than 20°. Lower limbs dip low- angle on average at 21°. Based on these findings we conclude that high degree of post-mortem bioturbation and hydrotaphonomic processes are the dominant signature of the Mare aux Songes Dodo-Lagerstätte. 137 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Insular Vertebrate Palaeontology poster V4

BONES, SHELLS AND MILLIPEDES: TAPHONOMY OF A JAMAICAN CAVE S.K. Donovan1*

1 Naturalis Biodiversity Center, Darwinweg 2, 2333 CR Leiden, The Netherlands

*[email protected]

Keywords: Red Hills Road Cave, karst, Pleistocene, preservation

The Red Hills Road Cave (or fissure), parish of St Andrew, near Kingston, Jamaica, is an insignificant remnant of a karstic feature that was quarried away during road building before the mid-1980s. Nevertheless, it is the most important site for Late Pleistocene terrestrial palaeontology on the island. The site is about 30,000 years old (oxygen isotope stage 3), but may span at least 15,000 years. Many taxa were probably washed in during hurricanes and tropical storms, either as dead skeletons or live organisms that drowned as the cave filled with water.

The invertebrate fauna includes both land snails and arthropods, largely or entirely derived from the surrounding area; none are obligate cave dwellers. The 62 species of land snails are the most diverse of any Jamaican cave; about half the snails in the cave live in the local area today. The arthropods include the only fossil millipedes, isopods and insects (fly puparia, beetle elytra) in the Jamaican fossil record, in addition to a land crab. Millipedes and isopods are well preserved because of a diagenetically early coating of calcite cement. The exoskeletons of these groups contain a small, but significant, calcite component not found in insects, spiders and scorpions.

The vertebrate fauna remains remain understudied, but include a rodent, three species of bat and possibly a marsupial; a flightless ibis, the Jamaican tody and various other birds; and reptile and amphibian remains. In contrast to the arthropods, the vertebrates are invariably disarticulated, apart from rare crania, jawbones retaining teeth and bones that are fused in life. A dead millipede could be coated in calcite when floating in the cave; a dead vertebrate carcass would have to rot to expose its bones, likely after the cave had dried out.

138 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Insular Vertebrate Palaeontology poster V5

OSTEOPHAGIA AND ANTLER MODIFICATION BY CRETAN DEER FROM BATE CAVE S. Altamura1 and M.R. Palombo1,2*

1Sapienza University of Rome, Department of Earth Sciences, P.le A Moro 5, 00185 Roma, Italy 2IGAG-CNR, Via Salaria km 29,300, 00015 Monterotondo (Roma), Italy

*[email protected]

Keywords: Candiacervus, Crete, Pleistocene, antler chewing, fork formation

During the Middle to Late Pleistocene, highly derived deer species, differing in size and ecological behaviour, were among the most common endemic mammals inhabiting the Crete island. Eight morphotypes/species belonging to a single genus (Candiacervus) have been described, attesting for an evolutionary radiation process to occupy an array of empty niches, ranging from forest to rocky environment. This research describes some modifications observed in shed antlers belonging to the smallest Cretan deer (Candiacervus ex gr. C. ropalophorus) in aim to identify marks indicative of an osteophagic behaviour by deer. We have inspected 63 shed antlers belonging to the rich sample collected in the 70th by a team of Sapienza University of Rome (Italy) in an unexplored cavity (named “Bate cave” by the Italian discoverers), located in the NW Cretan coast (Rethymnon). Two amino-acid racemisation dates on deer teeth (152,000±20% to 105,000±20%) suggest a late Middle to early Late Pleistocene age for the accumulation of this fossil assemblage. The osteophagic behaviour is quite common among artiodactyls. Detecting the presence and estimating the extent of osteophagia in fossil species may give some support to palaeoecological investigation, and provide clues on environmental conditions and resource exploitation by herbivores. The osteophagic activity is due to a mineral deficiency in their diet, in particular the lack of elements such as calcium and phosphorus. The lack of these nutrients in plants, which may depend on the nature of the soil and climate factors, led some animals to chew antlers and dry and weathered bones. Among the shed antlers of small deer from Bate cave, the 76% shows chewing marks. Some a few antler tips have a typical forked shape, a number of tines and some antler burrs show grooves resulting from the molar dragging in the typical zigzag mastication movement of deer. The high percentage of remains showing osteophagic marks suggest that the antler chewing was a quite common practise among deer from Bate cave and their diet was poor in mineral nutrients, in line with the rocky environment they inhabited and the climate condition at that time.

139 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Insular Vertebrate Palaeontology poster V6

AN ENDEMIC RADIATION OF DEER IN THE LATE PLEISTOCENE OF MALTA 1 1 L. D’Souza and A.M. Lister *

1Department of Earth Sciences, Natural History Museum, London SW1 5BD, UK

*[email protected]

Keywords: Deer, Malta, Quaternary, endemism, taxonomy

Mediterranean Quaternary deposits reveal a remarkable history of islands inhabited by dwarfed populations of elephants, hippopotamuses and deer during the Late Pleistocene. Although many of these taxa are well documented, others remain poorly understood, due partly to imprecise dating and inadequate sampling. The present research aims to document an island mammal radiation using Malta’s endemic deer as a case study. Thousands of fossils recovered from Għar Dalam Cave (Malta) show exceptional variation in size and form, suggesting at least four size classes (potential species); this contrasts with the limited size variation found on the much larger island of Sicily, from where the ancestors of the Maltese deer are presumed to have originated. There is also remarkable variation in antler form, which may or may not have taxonomic significance. Morphological characters on bones and teeth, established to separate red, fallow and giant deer (the most plausible mainland ancestors), in combination with metric data, are used to investigate likely ancestry and possible taxonomic diversity. Adaptive diversity is also being studied; early results indicate that Maltese deer are not simply isometrically-scaled versions of modern species; some possess the robust distal limb proportions commonly seen in island dwarfs. Dental mesowear analyses suggest that all size groups contain both browsing and grazing individuals, but mixed feeders are mainly restricted to smaller size classes. These morphological and taxonomic analyses will be integrated with new dating and stratigraphic information to provide a chronological context for the evolutionary patterns observed.

140 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Insular Vertebrate Palaeontology poster V7

THE CLOCK HANDS OF SICILIAN BIOCHRONOLOGY 1 1 1 2 F. Masini *, D. Petruso , G. Surdi and A. Savorelli

1University of Palermo, Department of Earth and Marine Sciences (DISTEM), via Archirafi 22, 90123 Palermo, Italy 2University of Florence, Department of Earth Science, Via La Pira 4, 50121 Florence, Italy

*[email protected]

Keywords: Small mammals, endemics, Quaternary, Sicily, Italy

The Sicilian Quaternary terrestrial mammals are biochronologically arranged as "Faunal Complexes" (F.C.) with decreasing endemic features. Small mammals associated with the macro mammals of the Palaeoloxodon mnaidriensis F.C., (late Middle Pleistocene-early Late Pleistocene) and of the youngest Castello F.C, (Lateglacial) were less known than the endemic small mammals from the oldest F.C.'s (Monte Pellegrino F.C.-Early Pleistocene and Palaeoloxodon falconeri F.C.- Early Middle Pleistocene).

Riparo dell'Uzzo (Capo San Vito) first documented Pleistocene/Holocene and early Holocene macromammals, together with Microtus (Terricola), Apodemus and Crocidura, still extant in Sicily. A stratigraphically-controlled P. mnaidriensis F.C. large mammal assemblage from Contrada Fusco (Siracusa) yielded Leithia melitensis, Crocidura esuae and Maltamys wiedicitensis. These finds follow biochronologically the older P. falconeri F.C., which is documented in a few sites where L. melitensis, Maltamys gollcheri and C. esuae are associated with pigmy elephant. In two "pockets" of a karstic cavity at Contrada Pianetti (Ragusa) P. mnaidriensis is associated with Leithia (Pocket A), whereas Pocket B yielded Microtus (Terricola) ex gr. savii, Apodemus sylvaticus and Crocidura sicula together with several large mammalian taxa. Locality K22 (Capo San Vito), and Alcamo Cappuccini fissure yielded endemic dormice and , with P. mnaidriendsis F.C. large mammals. An upper level at K22 provided a small mammal assemblage like that of Contrada Pianetti. Non-endemic small mammals from San Teodoro Cave are associated with P. mnaidriendsis F.C. large mammals, among which Equus hydruntinus, an equid considered exclusive of the Castello F.C. This assemblage is characterized by the co-occurrence of older large mammals and new small mammalian incomers. It constitutes a new late Pleistocene (MIS 4-3) Grotta San Teodoro - Pianetti F.C. Finally, the lower deposits of site Isolidda 2 (Capo San Vito) gave remains of P. mnaidriensis, Maltamys wiedincitensis and Crocidura esuae and includes the last occurrence of Leithia preceding the first occurrence M. (Terricola) nov. sp. The site records the transition between the P. mnaidriensis - S. Teodoro- Pianetti F.C.'s. The analysis of M. (Terricola) revealed three dispersal events from southern Italy at MIS 6, 4 and 2 sea lowstands.

141 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Insular Vertebrate Palaeontology poster V8

DOES NEUTRAL THEORY APPLY TO INSULAR FOSSIL MAMMALS? A CASE STUDY OF THE PLEISTOCENE FAUNA FROM SARDINIA AND SICILY (MEDITERRANEAN) M.R. Palombo1,2* and R. Rozzi1

1Sapienza University of Rome, Department of Earth Sciences, P.le A Moro 5, 00185 Roma, Italy 2IGAG-CNR, Via Salaria km 29,300, 00015 Monterotondo (Roma), Italy

*[email protected]

Keywords: Mammal, Sardinia, Sicily, Pleistocene, Neutral Theory

The Unified Neutral Theory of Biodiversity and Biogeography (UNTB), inspired by MacArthur and Wilson’s theory of island biogeography, has attracted increasing attention over the last fifteen-years. Only few efforts, however, have been put into testing and applying it in the field of island biogeography and palaeobiology. This research focuses on Pleistocene mammalian faunal complexes (FCs) of the large central Mediterranean islands, Sicily and Sardinia, that differ in their palaeobiogeographic histories and composition of mammalian endemic fauna. The research aims to investigate whether: i) the dispersal parameters [m= probability that any disappearing individual is replaced by an immigrant and I (=m(J-1)/(1-m))], and the “fundamental biodiversity number” θ (asymptotically equal to Fischer’s α diversity), estimated for each FC match or not the trend of Fischer’s α calculated from the empirical data; ii) the trends are consistent or not with the palaeobiogeographic settings proposed for Sardinia and Sicily; iii) the log-normal and ZSM distributions appreciably fit or not the empirical data on the relative species abundance; iv) the neutral predictions are useful or not in evaluating the community inertia on islands at large temporal scales.

Results obtained show that the general trends of the neutral parameters m, I and especially θ are substantially in agreement with the evolutionary model expected for the studied insular faunas. A number of important points, however, need to be clarified. In particular, problematic issues concern the empirical over-representation of rare taxa, the effective best-fitting of the lognormal with respect to the zero-sum multinomial distribution (ZSM) and the existence of some disagreements between the estimated community inertia and the turnover index (TI) at the transition between both faunal complexes and faunal subcomplexes (FSCs). To answer the questions we need to clarify to what extent the period of persistence of the FCs/FSCs and the immigration and renewal rates on one hand, and the fact that our research is focused on mammals and species-poor insular communities on the other, might affect the shape of the species abundance distribution (SAD).

142 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Insular Vertebrate Palaeontology poster V9

NEW MAMMOTH REMAINS FROM THE LATE PLEISTOCENE OF ALGHERO (SARDINIA, CENTRAL MEDITERRANEAN) 1 2 3 M. Zedda , M.R. Palombo * and R.T. Melis

1University of Sassari, Veterinary Medicine Department, via Vienna 2, 07100 Sassari, Italy 2Sapienza University of Rome, Department of Earth Sciences, P.le A Moro 5, 00185 Roma, Italy - IGAG-CNR, Via Salaria km 29,300, 00015 Monterotondo (Roma), Italy 3University of Cagliari, Department of Chemical and Geological Sciences, Via Trentino, 51, 09127 Cagliari, Italy

*[email protected]

Keywords: Mammuthus, Sardinia, Late Pleistocene

Endemic elephants belonging to Palaeoloxodon lineage have been largely reported in the Middle and Late Pleistocene deposits of a number of Mediterranean islands (e.g. Sicily, Malta and the Aegean Islands, Crete, the Cyclades, Dodecanese Islands and Cyprus). Nonetheless, the straight-tusked elephants, in spite of their swimming ability, never colonized the Corso-Sardinian massif.

In Sardinia, elephant postcranial remains belonging to a single individual, were first reported at the end of the 19th century by in the Morimenta area (Gonnesa, SW Sardinia), and then described as the new species Elephas lamarmorae Major, 1883. The morphology of these remains does not contradict their attribution to a Mammuthus representative (Mammuthus lamarmorai) as confirmed by the Schreger line pattern of the tusk. Mammuthus is recorded in Sardinia from few molars discovered in alluvial deposits (possibly late Middle Pleistocene in age) filling the Campu Giavesu Basin (Sassari, north-western Sardinia), in the pre-Tyrrhenian pedogenized beach deposit (?MIS 7 or older) of San Giovanni di Sinis (Oristano, W Sardinia), and in the post-Tyrrhenian (post-MIS 5e) breccia cropping out at Tramariglio (Alghero). The modest fossil record and some size variation inferred poses intriguing questions as whether only one mammoth species inhabited Sardinia and the dimensional scaling results from a progressive dwarfing due to anagenetic evolution, or mainland mammoth entered the island more than one time, giving rise to different species, or the size depends on some introgressive hybridization.

This research aims to provide additional data by describing an incomplete radius, larger in size than that from Morimenta, which was found in cemented sandstone deposits, possibly deposited during the last glacial in a backshore environment, cropping out along the coast in the surroundings of Alghero (NW Sardinia).

Histological studies have been performed to scrutinize whether the histomorphology of the bone tissue of the studied radius differs from that of mailand species. Transverse and longitudinal sections of the mesosteal zone of the radius diaphysis were observed and studied by means of an Axiophot microscope. Osteons were examined on order to number their lamellae and osteocytic lacunae.

143 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Insular Vertebrate Palaeontology poster V10

THE INTRIGUING QUESTION OF THE PLEISTOCENE SARDINIAN OTTERS 1 2 M.R. Palombo * and M. Zedda

1Sapienza University of Rome, Department of Earth Sciences, P.le A Moro 5, 00185 Roma, Italy - IGAG-CNR, Via Salaria km 29,300, 00015 Monterotondo (Roma), Italy 2University of Sassari, Veterinary Medicine Department, via Vienna 2, 07100 Sassari, Italy

*[email protected]

Keywords: endemic otters, Sardinia, taxonomy, ecology, evolution.

Endemic otters showing different adaptation, ecological behaviour and trends towards change in body mass are known in the Mediterranean islands since the Late Neogene (i.e. Tyrrhenolutra helbingi, Paludolutra maremmana, and Paludolutra campanii and Paralutra garganensis from the Late Miocene of Tusco-Sardinian and Abruzzi-Apulian palaeobioprovinces respectively). An apparent long temporal gap separates these otters from the Quaternary ones mainly recorded from single findings (either nearly complete skeletons or scattered remains), whose age has been tentatively estimated to the Middle or Late Pleistocene. From the Corso- Sardinian massif four species have been described, the Sardinian species Sardolutra ichnusae and Megalenhydris barbaricina known by almost complete skeletons, Algarolutra majori known in Sardinia and Corsica from some upper molars and a few limb bones, and Lutra castiglionis thus far only recorded in Corsica in the late Middle Pleistocene deposits of Castiglione Cave, and believed to be morphologically close to the mainland ancestor Lutra simplicidens, which is latest recorded in the Italian peninsula in Early Pleistocene deposits of Upper Valdarno Basin.

This researches focuses on an almost complete skeleton, undescribed until today, found in Sardinia in the 70ths of the last century at Grutta Sa Folla (Nuxis, SW Sardinia), a cave nowadays located in a hilly area, about 22-25 km far from the sea. The otter, slightly larger than L. castiglionis and S. ichnusae, shares with L. castiglionis some key features (e.g. absence of a sagittal crest, humerus less curved than in S. ichnusae, no fused tibia and fibula, possibly the baculum size). Some of these traits are consistent with a riverine aquatic life as suggested for the Corsican otter and supported by the environmental context of the cave. Although the comparison among these three otters cannot be exhaustive due to incompleteness of the Corsican remains, available evidences poses some intriguing questions as whether intraspecific variation in size, the role of dietary behaviour and environment in giving rise to different ecomorthypes belonging to the same spacies/genus, the actual presence in Sardinia and Corsica of two different genus, and the evolutionary pattern of Corso-Sardinian otters in the course of the Pleistocene.

144 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Insular Vertebrate Palaeontology poster V11

MANDIBULAR FORCE PROFILES AND IMPLICATIONS FOR THE FEEDING BEHAVIOUR OF THE ENDEMIC DOG CYNOTHERIUM M.R. Palombo1*

1Sapienza University of Rome, Department of Earth Sciences, P.le A Moro 5, 00185 Roma, Italy - IGAG-CNR, Via Salaria km 29,300, 00015 Monterotondo (Roma), Italy

*[email protected]

Keywords: Biomechanics, bite force, mandible, Pleistocene, Sardinia

Non-aquatic Carnivora are uncommon in impoverished, disharmonic insular faunas. The fossil record of large Pleistocene Canidae includes few taxa know from Java (Megacyon merriam and Mececyon trinilensis) and the Corso-Sardinian massif (Cynotherium sp., Cynotherium malatestai, Cynotherium sardus). C. sardous is the best known endemic carnivorous species, mainly recorded from Late Pleistocene deposits, although the origin of the endemic lineage possibly goes back to the latest Early Pleistocene. The ancestor, the hypercarnivorous Lycaon-like dog Xenocyon lycaonoides, entered Sardinia by the end of the Early Pleistocene. The dentition of its descendants retained some hypercarnivorous traits, though a change in diet to small mammals (i.e the extraordinary abundant ochotonid Prolagus) and birds likely triggered a progressive reduction in size and the appearance of traits presumably related to pedomorphsis in the skull morphology of C. sardous. As a result the species shows a unique combination of a strong dentition with hypocarnivorous cranial features.

A biomechanical approach to model mandibular force profiles from mandibular dimensions has been applied to provide new evidence on the feeding behaviour of Cynotherium. The method permits to infer the bending strength on both the dorso- ventral (Zx/L) and labio-lingual (Zy/L) planes of the mandible, and the relative mandibular force (or overall mandibular shape) (Zx/Zy). Mandibular force profiles of C. sardus mandibles belonging to the rich sample from Medusa-Dragonara cave (NW Sardinia, Last Glacial) show that the dorso-ventral force was lower at the canine than at the P4 –M1 interdental gap, and particularly high behind M3. This dorso-ventral buttressing is typical of those extant canids that use their post-carnassial teeth for cracking bones. Labio-lingual force is conversely almost uniformly distributed along the premolar-molar tooth-row. A roughly constant relative force value in the region where flesh is processed reflects some adaptation to slicing. The relative mandibular force confirms on one hand a dorso-ventral buttressing behind the carnassial, particularly behind M3, on the other the weak canine bite. This feature is consistent with the hunting behaviour of canids, which hunt in packs and kill their prey by repeated superficial bites, or prey on small mammals.

145 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday plenary poster R1

REVISION OF THE EARLY EOCENE PRIMATE AGERINIA ROSELLI FROM ITS TYPE LOCALITY, LES SALERES (NE IBERIAN PENINSULA) J. Femenias-Gual1*, R. Minwer-Barakat1, J. Marigó1,2 and S. Moyà-Solà3.

1Universitat Autònoma de Barcelona, Institut Català de Paleontologia Miquel Crusafont, 08193, Cerdanyola del Vallès, Barcelona, Spain 2Muséum National d’Histoire Naturelle, UMR 7207 CR2P - C.N.R.S., M.N.H.N., U.P.M.C.- Paris6. Département Histoire de la Terre, Paris, France 3Universitat Autònoma de Barcelona, ICREA at Institut Català de Paleontologia Miquel Crusafont and Unitat d’Antropologia Biològica (Dept. BABVE), 08193, Cerdanyola del Vallès, Barcelona, Spain

*[email protected]

Keywords: Adapiformes, Cercamoniinae, Notharctidae, Paleogene, Spain

The scarcity of early Eocene primate remains in Europe increases the importance of each new finding and the relevance of the information that they provide. Several unpublished dental remains of Agerinia roselli (Primates, Adapiformes) from its type locality, Les Saleres (NE Spain), have recently been found in the collections of Institut Català de Paleontologia Miquel Crusafont. The whole sample from this site (including both already published and unpublished remains) consists of twelve lower teeth from P3 to M3 and two root fragments placed mesially with respect to the P3. Its detailed analysis has allowed for finding some inaccuracies in the previous descriptions of this species (Crusafont-Pairó, 1967; Szalay, 1971). This new study solves these mistakes and emphasises several important traits of A. roselli that remained unknown up to date, such as the distinct hypoconid and entoconid on the P4 or a presence of a tiny paraconid on the M1. This latter feature seems to be the most consistent criterion to discern between the first and second lower molars when they are found isolated, instead of the trigonid length, as proposed in previous works. Regarding the anterior roots, they can either correspond to a double-rooted P2 or, more probably, to single rooted P1 and P2, since the most mesial root is strongly shifted towards the buccal side of the mandible. Furthermore, the body mass of A. roselli has been estimated on the basis of the size of the lower molars, ranging from 650 to 900 g. To conclude, this review updates the knowledge about the dental anatomy of A. roselli and demonstrates that it is a valid taxon clearly distinguishable from other cercamoniines.

This study has been supported by the Spanish Ministerio de Economía y Competitividad (CGL2014-54373-P) and by the Generalitat de Catalunya (2014/100604; 2014 SGR 416, GRC). JF-G has been financed by BES-2012-052951 and EEBB-I-14-08986 and JM by Fondation Fyssen.

146 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday plenary poster R2

THE STUDY OF THE RODENTS FROM SANT JAUME DE FRONTANYÀ-1 (MIDDLE EOCENE, NORTHEASTERN SPAIN) AND ITS BIOCHRONOLOGICAL SIGNIFICANCE I. Bonilla-Salomón1*, R. Minwer-Barakat1, M. Vianey-Liaud2 and S. Moyà-Solà3

1Universitat Autònoma de Barcelona, Institut Català de Paleontologia Miquel Crusafont, 08193, Cerdanyola del Vallès, Barcelona, Spain 2Université de Montpellier, Institut des Sciences de l’Evólution de Montpellier (ISE-M, UMR 5554, CNRS/UM/IRD/EPHE), Laboratoire de Paléontologie, c.c. 064, Place Eugène Bataillon, F-34095 Montpellier cedex 5, France 3Universitat Autònoma de Barcelona, ICREA at Institut Català de Paleontologia Miquel Crusafont and Unitat d’Antropologia Biològica (Dept. BABVE), 08193, Cerdanyola del Vallès, Barcelona, Spain

*[email protected]

Keywords: Gliridae, Theridomyidae, Rodentia, Robiacian, Iberian Peninsula

Eocene mammals from Spain have received increasing attention in the last decades. Sant Jaume de Frontanyà (Eastern Pyrenees) represents the most significant middle Eocene site from the Iberian Peninsula, including four fossil-bearing levels with an extraordinary abundance and diversity of mammal remains. However, the fauna from this locality has not been studied in detail. Late works have been mostly focused on the primates from the oldest levels, including the description of several new species. Recently, the new primate Necrolemur anadoni has been described from the uppermost level, Sant Jaume de Frontanyà-1 (SJF-1). On the contrary, other mammal groups have been practically undescribed to date. This is the case of rodents, which are especially useful for biostratigraphic correlations.

The rodent remains from SJF-1 have been now described in detail. Three different forms have been identified: one glirid, ascribed to Glamys aff. robiacensis, and two new theridomyid species that represent the most primitive members of Paradelomys and Elfomys. These three forms show less advanced traits than other species of these genera from localities assigned to the MP16 such as Robiac, Grisolles, and Le Bretou. Therefore, SJF-1 is assigned to the MP15, with an age slightly older than Chéry-Chartreuve, and represents the first occurrence of Paradelomys and Elfomys. This dating allows us to place in a correct chronological framework the only record of Necrolemur from Spain, shedding new light on the evolution of this primate genus. Further studies of the abundant rodent remains from the oldest levels of Sant Jaume de Frontanyà, SJF-3C and 3D, will notably improve the knowledge about the still poorly studied mammal assemblages from this time span.

This study has been supported by the Spanish Ministerio de Economía y Competitividad (CGL2014-54373-P) and by the Generalitat de Catalunya (2014/100604; 2014 SGR 416, GRC

147 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday plenary poster R3

THE OLDEST SIRENIAN REMAIN FROM EUROPE (EARLY

LUTETIAN, HUESCA, SPAIN) E. Díaz-Berenguer1*, R. Silva-Casal2, A. Badiola3, J.I. Canudo4, A. Payros3 and 2 M. Aurell

1Grupo Aragosaurus-IUCA, Dpto. Ciencias de la Tierra, Facultad de Ciencias, Universidad de Zaragoza, Calle Pedro Cerbuna 12, 50009 Zaragoza, Spain 2Área de Estratigrafía, Dpto. Ciencias de la Tierra, Facultad de Ciencias, Universidad de Zaragoza, Calle Pedro Cerbuna 12, 50009 Zaragoza, Spain 3Dpto. Estratigrafía y Paleontología, Facultad de Ciencia y Tecnología, Universidad del País Vasco (UPV/EHU), Apartado 644, 48080 Bilbao, Spain 4Grupo Aragosaurus-IUCA, Museo de Ciencias Naturales, Universidad de Zaragoza, Calle Pedro Cerbuna 12, 50009 Zaragoza, Spain

*[email protected]

Keywords: sirenian, early Lutetian, pachyosteosclerosis, Europe

The osteoesclerosis, or inner bone compaction, and the pachyostosis, or hyperplasia of bone cortices, is one of the best diagnostic characteristic of the sirenian bones. The bone pachyosteosclerotic condition observed in the recently found early Lutetian fossil remain of the Arguis locality (Southern Pyrenees, Spain) allows identifying it as a sirenian rib. It represents the earliest accurately dated sirenian from the Lutetian of Europe and the oldest known from this continent. The Lutetian sirenian fossils are scarce in Europe, and excepting the Hungarian taxa Sirenavus hungaricus Kretzoi, 1941, both the age and the status of the existing material are uncertain and lack precise dating. The fossil comes from the Guara Formation, Isuela Section (Southern Pyrenees, Huesca, Spain). Isuela section is one of the most studied outcrops in the External Sierras, giving a strong biostratigraphic, magnetostratigraphic and lithostratigraphic framework to the Guara Fm. This unit is manly composed of carbonate facies deposited in a transitional to middle ramp environments. The sirenian rib appeared in transitional facies, in the lower Guara Fm. This unit contains several larger foraminifers, related to the SBZ 13. Moreover, available magnetostratigraphic data allows a precise age of the fossil, located close to the C22r - C21n chron transition, giving an age of 47.349 Ma. The rib fragment of Arguis shows a lateromedially flattened shaft with an oval cross-section. The inner bone is composed by dense tissue and lacks the medulla canal. It probably belongs to the first third of an undetermined rib. Recently discoveries have proven that the southern Pyrenees is an unrivaled area to study the early stage of the evolution of this group of afrotherian mammals. In the nearby site of the Castejon of Sobrarbe (Ainsa Basin, Sobrarbe Formation, Huesca) the most complete fossil collection of the Lutetian sirenian known in Europe has been recovered, which corresponds to the first taxon with functional hindlimbs of the northern of Tethys, whereas in the Early Lutetian deposits of Arguis the oldest sirenian record known from Europe is described.

148 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday plenary poster R4

CONCERNING THE EAST CARPATHIAN FORELAND MAMMAL BIOSTRATIGRAPHY (EASTERN ROMANIA, MOLDOVA AND WESTERN UKRAINE) D.J. Scager1* , W. Wessels1 and A. de Leeuw2

1Utrecht University, Utrecht, the Netherlands 2CASP, Cambridge, United Kingdom

*[email protected]

The East Carpathian Foreland is a high potential area for the construction of a consistent local mammal biostratigraphy due to its rich and high quality records, internally consistent marine biostratigraphy, and the advances in Paratethys magnetostratigraphy and isotopic dating of the past 10 years.

However, information on the existing fossil localities is distributed over a plethora of publications in at least four different languages. Various biozonations for both marine and terrestrial domains have furthermore been devised over time. The terrestrial biozonations are not yet in agreement with one another, due to the lack of unanimous substage boundary definitions and differing correlations to the MN (Mammals Neogene) system. In order to overcome these problems and access the full potential of the fossil record, an overview is made of the state of the art of mammal biostratigraphy for the Middle Miocene to Early Pleistocene of the East Carpathian Foreland. A detailed inventory is made of the existing fossil localities and their faunas, the spatio-temporal distribution is assessed and the resulting implications for Paratethys timescales are discussed. The inventory will form the starting point for an improved local mammal biostratigraphy and plans for publication of the results are underway.

149 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday plenary poster R5

NANOSCALE STRUCTURE OF DOLINASOREX GLYPHODON

(SORICINAE, MAMMALIA) TOOTH ENAMEL 1 1 1 R. Moya-Costa *, B. Bauluz and G. Cuenca-Bescós

1Aragosaurus-IUCA, Dpt.Earth Sciences, University of Zaragoza, 50009 Zaragoza, Spain

*[email protected]

Keywords: biomineral, iron, soricinae, Dolinasorex, enamel

Soricinae is a subfamily of mammals of the order that have red enamel in the cusps of their teeth. They live in humid habitats and have a fast metabolism. Studies of current species point that soricines have a complex structure of the enamel and that the red colour is due to the content of iron (Fe). Dolinasorex glyphodon is one species of giant and venomous soricine that was defined in the Early Pleistocene, TD6 level of the Gran Dolina site (Atapuerca, Burgos, Spain). The objective is to characterise the phase with Fe and the structure of the enamel in D. glyphodon.

A lower incisor of D. glyphodon, from the level TD6 of Gran Dolina site, was embed in resin and cut perpendicularly to the long . Then it was polished and carbon coated. The section was observed by field emission electron microscopy with secondary and backscattered electrons. Compositional images and chemical analyses were taken to localise the zone with highest Fe content. Next a thin slab (6µmx5µmx50nm) of this zone was extracted with a Dual Beam instrument (Helios 600), and subsequently it was observed by transmission electron microscopy (TEM Tecnai F30).

TEM images show that the enamel of D. glyphodon is composed by prisms of apatite that change of direction from the dentine to the edge, forming different layers and zones. Fe rich area in the enamel is an inner layer of the tooth. It is composed for elongated apatite crystals that are disposed perpendicularly to the long axis of the tooth and to the enamel edge. These apatite crystals are disposed forming “fans” were crystals are contained in the same plane but with different directions. Compositional images display that among the apatite crystals and more abundantly among the “fans” aggregates of round nanocrystalline particles of Fe oxides or hydroxides (1nm sized diameter) are observed. These particles form domains with similar orientations changing progressively their orientations in the space.

Acknowledgements. RMC has a grant from the Spanish Ministry of Education, Culture and Sport (FPU14/05528). This work has been supported by the MINECO Projects CGL2012-38434-C03-01 and CGL2013-46169-C2-1-P.

150 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday plenary poster R6

NEW DETERMINATION OF THE SPECIMEN OF MICROCHOERUS (OMOMYIDAE, PRIMATES) FROM SANT CUGAT DE GAVADONS (LATE EOCENE, NE SPAIN) 1 1,2 3 R. Minwer-Barakat *, J. Marigó , and S. Moyà-Solà

1Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, 08193, Cerdanyola del Vallès, Barcelona, Spain 2UMR 7207 CR2P - C.N.R.S., M.N.H.N., U.P.M.C.-Paris6. Département Histoire de la Terre, Muséum National d’Histoire Naturelle, Paris, France 3ICREA at Institut Català de Paleontologia Miquel Crusafont and Unitat d’Antropologia Biològica (Dept. BABVE), Universitat Autònoma de Barcelona, 08193, Cerdanyola del Vallès, Barcelona, Spain

*[email protected]

Keywords: Microchoerinae, Headonian, Ebro Basin, Iberian Peninsula

The species Microchoerus ornatus was described on the basis of a fragment of maxilla bearing P2-M3 from the Swiss locality of Mormont Entreroches, and further identified at Gösgen-Kanal, also in Switzerland. In the 1960s, Miquel Crusafont-Pairó first mentioned this species in Spain, assigning to M. ornatus a fragment of maxilla with two teeth that he interpreted as P4 and M1 from Sant Cugat de Gavadons (Late Eocene, Ebro Basin). Following that taxonomic ascription, some later works have considered that the paleogeographical range of this species extended far from Central Europe and included the Iberian Peninsula.

We have reexamined the single specimen found from this Spanish locality, providing detailed descriptions, measurements, illustrations and comparisons with other species. The two teeth preserved in this maxillary fragment, previously interpreted as P4 and M1, correspond in fact to P3 and P4. The large size of the specimen from Sant Cugat de Gavadons and some morphological features (particularly the simple paracone and the distinct hypostyle in the P3) allow distinguishing it from the holotype of M. ornatus. Moreover, the premolars from Sant Cugat also differ from those of the other known species of Microchoerus (M. erinaceus, M. wardorum, M. creechbarrowensis), mainly by the more developed protocone, hypocone and hypostyle. Besides, the studied teeth are clearly larger than those of M. wardorum and M. creechbarrowensis from Creechbarrow and slightly smaller than those of M. erinaceus from Hordle Cliff.

Therefore, the material from Sant Cugat de Gavadons shows clear differences with the type of M. ornatus and with all the other species of the genus, but its scarcity and the advanced wear of the two available teeth do not allow the description of a new taxon, so we ascribe this specimen to Microchoerus sp. This way, the known geographic distribution of the species M. ornatus, previously considered to include the Iberian Peninsula, is restricted to Central Europe.

This study has been supported by the Spanish Ministerio de Economía y Competitividad (CGL2014-54373-P) and by the Generalitat de Catalunya (2014/100604; 2014 SGR 416, GRC). JM is financed by Fondation Fyssen.

151 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday plenary poster R7

TEETH HISTOLOGY OF TWO EQUID SPECIES FROM THE

LATE PLEISTOCENE OF THE IBERIAN PENINSULA 1 1 2 1,3 C. Nacarino-Meneses *, X. Jordana , J. Madurell-Malapeira and M. Köhler

1Institut Català de Paleontologia Miquel Crusafont, Department of Evolutionary Paleobiology, 08193 Cerdanyola del Vallès, Barcelona, Spain 2Institut Català de Paleontologia Miquel Crusafont, Department of Neogene and Quaternary Faunas, 08193 Cerdanyola del Vallès, Barcelona, Spain 3ICREA at Institut Català de Paleontologia Miquel Crusafont, Department of Evolutionary Paleobiology, 08193 Cerdanyola del Vallès, Barcelona, Spain

*[email protected]

Keywords: teeth histology, incremental marks, enamel, dentine, Equus

The evolution of Equidae is a classical research topic in Paleontology. Most remarkable is the body size variation observed within this lineage along a general trend towards size increase (Cope’s Rule). Specifically, the genus Equus experienced important size changes during the Plio-Pleistocene period in Europe. Body mass is an important life history trait and, as such, under strong influence of the ecological conditions. It correlates with other life history traits, for instance the age at weaning, which is associated with the eruption time of the first permanent molar (m1) in mammals. The study of histological sections of this tooth has been widely used to reconstruct this and some other life history traits in extinct species.

Here, as part of a more general project aimed to reconstruct the life history strategy of European Pleistocene horses, we present the preliminary results obtained from the histological study of several permanent first lower molars of two different sized equid species from the Late Pleistocene of the Iberian Peninsula: Equus hydruntinus (180- 200 kg) and Equus ferus (≈ 350 kg). In addition, a slice of a lower m1 of the extant species Equus hemionus was prepared for comparison. Fossil material used in the present study comes from La Carigüela site, a Late Pleistocene cave located in Piñar (Granada, Spain), while the tooth of E. hemionus belongs to the collections of the Zoological Institute of Hamburg University (Hamburg, Germany). Histological sections of teeth were observed under polarized light to study the enamel and dentine tissues. Different incremental marks (daily, circa-weekly) were identified in all samples, allowing the estimation of dental histological parameters (i.e. daily secretion rate, repeat interval) and rates and time of teeth formation. The results obtained from this research have provided valuable information about the growth and pace of life of these fossil species of European equids.

152 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday plenary poster R8

SYSTEMATICS AND PHYLOGENY OF OLD WORLD

ACERATHERES (MAMMALIA, RHINOCEROTIDAE) 1 2 3 L. Pandolfi *, N. Handa and X. Lu

1University of Roma Tre, Department of Sciences, Largo S.L. Murialdo 1, 00146 Roma, Italy 2Shinshushinmachi Fossil Museum, 88-3 Kamijo, Shinshushinmachi, Nagano 381-2404, Japan 3Henan Unicersity of Chinese Medicine, Department of Base Medicine, 450046 Zhengzhou, China

*[email protected]

Keywords: Neogene, , systematics, "hornless" rhinoceros, paleobiogeography

Aceratheres are an extinct widespread group of rhinoceroses around Eurasia, North America and Africa during the Neogene, characterized by the absence of the nasal or frontal horn. The systematic position and the phylogenetic relationships of several genera and species identified as aceratheres remain debated or poorly known. Here we provide a systematic and phylogenetic revision of the Old World representatives of aceratheres at genus-level. A cladistic analysis (maximum parsimony analysis) based on 314 characters (214 craniodental, 100 postcranial) and scored from 35 taxa has been performed in order to identify the synapomorphies of the group and to highlight their inner phylogenetic relationships. Hyrachyus eximius, Tapirus terrestris, Trigonias osborni and Ronzotherium filholi were treated as outgroup. The strict consensus tree obtained from six most parsimonious trees showed the presence of two major clades: one groups taxa previously assigned to Elasmotheriinae together with the African Chilotheridium pattersoni and Turkanatherium acutirostratum; the other (Rhinocerotinae) groups taxa assigned to aceratheres sensu lato (such as Protaceratherium minutum and Dromoceratherium mirallesi), and members of Rhinocerotini (extant rhinoceroses and closely related species, Brachypotherium and Diaceratherium) and Aceratheriini (Alicornops, Aceratherium, Hoploaceratherium, Persiatherium, Acerorhinus, Chilotherium, Shansirhinus, Plesiacetatherium and Subchilotherium). A major clade composed of Persiatherium, Acerorhinus, Chilotherium, Shansirhinus, Plesiacetatherium and Subchilotherium was recovered; but, Aceratherium, Alicornops and Hoploaceratherium formed a separated group sister to Rhinocerotini. The early representatives of Aceratheriini have been reported from the lower Miocene of Eurasia and Africa, suggesting a complex origin of this tribe. Probably, increasing the number of characters and taxa in the analysis, particularly the new early African representative, could change the topology in several details.

153 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Friday plenary poster R9

PRELIMINARY REPORT ON THE DECIDUOUS PREMOLARS OF “SASHA,” THE FIRST INFANT WOOLLY RHINO (COELODONTA ANTIQUITATIS) TO BE DISCOVERED

W. Dirks1*, O. Potapova2, C. Witzel3, U. Kierdorf3, H. Kierdorf3 and A. 4 Protopopov

1Durham University, Dawson Building, Department of Anthropology South Road, Durham, DH1 3LE, UK 2The Mammoth Site of Hot Springs, SD, Hot Springs, SD 57747, USA 3University of Hildesheim, Department of Biology, Hildesheim, 31141, Germany 4Yakutian Academy of Sciences, Department of Mammoth Fauna Studies, Yakutsk 677007, Russia

*[email protected]

Keywords: hypoplasia, enamel, dentine, age determination

The world’s first frozen mummy of an infant woolly rhino was discovered in the Abyysky district of Yakutia, Russia in 2014, and given the nickname “Sasha.” The partial mummy is remarkably well preserved, with soft tissue, including skin and fur, as well as most of the skull and teeth. It has been dated to over 45,300 calBP, from the Karginsk interstadial of the Weichselian glacial. Our team is undertaking the analysis of the DP3 (slight occlusal wear) and DP4 (unerupted) These analyses will include determination of the age at death from histological sections, trace elemental analyses to determine the process of weaning, and isotopic analyses to determine season of death. Eruption order of the deciduous premolars in both black and white rhinos is DP3, DP2, DP4, DP1. The first deciduous premolar erupts at ~ 12-14 months. In Sasha, the order of deciduous premolar eruption appears to differ from that in the extant African rhinos. The DP1 is already in place but the dp1 has not erupted through the gingiva. Our preliminary investigations suggest that Sasha is much younger than a year. The position of the neonatal line, visible in the dentine of the DP4, is very close to the end of crown formation, indicating that most of the crown was formed prenatally. The root stock has partially formed but not yet divided, also indicating that the calf was quite young at death, probably only a few months old, which will be determined with histological analysis. Neither the enamel nor the dentine is fully mineralized. Hypoplastic enamel defects are visible in both analyzed deciduous premolars. Trace element analyses combining dental histology and LA- ICP-MS will determine the age at which the calf began to eat solid food, which can be as young as a week in extant rhinos. SEM observation revealed that the enamel is highly decussated, as was reported in previous studies of rhino enamel. Our analyses will add significantly to the understanding of the biology and ontogeny of woolly rhinos.

154 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary talks

THE SCIENCE OF ‘SOPHIE’ THE STEGOSAURUS: BUILDING A RESEARCH PROGRAMME AROUND A NEW ACQUISITION P.M. Barrett1*

1Natural History Museum, Department of Earth Sciences, Cromwell Road, London SW7 5BD, UK

*[email protected]

Keywords: Late Jurassic, body mass, feeding mechanics, palaeobiology, Stegosauria.

Stegosauria is a familiar dinosaur clade, but surprisingly little is known about its palaeobiology due to the scarcity of well-preserved skeletal material. The acquisition of a substantially complete, three-dimensionally preserved individual of Stegosaurus stenops by the Natural History Museum, London has enabled development of a research programme to investigate the anatomy and functional morphology of this taxon. This specimen (nicknamed ‘Sophie’) was collected in 2003 from the Late Jurassic Morrison Formation of Wyoming, USA and purchased for the museum in 2013. Its excellent preservation enables application of different modelling and imaging techniques to this taxon for the first time. Sophie provides new information on the anatomy of Stegosaurus, including better constraints on the position and number of dermal armour plates. In addition, the 3D preservation has allowed construction of accurate virtual models of the skeleton (generated using 3D-laser scanning and photogrammetry) that have formed the basis for studies on body mass estimation and locomotion. Convex-hulling was used to provide volumetric data that enabled body mass calculations with varying amounts of soft tissue reconstruction that, in turn, could be compared with estimates based on limb dimensions alone. Comparisons of these techniques highlighted the importance of ontogenetic stage in the application of body mass estimation methods and converged on a body mass of approximately 1,500 kg for Sophie. The skull is naturally disarticulated and almost complete, offering numerous insights into cranial arthrology and function. CT- scanning and retrodeformation of each element allowed construction of a three- dimensional craniomandibular model that was subjected to finite element (FE) and multibody dynamic analyses. These models show that Stegosaurus had a surprisingly high bite force in comparison with those of other herbivorous dinosaur clades, despite their reduced tooth morphology. Patterns of cranial stress and strain also differ between Stegosaurus and other ‘unspecialised’ dinosaur herbivores, such as therizinosaurs and basal sauropodomorphs, showing that anatomical and functional similarly do not occur in concert. Work continues on the specimen, with projects planned on locomotion and plate function. In addition to this research programme, Sophie has also become a focus for public outreach, raising awareness of palaeontology among the general public.

155 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

LATITUDINAL BIODIVERSITY GRADIENTS IN DEEP TIME: A CASE STUDY USING THE DINOSAURS OF THE MORRISON FORMATION, WESTERN USA S.C.R. Maidment1* and A.R. Muxworthy1

1Imperial College, Department of Earth Science and Engineering, South Kensington Campus, London, SW7 2AZ, United Kingdom

*[email protected]

Keywords: Dinosauria, Morrison Formation, latitudinal biodiversity gradient, sequence stratigraphy, magnetostratigraphy

The Morrison Formation is an ideal candidate for examining biodiversity distributions and population ecology at a time in Earth’s history when climatic conditions were different to today. It outcrops over 12 degrees of latitude, from Montana in the north to New Mexico in the south, making it possible to examine latitudinal biodiversity gradients at a time when temperature gradients between the equator and poles were significantly reduced. The Formation was deposited over about 10 million years, making it possible to examine high-resolution changes in diversity through time in terrestrial vertebrate communities. The intensive study of the Formation since the discovery of the Morrison’s diverse and well-preserved dinosaurian fauna in the latter part of the 19th century has generated a very large body of data that should allow diversity studies to be possible. However, attempts to use the Morrison for such studies are hampered by a lack of long-range correlation: the chronostratigraphy of the Formation is entirely unknown. Sedimentological logging and magnetostratigraphic sampling were carried out at 19 sites in order to use terrestrial sequence stratigraphy and magnetostratigraphy to produce a chronostratigraphic framework for the Formation. Terrestrial sequence stratigraphy divided the Formation into three sequences, comprising six systems tracts. The Formation youngs to the north, as would be expected as the Sundance Sea retreated. Magnetostratigraphic results were generally unsatisfactory due to magnetically weak samples with low intensities, the alteration of magnetic phases on heating, and poor sample coverage as a result of friable mudstones and unconsolidated sandstones. However, where magnetostratigraphic data has been recovered, it appears to support the chronostratigraphic framework based on sequence stratigraphy, and this framework is also supported by a number of smaller-scale studies in the literature. Sample- standardized dinosaur biodiversity for the two systems tracts which cover a broad latitudinal range indicate that biodiversity did not increase with decreasing latitude, in contrast to the pattern seen across the area today. This finding suggests that latitudinal temperature gradients may be a principal driver of today’s latitudinal biodiversity gradient, and significantly reduced latitudinal temperature gradients in the Upper Jurassic resulted in significantly different biodiversity distributions at that time.

156 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

THE PERMO-TRIASSIC DIVERSIFICATION OF ARCHOSAUROMORPHS

R.J. Butler1,2*, M.D. Ezcurra1,2,3 and R.B. Sookias1,3

1University of Birmingham, School of Geography, Earth and Environmental Sciences, Birmingham, United Kingdom 2Museo Argentino de Ciencias Naturales "Bernardino Rivadavia", Sección Paleontología de Vertebrados, Ángel Gallardo 470, C1405DJR, Buenos Aires, Argentina 3Ludwig-Maximilians-Universität München, GeoBio-Center, Munich, Germany

*[email protected]

Keywords: Archosauromorpha, macroevolution, mass extinction, Permo-Triassic, phylogeny

One of the key evolutionary events during the Triassic was the rise of archosaurs, the group that includes dinosaurs, birds, crocodylomorphs and pterosaurs. The Archosauria forms one component of a broader evolutionary radiation of diapsid reptiles, Archosauromorpha, but this broader context has often been ignored in analyses of Triassic faunal turnover events. Here, we present an overview of our recent and ongoing work revising and clarifying the diversity and phylogenetics of early archosauromorphs, and documenting and quantifying temporal and spatial changes in their morphological diversity and species richness. Archosauromorphs originated by the middle–late Permian (c. 260–255 Ma), but just four Permian species are currently known. However, their broad palaeogeographic distribution and phylogenetic interrelationships suggest that substantial archosauromorph diversification occurred before the end of the Permian, possibly in response to the end-Guadalupian extinction event. Archosauromorph fossil remains increase in abundance in the immediate aftermath of the Permo-Triassic mass extinction (PTME; c. 252 Ma), forming part of a short-lived, broadly distributed, earliest Triassic ‘disaster fauna’ with low morphological diversity. A major increase in archosauromorph morphological diversity, abundance, species richness and maximum body size occurred around the transition between the Early and Middle Triassic (Olenekian– Anisian boundary, c. 247 Ma), including the origin of large, hypercarnivorous erythrosuchids. This evolutionary event may reflect the post-PTME recovery and stabilization of global ecosystems, following the unstable ‘boom and bust’ world of the Early Triassic. The earliest archosaur body fossils, the sail-backed ctenosauriscids, occur at this time, with a broad palaeogeographic distribution. Phylogenetic relationships and footprints suggest that much of the Early Triassic diversification of archosaurs remains to be discovered. Cranial disparity of non- archosaurian archosauromorphs in the Anisian was notably high, and exceeded that of the crown archosaurs. However, non-archosaurian archosauromorphs declined in the early Late Triassic, coincident with the morphospace expansion and diversification of crown archosaurs. This turnover may reflect a competitive replacement. Only crown archosaurs survived into the Jurassic, going on to dominate the Mesozoic world.

157 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Hominins

DUBOIS: TRINIL, TEYLERS, TEGELEN J. de Vos1,2,3*

1Naturalis Biodiversity Center, Darwinweg 2, 2333 CR Leiden, The Netherlands 2Geologisch Museum Hofland, Hilversumseweg 51, 1251 EW Laren, The Netherlands 3Teylers Museum, Spaarne 16, 2011 CH, Haarlem, The Netherlands

*[email protected]

Keywords: Palaeoanthropology, Pithecanthropus, Homo erectus, Villafranchien, Dmanisi.

The Dutchman Eugène Dubois is the founder of our profession, palaeanthropology. Dubois was the first one person to search for hominin fossils on purpose, at the end of the 19thy century. He went to the Indonesian island Java, and, in September 1891, he found a third molar of a primate at Trinil, along the Solo River. The now-famous skullcap with a brain capacity just between Man and Ape was unearthed in October 1891. In August of the following year a third primate fossil was discovered, this time an almost completely preserved left femur. The femur is very human-like, and at first instance Dubois called it Anthropopithecus erectus. By then he realised that he had his transitional form and renamed it Pithecanthropus erectus, nowadays Homo erectus. There was a lot of discussion, but Dubois’ time was no different from today.

Dubois returned to the Netherlands in 1895. He became a Professor at the University of Amsterdam, and the Director of his own collection, the Dubois collection at the Rijksmuseum van Natuurlijke Historie (now part of Naturalis) in Leiden. In 1896, he became Curator of the Paleontological and Mineralogical Cabinet at Teylers Museum. The directors of Teylers Museum donated a safe for Dubois to store his precious fossils. Today, Teylers Museum still contains a collection of hominin casts collected under Dubois’ care.

Dubois continued to collect fossils, this time from Tegelen in the south of the Netherlands, and after which the Tiglian is called. The fauna from Tegelen consists of a Villafranchien fauna including Hypolagus sp., Mammuthus meridionalis, Stephanorhinus etruscus, Equus stenonis/robustus, Cervus rhenanus, Eucladoceros ctenoides, Hystrix etrusca, Ursus etruscus, Panthera onca gombaszoegensis and (Pachy)crocuta perrieri. This faunal list is almost similar to the one from Dmanisi, the site with the Homo erectus skulls. Also the age of Tegelen is more or less similar (1.7 Ma) to that of Dmanisi. Although he never said it, Dubois also hoped, according to Von Koenigswald, to find a Pithecanthopus skull in Tegelen. As the Dmanisi skulls show that it Homo erectus was associated with a Villafranchien fauna, who knows.

158 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

THE FIRST HOMO OUT OF AFRICA D. Lordkipanidze1*

1Georgian National Museum, 3 Purtseladze Street, 0105 Tbilisi, Georgia

*[email protected]

Keywords: Hominins, Pleistocene, Dmanisi.

This paper presents the case from Dmanisi (Georgia) where the earliest traces of humans that emerged out of Africa have been found. This case offers very comprehensive data, including well-preserved human remains, animal bones and primitive stone tools. This discovery has transformed our comprehension of the bio- geography of early Homo and revolutionized our understanding of their morphology, population, environment and behaviour. Dmanisi yielded a spectacular collection of skulls and skeletal bones from at least five individuals dating from 1.8 million years ago. The Dmanisi discoveries document the first expansion of hominins out of Africa and show that this was neither due to increased brain size, nor to improved technology. These discoveries include “Skull 5”, the most complete early Homo skull ever found. The variation in the five different skulls from the same place and time yields exciting new evidence on the evolutionary biology of early Homo and supports the idea of a single evolving lineage of early Homo.

159 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

DEFINING NEW BOUNDARIES? PALAEOLITHIC RESEARCH IN THE NETHERLANDS AND ADJACENT DOGGERLAND L. Amkreutz1* and G. Dusseldorp2

1National Museum of Antiquities, Papengracht 30, PO Box 11114 2301 EC Leiden, the Netherlands. 2Leiden University, Faculty of Archaeology, the Netherlands

*[email protected]

Keywords: Doggerland, Neanderthals, Palaeolithic, Pleistocene, North Sea.

Our knowledge of the Pleistocene human occupation of the Netherlands and its faunal context is limited. We present preliminary results of research efforts to supplement the traditional dryland data with data recovered from the North Sea. We try to integrate the terrestrial and marine archaeological records to arrive at a general overview of the Late Pleistocene occupation of the North Sea Basin.

Research into Pleistocene deposits aimed at documenting human occupation of the Netherlands goes back to the late 19th and early 20th century. In situ remains of fauna and artefacts remain elusive to this day, however. Recently, large scale excavations at quarry sites such as Maastricht Belvédère (NL) and Veldwezelt and Kesselt (B), provided valuable information on Neanderthal ways of life in the south of the Netherlands. Nevertheless, such research projects are rare and skew our interpretations to regional contexts with little faunal preservation, leading to an emphasis on lithics. In contrast to adjacent countries, Pleistocene deposits are generally not investigated under current rescue-archaeology regimes. Complementary evidence from the North Sea has come to light over the past decades. From the 1970s onwards artefacts and abundant faunal remains recovered from fishing nets sparked the imagination about this lost area in front of the Dutch coast. Much of the recovered faunal material appeared of Pleistocene age. In recent years sand supplementation on the beaches and large-scale infrastructural projects have added to this picture.

We give an overview of recent research into both ‘dryland’ and North Sea archaeology. We present preliminary results of a project currently headed by the Museum of Antiquities in Leiden inventorying collections of North Sea artefactual materials. We contextualise the known artefacts from the North Sea area with evidence derived from marine geological and prospective research and elaborate on the context and significance of a Neanderthal fossil from the North Sea. By trying to connect the maritime discoveries with the results of recent land-based research in both the Netherlands and the United Kingdom we attempt to arrive at a more complete picture of the waxing and waning of the Pleistocene occupation of north western Europe.

160 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

‘NATIVITY’ IN THE BONE BEDS OF BETHLEHEM

A.M. Lister1* and R. Rabinovich2

1Department of Earth Sciences, Natural History Museum, London SW7 5BD, UK 2Institute of Archaeology and Institute of Earth Sciences, Hebrew University, Jerusalem 91904, Israel

*[email protected]

The ‘bone beds’ of Bethlehem were excavated in the 1930s and 40s and yielded an important mammalian fauna, dominated by elephant remains, as well stone and bone objects initially interpreted as artefacts but later dismissed. Our new research into surviving excavation notes and plans, and other contemporary archives, allows us to contextualise the finds. Although the site no longer exists, current understanding of regional geology concurs with mammalian biostratigraphy (presence of Hipparion and very primitive elephant) in dating the accumulation, probably a karstic collapse structure, to at least 3 Ma. In light of the discovery of lithic industry in E Africa ca. 3.4 Ma, and the paucity of evidence from the Levant and SW Asia as a whole, the possible presence of humans in this region at the time of Bethlehem cannot be excluded. Re-study of the lithic assemblage is planned, but we have started with the fauna, especially the elephants, whose evolutionary and migrational history to some extent parallels that of humans. The family (Elephantidae) arose in Africa ca. 7 Ma, and the earliest mammoths (Mammuthus) and Asian elephant progenitors (Elephas) dispersed into Europe and S Asia, respectively, both at around 3.5 Ma. The Bethlehem molars are more primitive than any Eurasian members of either genus known until now. They are closest to stem African Mammuthus, but a tusk from Bethlehem has the untwisted shape of Elephas. This raises the possibility that the site includes the earliest members of both genera, ‘en route’ to Europe and southern Asia, respectively.

161 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

A GENERAL OVERVIEW OF PLEISTOCENE HOMININ REMAINS FROM TURKEY

A. İ. Aytek1*

1Mehmet Akif Ersoy University, Faculty of Arts and Science, Deparment of Anthropology, Istiklal Campus, Burdur, Turkey

*[email protected]

Keywords: Homo erectus, Homo neanderthalensis, Homo sapiens, Pleistocene, Turkey

Anatolia, thanks to its geographical location, has been taken place on the crossroads of three major continents and this feature enabled it to host hominin groups, and again due to the same feature, it served as a corridor and a bridge to the migrations throughout the history. Despite its geographic position, paleoanthropological research in Turkey has been limited, and therefore the known fossil human record from this region is quite sparse. Nevertheless, the Pleistocene human remains are likely represented. Furthermore, most of this fossil record suffers from a lack of proper description and documentation, as well as a dearth of information regarding its geological and chronological context. Although most of the known fossil human remains were found during early investigations, in the last decade new findings have further highlighted the region’s potential for paleoanthropological research. These researches reveal that at least three different human species - Homo erectus, Homo neanderthalensis and Homo sapiens - populated in Turkey during the Pleistocene period. The literature about the earliest Paleoanthropological research for the Pleistocene period is not kept in view and most of the fossils are not clearly known in scientific area. Therefore, this study aims to review the human fossil record from Turkey, their morphological features, finding spots and related literatures.

162 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

BIOMECHANICS AS A PREDICTOR OF ENAMEL THICKNESS IN EXTINCT PRIMATES

J. Marcé-Nogué1* , A. Dasch1, S. de Esteban-Trivigno2,3, P.González4, J. L. Cantalapiedra5 and T. M. Kaiser1

1Centrum für Naturkunde, University of Hamburg, Martin-Luther-King-Platz 3, 20146 Hamburg, Germany. 2Institut Català de Paleontologia, Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Spain 3Transmitting Science, Piera, Spain 4Instituto de Genética Veterinaria "Ing. Fernando N. Dulout". CONICET-FCNyM-UNLP, La Plata, Argentina 5Museum für Naturkunde, Invalidenstraße 43,10115 Berlin, Germany

*[email protected]

Keywords: biomechanics, primates, teeth, bite forces, lower jaw

Biomechanics and forces as well as properties of teeth are demonstrated to be crucial to understand diet, dental functional traits and evolution in primates. In this work, biomechanics of extant and extinct primate jaws during chewing were analysed using a classical two-dimensional lever approach. First, we aim to test for correlation between indicators of jaw biomechanical performance and geometrical and physical properties of the teeth with the diet. Second, we will predict the biomechanical behaviour and the diet of extinct taxa -such as hominoids and from other families of primates- from physical characteristics of the teeth and different biomechanical indicators respectively.

Lever models from mechanics were used to estimate muscular and biting forces as a load response in the models. As a comparative framework, some considerations were taken into account when building the biomechanical models such as orientation of the jaws or obtaining dimensionless parameters, etc. Correlations were tested using Ordinary Least Square (OLS) regressions and phylogenetically-informed regressions (PGLS). Geometrical and physical properties of the teeth were gathered from the literature.

In the extant data, our results show a strong coevolution between biomechanical parameters of the chewing mechanism with the teeth properties (especially mechanical advantage and geometrical ratios). Regarding the extinct species, one specimen of an extinct primate Oreopithecus bambolii was used to verify the correlations. The results show that this method is useful to predict enamel thickness among primates.

163 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

THE FIRST MODERN HUMANS OUTSIDE AFRICA

M.J. Sier1*

1Paleomagnetic Laboratory ‘Fort Hoofddijk’, Department of Earth Sciences, Faculty of Geosciences, Utrecht University, Budapestlaan 17, 3584 CD, Utrecht, The Netherlands

*[email protected]

In the early days of palaeoanthropology Asia played an important role that became more marginal with the research and discoveries in the African continent. However, in recent times new findings are putting back Asia into the human evolution headlines. Recently, we published evidence of Homo sapiens presence in China with a robust chronology between 80.000 and 120.000 years ago. The 47 human teeth found at the Fuyan (Daoxian) cave in South China can be unequivocally attributed to our own species and currently represent the earliest and soundest evidence of the presence of modern humans in China in the early-Late Pleistocene.There have been claims of Homo sapiens fossils in China with an early-Late Pleistocene age for some time. However, some doubts lingered due to the lack of precise chrono-stratigraphic context and/or an unclear taxonomic status, in combination with the reluctance of the scientific community to change longs-standing models. In this talk I will present the fossil and geological evidence of the Fuyan Cave and will briefly talk about how our views on modern human migrations have changed in the light of this new evidence.

164 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

REVISITING TRINIL: GETTING TO KNOW HOMO ERECTUS

J.C.A. Joordens1,2*, T. Simanjuntak3, R. Janssen2 and A. Verpoorte1

1Leiden University, Faculty of Archaeology, Einsteinweg 2, 2333CC Leiden, The Netherlands 2VU University Amsterdam, Faculty of Earth and Life Sciences, De Boelelaan 1-85, 1081HV Amsterdam, The Netherlands 3Pusat Arkeologi Nasional, Jalan Raya Condet Pejaten 4, Jakarta, Indonesia

*[email protected]

Keywords: Pleistocene, Java, shell, engraving, geochronology

The hominin Homo erectus is our most likely direct ancestor, and its former existence is known since its fossil remains were first discovered at Trinil on Java (Indonesia) in the 1890s. Trinil has not only yielded the first iconic remains of our extinct relatives, but also tens of thousands of other vertebrate and molluscan fossils that were excavated by Dubois and Selenka between 1891 and 1908. These collections are currently housed at Naturalis in Leiden (The Netherlands) and the Museum für Naturkunde in Berlin (Germany). Together, the Trinil site and collections constitute a treasure trove with an enormous, so far underexplored, potential to shed new light on the origins of the human lineage. This is evidenced by the worldwide excitement after recent discoveries in the Dubois Collection, showing that Homo erectus around half a million years ago collected freshwater mussels for consumption, used their shells for tool production and –most spectacularly- as a substrate for a geometric engraving. The latter suggests that Homo erectus may have been more “creative” than previously recognised, which evokes new questions on its ecology, behaviour and cognition. Moreover, the dating of Trinil shell infills, yielding a much younger age than expected, calls for a thorough reassessment of the age and depositional context of the Trinil site. To address these questions, we are planning to conduct new research at Trinil and in the collections. In this presentation we will give an overview of the recent findings from Trinil, results from collection research and the plans for a re- excavation of Trinil starting in summer 2017. We will share our excitement about these research prospects especially in the light of the recent discoveries from South East Asia, such as the earliest figurative cave paintings from Sulawesi and the redating of Homo floresiensis presence on Flores.

165 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

32 YEARS OF EXCAVATION IN PAŞALAR SITE TURKEY, THE RICHEST HOMINOID LOCALITY OF THE WORLD

B. Alpagut1*, S. Mayda2, T.T. Kaya3, A. Demirel4, L.W. van den Hoek Ostende5 , D. Kanık4, A. Şarbak6, G. Güler1, P.Joniak7, P. Pelaez-Campomanes8, S. D. Kesici9, K. Halaçlar3, M. Bilgin3, A. Tan3 and S. Karakütük3

1Ankara University, Faculty of Languages, History and Geography, Anthropology Department, Sihhiye-Ankara, Turkey. 2Ege University, Faculty of Science, Department of Biology, Bornova, 35100 Izmir, Turkey 3Ege University, Natural History Research and Application Center, Bornova, 35100 Izmir, Turkey 4Mehmet Akif Ersoy University, Faculty of Art & Sciences, Department of Anthropology, Burdur-Turkey 5Naturalis Biodiversity Center, PO Box 9517, 2300 RA, Leiden, The Netherlands 6Hittite University, Department of Anthropology, Faculty of Art & Sciences, Çorum, Turkey. 7Department of Geology and Paleontology, Comenius University, Mlynska dolina G, 842 15 Bratislava, Slovakia 8Museo Nacional de Ciencias Naturales, MNCN-CSIC, C/ José Gutiérrez Abascal, 2, 28006 Madrid, Spain 9 The Bodrum Museum of Underwater Archaeology

*[email protected]

Keywords: Paşalar, Hominoid, Middle Miocene, Turkey

The “Lignite Deposit Exploration in Turkey” project, conducted by a group of German and Turkish geologists, led to the discovery of the “Paşalar Hominoid Site” in 1969. The first excavations in Paşalar were carried out by H. Tobien and his team and the preliminary results of the faunal list, which included 47 micro and macromammalian taxa, were published. In 1983, the excavation restarted under the direction of Prof. Berna Alpagut on behalf of the Turkish Ministry of Culture and Tourism, Department of Antiquities, and the University of Ankara, Department of Paleoanthropology. In 1990, 55 taxa were reported, with some groups (carnivores, proboscideans) discussed briefly. At that time, around 1000 hominoid specimens were already recovered from the site and published in 1995 and 2008. These included the description of a new hominoid species, Kenyapithecus kizili. Up to this period, over 10.000 large mammal specimens have been identified and classified into 61 species. Between 2007-2012, as the excavations continued, Paşalar site attained its target: hosting the richest Miocene Hominoid collection of the world, encompassing more than 2055 Hominoid specimens. Here, we’d like announce the very first results of an extensive study by researchers from Turkey, The Netherlands, Spain and Slovakia, that has revealed the existence of 11 new taxa: Hyaenodontidae: Hyainailouros; Amphicyonidae: Cynelos, Amphicyon nov.sp., Pseudarctos, Gobicyon; Viverridae: Leptoplesictis; Hyaenidae: Protictitherium cingulatum, Protictitherium cf. crassum; Barbourofelidae: Sansanosmilus; Cervidea: Lagomeryx and Gomphotheriidae: Protanancus nov.sp. Hyainailouros, Pseudarctos, Gobicyon, Leptoplesictis and Lagomeryx are the first records from Turkey and particularly important as they will include new data and reveal peculiar paleobiogeographical connections and relations for the local and eastern Mediterranean biostratigraphy. Taking into consideration these biochronological data, we prefer to correlate the Paşalar fauna to early middle Miocene (late MN5). The palaeoecological data reflect a subtropical seasonal climate during the early middle Miocene (late MN5). In line with earlier studies, the surroundings encompassed some open areas.

166 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary talks

THE REPRODUCTIVE BIOLOGY OF PTEROSAURS

D.M. Unwin1* and D.C. Deeming2

1University of Leicester, School of Museum Studies, 19 University Road, Leicester, LE1 7RF, United Kingdom 2University of Lincoln, School of Life Sciences, Joseph Banks Laboratories, Lincoln, LN6 7DL, United Kingdom

*[email protected]

Keywords: pterosaur, Mesozoic, reproduction, embryo, superprecocial

Pterosaurs, Mesozoic flying reptiles, were first discovered more than 230 years ago, but next to nothing was known of their reproductive biology until quite recently. Several finds of pterosaur eggs, some with embryos and in one case preserved in association with the mother ('Mrs T', an example of Darwinopterus), have all been made in the last 12 years. Coupled with the identification, in several species (e.g. Darwinopterus, Pterodactylus, Pterodaustro), of near complete growth series (including hatchlings, juveniles and adults), these fossils have provided some key insights into reproduction and growth in these extinct fliers. The recent discovery that Mrs T is preserved in association with two eggs (rather than one, as originally thought) indicates the presence of paired oviducts as is typical for , but unlike birds which generally have only a single oviduct. Pterosaur eggs were relatively small and oval with pliable, fibrous shells. The latter features point to burial and incubation at ambient temperatures rather than contact incubation as in birds. Hatchling pterosaurs were, in many respects, like miniature versions of adults. Most notably, they had a well-developed flight apparatus and were ‘superprecocial’, probably capable of flying within hours or days of being born. Pterosaur reproductive biology thus appears to have been closely comparable to that of extant reptiles and quite unlike that of extant flapping fliers, birds and bats, with which they are otherwise convergent in many respects. Critically, the reproductive mode of pterosaurs suggests that their physiology was much more similar to that of reptiles than, as previously thought, to that of birds and bats. Interestingly, this idea is consistent with new reinterpretations of histological data (primarily for Rhamphorhynchus and Pterodaustro) which point to relatively slow growth rates in pterosaurs.

167 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

STUDY OF PRESERVED INTEGUMENTS FROM THE CRANIUM OF A TAPEJARID PTEROSAUR

A. Cincotta1,2*, P. Godefroit2 and J. Yans1

1University of Namur, Department of Geology, rue de Bruxelles 61, 5000 Namur, Belgium 2Royal Belgian Institute of Natural Sciences, Directorate ‘Earth and History of Life’, rue Vautier 29, 1000 Brussels, Belgium

*[email protected]

Keywords: taphonomy, integuments, pterosaur, SEM

The posterior part of the skull of the tapejarid pterosaur Tupandactylus imperator was collected in laminated limestones from the Early Cretaceous Crato Formation in northeastern Brazil. The supracranial crest is exquisitely preserved, including extensive soft-tissue impressions under the form of long blackish curved, parallel impressions. Moreover, small clusters of long thin light hair, or pycnofibers, are distributed along the posterior part of the skull. Scanning electron microscopy was performed on both blackish impressions and pycnofibers. Microbodies and imprints are observed on both tissue types. First, aggregates of oblong and rounded structures were observed within all samples collected from the crest. Further analyses by Energy Dispersive x-ray Spectrometry (EDS) reveal that these tiny corpuscles contain C, O, and Ca, as the diagenetic matrix. Pycnofibers also contain abundant microbodies, which are located on the surface and underneath. These structures are more elongated, thinner, and shorter than those from the crest. EDS analyses on pycnofibers show that the microbodies have a similar elemental composition than those on the crest. We suggest that both bacteria and melanosomes can be encountered together in the same sample. Indeed, microbodies found within pycnofibers show some organization, with bundles of similarly oriented structures, parallel to a thin overlying matrix. In the soft-tissue crest, microstructures are observed in three different arrangements: embedded in rhombohedral calcite crystals, which can suggest authigenic precipitation of calcium carbonate around them, or enclosed in an amorphous matrix, or as dense clusters of individual grains. As a consequence, oblong microbodies from the crest as well as elongate microbodies from the pycnofibers, all located within the integuments, are interpreted as probable fossil melanosomes. By contrast, dense and widespread corpuscles located on the surface likely represent autolithified bacteria. The presence of melanosomes in the crest would imply an epidemal origin for the soft tissue crest in pterosaurs: the supracranial crest of T. imperator was obviously formed by agglutinated and rigidified hair-like structures.

168 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

HIGH CONCENTRATIONS OF TURTLE EGGSHELLS (TESTUDOOLITHIDAE) IN AN EOCENE DELTAIC PLAIN (SOBRARBE DELTAIC COMPLEX, NORTHERN SPAIN).

M. Moreno-Azanza1*, R. Silva2, E.D. Berenguer3, J.I. Canudo3 and A. Badiola4

1Universidade Nova de Lisboa, Geobiotec. Dpto. de Ciências da Terra. Faculdade de Ciências e Tecnologia, FCT, 2829-526. Caparica, Portugal. Museu da Lourinhã 2Universidad de Zaragoza, Área de Estratigrafía, Dpto. de Ciencias de la Tierra, Facultad de Ciencias, Calle Pedro Cerbuna, 12, 50009, Zaragoza, España 3Universidad de Zaragoza, Grupo Aragosaurus-IUCA, Dpto. de Ciencias de la Tierra, Facultad de Ciencias, Calle Pedro Cerbuna, 12, 50009, Zaragoza, España 4Universidad del País Vasco (UPV/EHU), Dpto. Estratigrafía y Paleontología, Facultad de Ciencia y Tecnología, Apartado 644, 48080, Bilbao, España

*[email protected]

Keywords: vertebrate eggshells, taphonomy, Ainsa Basin, Southern Pyrenees, Lutetian

Here we describe an unusually high concentration of eggshell fragments from the Eocene Lutetian Sobrarbe Formation, in the Ainsa Basin, Huesca, Northern Spain. The eggshells were collected by screen washing of sediments from the Castejón de Sobrarbe 41 site, a sirenian dominated bonebed which has also provided several crocodilian and turtle specimens, including a complete carapace of a pleurodiran turtle. Sedimentological, taphonomical and paleoecological evidence supports that the accumulation was produced by the infilling of a tributary channel in the deltaic plain of the Sobrarbe Deltaic Complex. The eggshells are single layered, and are formed by subcilindrical shell units with radial ultrastructure. The outer surfaces are slightly sculptured, with compactituberculated ornamentation. The inner surfaces are cratered in most of the eggshell fragments. In thin sections, the eggshells are highly recrystallized, with its original composition completely lost. This contrasts with the apparent fine preservation of the ultrastructure under secondary electrons. All features are consistent with fossil and modern turtle eggshells, suggesting a very slow recrystallization of the original aragonitic eggshell, which preserves the fine morphology of the eggshells but not its crystal structure. The eggshells are assigned to the oofamily Testudoolithidae, and represent the third record of Eocene turtle eggshells in the world. Nevertheless, this scarcity is probably due to lack of sampling and the instability of their aragonitic composition. The high numbers of eggshell fragments, all of them assignable to the same ootaxon, and the absence signs of transport, suggests that the accumulation occurred in the proximities of a nesting area. Recent Pleurodira turtles are known to nest in different lentic and lotic environments within modern deltaic systems. Further prospection of the more proximal facies of the Sobrarbe Deltaic Complex may thus yield more complete specimens, including nests and clutches.

169 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

THE RISE AND FALL OF THE EUROPEAN CROCODYLIANS

M. Delfino1,2*

1Università di Torino, Dipartimento di Scienze della Terra, Via Valperga Caluso 35, I-10125 Torino, Italy 2Universitat Autònoma de Barcelona. Edifici ICTA-ICP, Institut Català de Paleontologia Miquel Crusafont, Carrer de les Columnes s/n, Campus de la UAB, E-08193 Cerdanyola del Vallès, Barcelona, Spain

*[email protected]

Keywords: Crocodylia, Cretaceous-Miocene, Planocraniidae, Diplocynodon, Crocodylus

If we restrict the meaning of the term Crocodylia to the crown Crocodylomorpha, the history of the European crocodylians can be traced back to the Late Cretaceous when non-crocodylian eusuchians and even non-eusuchians were still rather abundant and diversified. The discovery of new remains and redescription of old collections lead to the identification of several new species and even genera, and significantly improved the knowledge of the latter groups in the last 15 years. Conversely, the only presently recognized Cretaceous crocodylian from Europe is the gavialoid Thoracosaurus, confidently reported from the Maastrichtian but possibly present earlier. The K-Pg boundary marks the extinction of a significant portion of European non-crocodylian crocodylomorphs whereas Thoracosaurus was also retrieved from the Danian. Despite the rarity of outcrops, the fossil record indicates that besides gavialoids (Eosuchus and Thoracosaurus), also alligatoroids (Diplocynodon) and crocodyloids (“Asiatosuchus”) were already present in Europe during Paleocene. Later in the Eocene, the crocodylian faunas of Europe reached a diversification peak through the temporary occurrence of the extinct planocraniids, characterized by an altirostral snout (mediolaterally compressed and dorsoventrally deep), “ziphodont” teeth (labiolingually compressed and serrated), and hoof-like ungual phalanges. Tomistomine taxa inhabited Europe from the Eocene to the Miocene with Dollosuchus, Kentisuchus, Megadontosuchus, and Gavialosuchus/Tomistoma. However, the most representative European crocodylian is by far the endemic Diplocynodon whose origin is still quite enigmatic, even if relationships with North American Borealosuchus have been underlined. Diplocynodon shows a very substantial fossil record extending from at least the late Paleocene to the middle Miocene. Its remains have been cited or described from about 400 localities, and nine Diplocynodon species are currently considered valid (a few more are still waiting to be revised). In coincidence with the late Miocene Arctic glaciation, the crocodylian range experienced a marked latitudinal contraction and the last European remains were retrieved from late Miocene southern sites in proximity to the Mediterranean Sea. They are referable, in some cases on the basis of phylogenetically relevant characters, to Crocodylus, a newcomer that likely dispersed across the Mediterranean Sea from Northern Africa.

170 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Vertebrates of the Germanic Triassic basin and their Allies

THE EVOLUTION OF A UNIQUE BODY PLAN: NEW EVIDENCE FROM A TRIASSIC PLESIOSAUR SKELETON AND LONG BONE HISTOLOGY

P.M. Sander1*, T. Wintrich1, S. Hayashi2,3, A. Houssaye4, Y. Nakajima5 and T. Sato6

1University of Bonn, Division of Paleontology, Steinmann Institute for Geology, Mineralogy, and Paleontology, Nussallee 8, 53115 Bonn, Germany 2Osaka Museum of Natural History, Nagai Park 1-23, Higashi-Sumiyoshi-ku, Osaka, 546-0034, Japan. 3Osaka University, Division of Materials and Manufacturing Science, Graduate School of Engineering, 2- 1, Yamada-Oka, Suita, Osaka, 565-0871, Japan 4UMR 7179 CNRS/Muséum National d'Histoire Naturelle, Département Écologie et Gestion de la Biodiversité, 57 rue Cuvier CP-55, 75005 Paris, France 5The University of Tokyo, Atmosphere and Ocean Research Institute, 5-1-5 Kashiwanoha, Kashiwa-shi, Chiba, 277-8564, Japan 6Tokyo Gakugei University, Department of Astronomy and Earth Sciences, Faculty of Education, Koganei City, Tokyo 184-8501, Japan

*[email protected]

Keywords: Plesiosauria, Triassic, Bone histology

One of the major patterns in evolution is secondary aquatic adaptation. Several lineages of marine reptiles evolved during the Triassic as part of the ecosystem recovery after the end-Permian extinction. These include the familiar ichthyosaurs and the sauropterygians. The most diverse and best known clade of sauropterygians is that of the iconic plesiosaurs of the Jurassic and Cretaceous. Plesiosaurs had a unique body plan and mode of locomotion with the forelimbs and hind limbs developed as long and pointed flippers of the same shape that were used in paraxial movements for some kind of four-winged underwater flight. Except for some derived, large-skulled forms (“pliosaurs”), plesiosaurs bore their small head on a long, stiffened neck consisting of numerous vertebrae. Plesiosaurs are separated by a wide morphological gap from their closest relatives such as Bobosaurus and Pistosauridae, making their origins from Triassic stem sauropterygians and the origin of their paraxial locomotion unclear. The find of the first Triassic plesiosaur skeleton, from the Rhaetian of Germany (Bonenburg village, eastern North Rhine-Westphalia) provides new information on these issues. Like all later plesiosaurs, the new skeleton had reduced neck mobility and shortened zeugopodials (propodial/zeugopodial ratio >2.5). Phylogenetic analysis reveals the new skeleton also to be the most basal member of the clade. Long bone histology of the oldest and all later plesiosaurs is rather uniform and different from stem sauropterygians, suggesting the concurrent evolution of fast growth, an elevated metabolic rate (endothermy), and large offspring. These features help to explain the evolution of the unique plesiosaur body plan and mode of locomotion. The stiffened neck was an adaptation for rapid acceleration and sustained cruising but at the same time facilitated ambush predation on fish schools by providing visual and hydrodynamic camouflage. The shortened zeugopodials are part of the integration of all bones of the limb into a stiff hydrofoil, which was the prerequisite for rapid acceleration and sustained cruising. Plesiosaurian endothermy is thus consistent with two hypotheses for the origin of endothermy (aerobic scope hypothesis and parental care hypothesis) and may explain clade survival of the end-Triassic extinctions.

171 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

MUSCLE RECONSTRUCTION AND FINITE ELEMENT ANALYSES OF HUMERI IN PLESIOSAURS AND SEA TURTLES BY DISSECTION OF THE LATTER A. Krahl1*, U. Witzel1 and P.M. Sander2

1Ruhr-Universität Bochum, Faculty of Mechanical Engineering, Biomechanics Working Group, Universitätsstr. 150, 44801 Bochum, Germany 2Steinmann-Institut for Geology, Mineralogy and Paleontology, Division of Paleontology, Universität Bonn, Nußallee 8, 53115 Bonn, Germany

*[email protected]

Keywords: plesiosaur, , FE-analysis, muscle reconstructions

Plesiosaurs were globally distributed, highly aquatic marine diapsids. They had evolved a unique locomotory apparatus with four hydrofoil-like flippers that, for 135 Ma, was only subject to minor morphological changes. Long bones have traditionally been regarded as bending structures. Recent studies have shown that skulls are not bending structures but are loaded under compressive stress by tension chords, i.e., ligaments, tendons, and muscles. Reduction of the bending moment and a homogenous compressive stress distribution in all bony cross sections minimizes material and associated energy demands, which leads to lightweight constructions. Through comparison of humeri of an extant underwater flying reptile, Caretta caretta, with those of an extinct marine reptile, the plesiosaur Cryptoclidus eurymerus from the Middle Jurassic of England, we aim to apply this principle to long bones. Muscles were reconstructed for the Cryptoclidus eurymerus skeleton on display at the Goldfuß Museum, University of Bonn, Germany, using the extant phylogenetic bracket approach based on Testudines, Lepidosauria, and Archosauria. These muscle reconstructions were corroborated by studying osteological correlates in the fossil limb. For comparison, we studied the chelonioid locomotory apparatus to gain information on an extant underwater flying reptile. Dissection of a Caretta caretta took place at the Stazione Zoologica Anton Dohrn Napoli, Italy. Muscle forces were calculated for locomotory muscles spanning, inserting, and originating on the humerus. Agonistic and antagonistic muscles for the up- and downstroke in sea turtles and plesiosaurs were identified. Micro-CT scans of the right humerus of the plesiosaur specimen and of a Chelonia mydas from the Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany were produced. These micro-CT scans were post-processed and meshed with Simpleware ScanIP 5.1 to create a finite element (FE) model. FE models were further processed in ANSYS 12.0. Preliminary FE analyses show homogenous physiological compressive stress distributions for both the Chelonia mydas and Cryptoclidus eurymerus humeri. They demonstrate a shared structural optimization towards a lightweight construction of the skeletons in both taxa.

172 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

CRANIAL ANATOMY OF A NEW SPECIMEN OF YUNGUISAURUS LIAE AND TOOTH REPLACEMENT IN PISTOSAUROIDEA (DIAPSIDA, SAUROPTERYGIA) J.M. Neenan1,2*, T. Reich3 and T.M. Scheyer3

1Oxford University Museum of Natural History, Parks Road, Oxford, OX1 3PW, UK. 2University of Oxford, Department of Earth Sciences, South Parks Road, Oxford, OX1 3AN, UK. 3University of Zurich, Palaeontological Institute and Museum, Karl-Schmid-Strasse 4, 8006 Zurich, Switzerland

*[email protected]

Keywords: Pistosauroidea, Triassic, tooth replacement, cranial osteology

Pistosaurs are a rare grade of Middle-Late Triassic sauropterygians closely related to plesiosaurs, and are known from Europe, North America and China. Yunguisaurus represents the most complete and best-known pistosaur, with two almost complete specimens having been already described in recent years. Here we report a new specimen of Yunguisaurus (ZMNH M8813) from the Ladinian (Middle Triassic) Zhuganpo Member of the Falang Formation, Fuyuan, Yunnan Province, China, which also consists of a near-complete skeleton. Through the use of computed microtomography and traditional osteological study, we present a new interpretation of the cranial osteology of Yunguisaurus, as well as the first description of its dental structure and replacement patterns. Several osteological characters appear to differ with previously described specimens, including the position of the stapes, the relative size of the nasal and prefrontal, and a pineal foramen completely enclosed by the parietal. The highly specialised dentition of Yunguisaurus consists of a series of extremely elongate fangs on the premaxilla, maxilla and dentary that would have functioned as a basket to snare agile, smaller prey. These fangs have unusually elongate roots that, in the premaxilla, extend to the rostral midline; a condition that is also present in the closely-related Augustasaurus. Yunguisaurus displays an active tooth replacement cycle, with the majority of functional teeth being paired with a replacement. Indeed, replacement stages are bilaterally synchronous in Yunguisaurus, similar to pliosaurid plesiosaurs, and phases of replacement do not change between the maxilla and premaxilla. In contrast, Augustasaurus and Pistosaurus display far less tooth replacement, particularly in the premaxilla and maxilla, with replacement being unilateral, i.e. the left side being at a different replacement phase than the right. This is more similar to the basal sauropterygian condition seen in placodonts. Tooth replacement patterns represent a valuable phylogenetic character and are an important gap in the understanding of sauropterygian evolution. This work compliments recent publications on tooth replacement in placodonts and pliosaurid plesiosaurs, and is part of an ongoing project to document this process in the entire sauropterygian lineage.

173 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

UNUSUAL TAPHONOMY OF THE MARINE REPTILE BODY FOSSILS FROM THE LOWER ANISIAN OF WINTERSWIJK, THE NETHERLANDS J. Heijne1*, N. Klein1,2 and P.M. Sander1

1University of Bonn, Division of Paleontology, Steinmann Institute, Nußallee 8, 53115 Bonn, Germany 2Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1,70191 Stuttgart, Germany

*[email protected]

Keywords: Sauropterygia, Triassic, Muschelkalk, Germanic Basin

A unique taphonomic assemblage is found within the Winterswijk quarry (Vossenveld Fm., Lower Muschelkalk, lower Anisian) consisting of both marine vertebrate body fossils (mostly fish and sauropterygia) and unambiguous terrestrial reptile tracks. This quarry is located in the East of The Netherlands and is the north-westernmost outcrop of the Muschelkalk. The shallow marine Vossenveld Fm. and its fossil content are widespread, extending to the Teutoburg Forest in eastern North Rhine- Westphalia. The marine reptile fauna from Winterswijk consists mainly of isolated bones and partial skeletons of small pachypleurosaurs (Anarosaurus heterodontus), nothosaurs (Nothosaurus marchicus), Cymatosaurus sp., and several Placodontia. The current hypothesis regarding the taphonomy and depositional environment describes a tidal flat environment onto which carcasses and live animals were washed up by storm tides. However, this hypothesis has not been tested by taphonomic analysis of the skeletal material. Here we will present preliminary results regarding the most obvious taphonomic patterns observed in articulated and associated marine reptile skeletons. The study includes all previously described skeletons and new material from public and private collections. A striking first observation is the disarticulation pattern of the skeletons, which is unique to this locality. Despite dozens of partial skeletons being known, fully articulated and complete skeletons have not been found. Multi-taxon associations of more than one individual are known from at least one find. Although some fossils show a near- perfect articulation of some regions, other parts of the same individual are completely disarticulated. Isolated skulls, in particular of Nothosaurus, are common, but there are also partial skeletons with skulls. Isolated but articulated fore- and hind limbs are common as well. The specimens were documented using photogrammetry to assure future accessibility of taphonomic data. The patterns described here can be partially explained by the anatomy of the animals, as some bones were bound in vivo by strong tendons and ligaments. Other important factors affecting the disarticulation patterns were the environmental conditions such as sedimentation rate and current velocity. A further assessment of the sedimentological factors, as well as the documentation of additional patterns within the fossil assemblage, will be carried out in future research.

174 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

GEOCHEMISTRY OF SEDIMENTARY SEQUENCES FROM THE ANISIAN OF WINTERSWIJK WITH RELEVANCE TO UNDERSTANDING THE PALAEO-ENVIRONMENTAL LIVING CONDITIONS OF SAUROPTERYGIANS M.A.D. During1, J.J.L. van der Lubbe1, D.F.A.E. Voeten2, J. Pietersen1 and J.J.G. Reijmer1

1Faculty of Earth and Life Sciences, Vrije Universiteit Amsterdam, De Boelelaan 1085, 1081HV Amsterdam, The Netherlands 2Department of Zoology and Laboratory of Ornithology, Palacký University, 17. listopadu 50, 771 46 Olomouc, Czech Republic

*[email protected]

Keywords: Triassic, Isotope geochemistry, Sauropterygia, palaeo-environment, lithostratigraphy

The Winterswijkse Steengroeve quarry complex in the east of the Netherlands exposes a ~40 m thick sedimentary sequence comprising intertidal and shallow marine strata. The overall transgressive sequence contains a variety of microbially induced carbonates including laminated, argillaceous or bioturbated mudstones with several marlstone and dolomitised packstone beds. These deposits are assigned to the Vossenveld Formation, which corresponds to an interval of the Lower Muschelkalk Member of the Muschelkalk Formation. The Early to Middle Anisian strata were deposited in and alongside an epicontinental sea that covered large parts of Central and Eastern Europe.

Numerous beds within the succession preserve a shallow marine palaeofauna through a relative abundance of skeletal material from sauropterygians, fishes and crustaceans. Intertidal intervals exhibit numerous trackways of terrestrial vertebrates. The quarry has been subject of palaeontological and geological research for decades, although important questions considering the depositional environment and its ecosystem have remained unanswered. Here, we present a detailed lithostratigraphy supported by natural gamma ray analysis and stable carbon and oxygen isotope data of the sediment matrix from the Winterswijkse Steengroeve locality. Bulk sediment chemistry was analysed to gain insight in the environmental and depositional setting in which the described fauna were deposited and subsequently preserved post-mortem. Additionally, thin sections of distinct sedimentary facies were analysed to elucidate on the variations in depositional environments and on diagenetic modification of the sediments. Future stable isotope analyses on fossil dental and osseous material aims to test the geochemical preservation of the fossils and will additionally provide more information on dietary niches and habitat segregation of sauropterygians. To understand the depositional environment and diagenetic stability of the sediments and fossil material, it is essential to assess the environmental factors that likely contributed to the evolutionary success of the Sauropterygia in the recovery after the Permo-Triassic mass extinction event.

175 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

PRELIMINARY REPORT ON THE RE-DISCOVERY OF THE FORGOTTEN TRIASSIC LOCALITY FROM SILESIA (POLAND) WITH REMAINS OF TERRESTRIAL ARCHOSAURS Ł. Czepiński1

1University of Warsaw, Department of Palaeobiology and Evolution, Faculty of Biology, Biological and Chemical Research Centre, Żwirki i Wigury 101, 02-089 Warsaw, Poland

*[email protected]

Keywords: Archosauria, Silesia, Triassic

Remains of terrestrial vertebrates from the Triassic of Poland are the subject of ongoing studies. Localities in Krasiejów, Lisowice, Miedary and at other locations produce new fossil material every year.

Here we announce the rediscovery of the forgotten locality of Kocury (Silesia), possibly Late Triassic in age. In the second half of the 19th century, Kocury was investigated by the German botanist Georg Julius Ernst Gürich. There, he found remains that were later named Velocipes guerichi by Friedrich von Huene and thought to be a portion of a fibula of a theropod dinosaur. For a long time, the material was considered to have been lost during World War II. However, we were able to trace the specimen that turned out to have survived in the archives of the Geological-Palaeontological Institute and Museum of the University of Hamburg. Its re-evaluation along with other purported dinosaur remains from the present territory of Poland is currently in review. In addition, we also identified the original Velocipes Kocury locality and performed preliminary excavation works. This led to the collection of new remains of terrestrial archosaurs, including aetosaurian ; the first finds in Kocury since the 19th century.

The newly rediscovered locality will hopefully lead to future discoveries that contribute to a better understanding of the Triassic ecosystems of the Germanic Basin.

176 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

THE LATE TRIASSIC HERPETOFAUNA OF THE JAMESON LAND BASIN (EAST GREENLAND): REVIEW AND UPDATES M. Marzola1,2,3,4*, O. Mateus1,3, O. Wings5, N. Klein6, J. Mìlan7,8 and L.B. Clemmensen2

1Universidade Nova de Lisboa, GeoBioTec, Departamento de Ciências da Terra, Faculdade de Ciências e Tecnologia, Quinta da Torre, 2829-516 Caparica, Portugal 2University of Copenhagen, IGN, Department of Geosciences and Natural Resource Management, Øster Voldgade 10, DK-1350 Copenhagen K, Denmark 3Museu da Lourinhã, Rua João Luís de Moura, 95, 2530-158 Lourinhã, Portugal 4Geocenter Møns Klint, Stengårdsvej 8, DK-4751 Borre, Denmark 5Landesmuseum Hannover, Willy-Brandt-Allee 5, 30169 Hannover, Germany 6State Museum of Natural History Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany 7Geomuseum Faxe/Østsjællands Museum, Østervej 2, DK-4640 Faxe, Denmark 8University of Copenhagen,Natural History Museum of Denmark, Øster Voldgade 5-7, DK- 1350 Copenhagen K, Denmark

*[email protected]

Keywords: Late Triassic, Norian-Rhaetian, Fleming Fjord Formation, Jameson Land Basin, Testudines

The Norian-Rhaetian Fleming Fjord Formation (lacustrine and fluvial deposits) in the Jameson Land Basin (East Greenland) is rich in vertebrate fossils, recording all main groups of vertebrates known from the Late Triassic. Fishes, amphibians, a plethora of reptilians (including Testudines, Aetosauria, Phytosauria, Pterosauria, and Dinosauria), and early mammals compose the richness and completeness of the vertebrate record from this region of Greenland, explored with expeditions since the 1970’s. Thus far, only the temnospondyl pulcherrimus, the pterosaur Eudimorphodon cromptonellus, and the mammaliaform Haramiyavia clemmenseni have been subject of detailed anatomical and phylogenetic studies. Besides the reptilian and amphibian material known from the 90’s, which includes among others a skull, a basal testudine, two species of aetosaurs, and prosauropod dinosaurs, an expedition in 2012 collected a new testudine specimen and at least four individuals of The record of Testudines from the Fleming Fjord Formation seems particularly rich. At least three specimens have been found and are currently under study. The only already published specimen (MCZ field number 22/88G) was putatively described in 1994 as cf. Proganochelys; at that time the only well-known Late Triassic turtle in the world. This individual is the stratigraphically youngest of the three Greenlandic collected specimens (circa 208 Ma). Its association with Proganochelys was based on the pairs of gular and intergular projections and on the dorsal epiplastral process. Since the report, gular projections have also been described in Odontochelys, from the Late Triassic of China, making this character not autapomorphic for Proganochelys anymore. In addition, the dorsal epiplastral process is currently known both in Odontochelys and Kayentachelys, an Early Jurassic turtle from Arizona. The specimen collected in 2012 (NHMD-VP-9516) is older in geological age (about 209 Ma) and presents a unique imbrication pattern of the carapace bones: each costal overlaps the previous one up to 1 cm, and the peripherals constantly overlap the costals. A third previously unreported specimen collected in 1995 (NHMD-VP-2014) is yet undated, but possibly covers the gap between 209 and 208 Ma. Some of its well-preserved limb bones present distinct new characters among the known Late Triassic testudines.

177 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Vertebrates of the Triassic Germanic Basin and their allies poster R1

THE OCCIPITAL SKULL OPENINGS IN SIMOSAURUS AND NOTHOSAURUS (SAUROPTERYGIA, NOTHOSAUROIDEA), AND THEIR INTERNAL ASPECTS AS REVEALED BY DIGITAL IMAGERY T. Reich1*, D.F.A.E. Voeten2,3 and T.M. Scheyer1

1University of Zurich, Palaeontological Institute and Museum, Karl-Schmid-Strasse 4, 8006 Zurich, Switzerland 2Palacký University, Department of Zoology and Laboratory of Ornithology, 17. Listopadu 50, 77146 Olomouc, Czech Republic 3European Synchrotron Radiation Facility, 71 Avenue des Martyrs, 38000 Grenoble, France

*[email protected]

Keywords: Middle Triassic, Sauropterygia, endocast, eustachian tube, ossification

The Sauropterygia represent one of the most successful radiations of secondarily marine tetrapods in the history of life, which resulted in a remarkable morphological and ecological diversity. Lower and middle Muschelkalk deposits in the western Tethyan province yielded sauropterygian assemblages that document a rapid diversification of Triassic sauropterygians with different ecologies. Within the Nothosauroidea from the middle to early Late Triassic, the monospecific taxon Simosaurus is the sister-group to all other nothosaurs. Simosaurus is endemic to the Germanic Basin and pursued a different trophic strategy than Nothosaurus. Since the alpha taxonomy of nothosaurs is chiefly founded on skull morphology, a better understanding of their cranial osteology will help elucidate significant morphological adaptations to an aquatic lifestyle and the evolutionary history of these animals. Here we provide novel computed tomography data with focus on the occiput and braincase elements of several nothosaur skull remains to increase our understanding of three- dimensional endocranial structures and to assess their degree of ossification. The selected crania of Simosaurus and Nothosaurus were subjected to conventional microtomography (µCT) and Propagation Phase Contrast X-ray synchrotron microtomography (PPC-SR µCT). Virtual endocasts were produced to visualise the internal aspects of each skull opening and to reconstruct the embedded neurovascular canals. In posterior view, the skulls of both referred taxa exhibit three paired openings, which are formed by braincase elements adjacent to the foramen magnum. They once accommodated soft-tissues harbouring essential pathways of the nervous and circulatory systems, or passages such as the eustachian tube connecting the middle ear vault with the pharynx. The bone elements bordering some openings differ between the two genera, whereas an ontogenetic closure or reduction of certain foramina had been suggested for Nothosaurus. Our findings provide evidence for ontogenetic plasticity of some of the referred traits in Nothosaurus. The putative “eustachian foramen” does constitute the only paired ventral opening of the endocranial vaults, besides the internal nares. However, these foramina do not internally communicate with the dorsal endocast in Simosaurus, but do so to variable degrees in young specimens of Nothosaurus. Our findings suggest a secondary ossification of the eustachian passage in Nothosauroidea.

178 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Vertebrates of the Triassic Germanic Basin and their allies poster R2

EOSAUROPTERYGIAN REMAINS FROM THE MIDDLE TRIASSIC OF CANALES DE MOLINA (GUADALAJARA, SPAIN) C. de Miguel Chaves1*, F. Ortega1 and A. Pérez-García1

1Universidad Nacional de Educación a Distancia, Grupo de Biología Evolutiva, Facultad de Ciencias, Paseo de la Senda del Rey 9, 28040 Madrid, Spain.

*[email protected]

Keywords: Nothosauridae, Sauropterygia, Muschelkalk, Iberian record

The Spanish record of Triassic sauropterygians is mostly composed of isolated elements of placodonts, pachypleurosaurs, nothosauroids and pistosauroids. These remains represent teeth, vertebrae, ribs, osteoderms and appendicular elements, recovered from several Spanish regions (i.e. Aragon, Catalonia, Balearic Islands, Castilla-La Mancha and Andalusia). We present here several undescribed sauropterygian remains from the Muschelkalk of Guadalajara (Castilla-La Mancha, Spain).

These remains were discovered in 1980 near the locality of Canales de Molina (Guadalajara) in dolomitic limestones from the Middle Triassic of the Royuela Dolostones, Marls and Limestones Formation (Muschelkalk Facies). The vertebrate remains from Canales de Molina include teeth and scales of fishes, and vertebrae, osteoderms, teeth and appendicular elements of placodonts and eosauropterygians.

Several platycoelous vertebral centra with the “cruciform” articular surface that is characteristic for the Eosauropterygia were identified. Neural arches with different sizes and morphologies attributable to this clade have also been recovered. The proximal and distal fragments of a humerus are also preserved, as well as some femoral remains. Several characters observed in the vertebral centra and in the appendicular elements exhibit a similar condition as forms such as Nothosaurus and thereby permit attribution of the Canales de Molina remains to Nothosauridae. In addition, the teeth recovered from Canales de Molina are curved and have ornamented enamel, as is also characteristic of the clade Nothosauridae. The isolated remains from Canales de Molina correspond to several individuals of nothosaurs that range from small to medium size. The attribution of these elements to different taxa or to different ontogenetic stages of a single taxon is evaluated here.

179 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Vertebrates of the Triassic Germanic Basin and their allies poster R3

REMAINS OF SMALL VERTEBRATES FROM THE RÖT (LOWER TRIASSIC) OF GOGOLIN (OPOLE REGION, POLAND) K. Kardynal1,2*, J. Kowalski1,2, A. Bodzioch1 and M. Kowal-Linka3

1Opole University, Department of Biosystematics, Laboratory of Palaeobiology, ul. Oleska 22, 45-052 Opole, Poland 2Opole University, European Centre of Palaeontology, ul. Oleska 48, 45-052 Opole, Poland 3A. Mickiewicz University, Institute of Geology, ul. Maków Polnych 16, 61-606 Poznań, Poland

*[email protected]

Keywords: Upper Silesia, Röt, Triassic, reptiles, fishes

Accumulations of vertebrate remains discovered near Gogolin over the last years represent one of the oldest assemblages of Mesozoic reptiles known to date. Increased understanding of that ecosystem and its components will likely improve insight in the early Mesozoic evolution of several lineages. The remains occur in a thin littoral limestone/marl sequence just below the Röt/Muschelkalk boundary (Olenekian, ~247 Mya), where they were concentrated in at least three horizons deposited at the surface of cyanobacterial mats. The fossil assemblage is dominated by skeletal elements of fishes and reptiles, which are accompanied by unidentified bones and invertebrate remains. Although vertebrate skeletons are completely disarticulated, the bones show little evidence of abrasion that may indicate transport. A portion of the bones suffered brittle deformation from postdepositional compressive compaction, which complexifies reconstruction after chemical preparation (dissolution of rocks in acetic acid).

Fishes are represented mainly by ganoid scales, which morphologically correspond to the osteichthyan genus Paleoniscus sensu lato, and by teeth typical for the chondrichthian family Hybodontidae. Less common skull bones and other fish remains (e.g. vertebrae, fin rays, ribs), can sometimes be classified as the genus Saurichthys.

Reptilian remains are represented mainly by vertebrae (intercentra or neural arcs, sometimes articulated), ribs, long bones and isolated teeth, although fragments of skulls or pelvic and pectoral girdles have also been collected. Several species of the order Pachypleurosauria dominate the lower and middle bone horizons, where they are accompanied by other reptilian material, most probably from Ichthyosauria and . In the uppermost horizon, which forms the physical Röt/Muschelkalk boundary (but is still of Olenekian age), also nothosaurian bones and teeth occur in the assemblage.

Many of the remains are still problematic, but may include lariosaurian vertebrae, plagiosaurian scales and crushed reptilian eggshells.

180 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Vertebrates of the Triassic Germanic Basin and their allies poster R4

BIODIVERSITY OF SMALL VERTEBRATES FROM THE LATE TRIASSIC OF KRASIEJÓW J. Kowalski1,2*, A. Bodzioch1, P. Janecki1,2 and M. Ruciński3

1Opole University, Department of Biosystematics, Laboratory of Palaeobiology, ul. Oleska 22, 45-052 Opole, Poland. 2Opole University, European Centre of Palaeontology, Oleska 48, 45-052 Opole, Poland. 3Adam Mickiewicz University, Institute of Geology, ul. Maków Polnych 16, 61-606 Poznań, Poland.

*[email protected]

Keywords: Norian, fishes, amphibians, reptiles.

Investigations of Late Triassic (Norian) terrestrial vertebrates from Krasiejów have thus far mainly focused on large-bodied animals (temnospondyls and archosaurs), whereas remains of other groups (fishes, small amphibians and reptiles) have not received the same attention. Nevertheless, small-bodied fauna is usually more diversified and occurs in larger populations than larger taxa, which offers a broader foundation for the interpretation of palaeoecosystems. This, in turn, is very important for the reconstruction of the evolution of terrestrial faunal communities, in particular those observed at the end of the Triassic. For this reason, the “Opole Group” has paid more attention to search and study remains of small vertebrates in the last years.

Small skeletal elements (rarely larger than 1 mm) are concentrated by washing the sediment with clear water, after which the residue is manually separated using a stereo microscope. In this way, we acquire several thousand specimens yearly, and this still growing collection shows a highly diversified fossil assemblage.

Various kinds of teeth are the most commonly encountered skeletal remains. They represent fishes (chondrichthyan: hybodonts and possible xanacanths, dipnoans and actinopterygians), small (or juvenile) temnospondyls, lepidosauromorphs and archosaurs (including pterosaurs). Some of teeth show morphological similarities to cynodonts or early dinosaurs, and in some cases it is difficult to assign a teeth to any particular group of vertebrates.

Both ganoid and cycloid scales are very common, while fragments of bones are rare. So far, bones of fish skulls, fragments of sphenodonts jaws, and unidentified fragments of reptilian long bones have been found.

This preliminary reported assemblage of small vertebrates comprises both evolutionary old lineages and younger lineages that are typical for the fauna that developed after the Permian/Triassic crisis. It therefore seems that the Krasiejów palaeontological site contains a good record of the transition of terrestrial vertebrate communities during Late Triassic times.

181 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Vertebrates of the Triassic Germanic Basin and their allies poster R5

FISH SCALE DIVERSITY IN LATE TRIASSIC DEPOSITS OF KRASIEJÓW M. Antczak1* and A. Bodzioch2

1A. Mickiewicz University, Department of Palaeontology and Stratigraphy, ul. Maków Polnych 16, 61-616 Poznań, Poland 2Opole University, Department of Biosystematics, ul. Oleska 22, 45-052 Opole, Poland.

*[email protected]

Keywords: ganoid scales, cycloid scales, Dipnoi, intraspecific variation, ornamentation

Taxonomy of extinct fishes is mostly based on body shape and skeletal morphology. Although it sometimes considers skin coverage, this has not been regarded at species level. However, analysis of modern fishes has shown that scale morphology can indeed provide taxonomic separation for lower taxonomic ranks as well. Although intraspecific morphological variation among fish scales can be broad, several specific characters successfully distinguish species within one genus. Analysis of fossilized fish scales found in the Late Triassic deposits of Krasiejów (SW Poland) reveals that the external scale microstructure can also be employed as a taxonomic tool in the fossil record.

In fine-grained continental deposits of Krasiejów, small ganoid (Actinopterygii) scales and large cycloid (Dipnoi) scales were found. Within ganoid scales, two types of connections between individual scales were observed, which might represent morphological variation between two genera or two species. In the first model, a broad bar at one scale contacts the edge of the second scale. In the second model, a ‘tooth’ at a ridge on the scale surface communicates with an indentation at the edge of the adjacent scale. Cycloid scales are mostly recovered incomplete, although a collection comprising several dozens of scales at Opole University (UOPB1032- UOPB1082) permits reconstruction of the complete scale morphology. Distinct differences between cycloid scales are visible in the ornamentation of posterior and lateral fields. The distribution of certain sets of features (circulli shape, ‘lepidont’ characters, and lateral fields sculptures) suggests the presence of at least two scale morphotypes in the assemblage. Based on the variation found in modern , we conclude that these morphotypes may reflect intraspecific variation.

182 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary talks

THE TALE OF THE CROUCHING DINOSAUR. HOW DINOSAURS WERE MADE IN EARLY 20TH CENTURY GERMANY I.J.J. Nieuwland1*

1Huygens ING, Netherlands Academy of Arts and Sciences, Willem-Alexanderplein 5, 2595BE Den Haag, The Netherlands

*[email protected]

Keywords: Gustav Tornier, Otto Jaekel, popularization, sauropoda, History of science

The finds of large dinosaur remains in German East Africa in 1906 proved to be a game-changer in the German attitude towards the study of dinosaurs. Suddenly the scientific elites of a nation that was the world leader in scientific activity had to form an opinion about creatures it had never seriously occupied with.

Never was that better illustrated than when a large part of the German zoological and paleontological elites were pitted against one another, and against their American colleagues, over questions of sauropod anatomy after 1909. Although the discussion in scientific journals remained brief, the consequences for the popular image of dinosaurs in Germany were profound.

This episode has usually been depicted as a straight-out example of nationalist scientific rivalry between German and American élites (Desmond 1975, Parsons 2001 & 2003), but closer inspection reveals more intricate processes at work. Social, disciplinary and political competition all played their roles in provoking a number of prominent German scholars to re-think the whole body of literature on sauropods, and to use both scientific and popular channels for the promulgation of their conclusions.

In this presentation, I will show what caused these scientists to promulgate such an idiosyncratic (and often-ridiculed) image of sauropods, and how the German public was involved in this controversy. It throws a light on the relation between the scientific world and the public sphere, and the power of popularization, that is as relevant today as it was then.

183 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

PRELIMINARY SURVEY OF THE MACKEREL-LIKE FISHES FROM THE OLIGOCENE OF THE “GRUBE UNTERFELD” (S. GERMANY) K. Monsch1,2

1Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, the Netherlands. 2Hessisches Landesmuseum Darmstadt, Friedensplatz 1, 64283, Darmstadt, Germany

*[email protected]

Keywords: Mackerel-like fishes, ichthyofauna, Oligocene, Paratethys Sea, palaeoecology

The famous fossil locality Grube Unterfeld, near Rauenberg in South Germany is a famous fossil locality, which has in recent years produced many new fossils of various animal and plant taxa. There have been, most notably many fishes, amongst which a considerable number of mackerel-like fishes (Scombroidei, Perciformes). The exact taxonomic records of these are still poorly known, not only because most taxonomies are based on old reference collections and data, but also because the taxonomy of scombroid fishes is in a state of flux, for Recent as well as fossil taxa. Bringing the current interests in the Grube Unterfeld (also known as the “Frauenweiler clay pit”) and scombroid systematics together, resulted in a preliminary inventory of all scombroid specimens that are known from this locality. We have investigated these specimens in all German repositories where they are available. This preliminary inventory has been done in the first place to create a catalogue. As it stands now, there may be an estimated number of 28 species, several of which are new to science. The Grube Unterfeld locality is part of a larger palaeontological seaway the Paratethys Sea. The scombroid palaeofauna of Grube Unterfeld is similar (but not identical) to that of other Paratethys localities, including famous ones from the former USSR. Knowledge of the “Frauenweiler” palaeofauna will not only allow the reconstruction of the palaeoecology of that locality itself, it will also aid in reconstructing the history of the Paratethys Sea.

184 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

NEW AND RARE FOSSILS FROM THE LOWER OLIGOCENE OF THE GRUBE UNTERFELD (“FRAUENWEILER”) CLAY PIT N. Micklich1*

1Natural History Department, Hessisches Landesmuseum Darmstadt, Friedenplatz 1, D.64283 Darmstadt, Germany

*[email protected]

Keywords: Grube Unterfeld, lower Oligocene, Echeneidae, Leiognathidae, Chaetodontidae

The Grube Unterfeld clay pit is located in southern Germany, about 12 km south of Heidelberg, in the municipality of Rauenberg. Its sediments date back to the lower Oligocene (approx. 30.1 Mya) and were deposited at the bottom of a small sea strait that occasionally connected the ancient North Sea with the pre-alpine marginal sea at that time. The locality became famous for a rich fossil record, particularly so of fish. Some taxa, such as small herrings, are represented by numerous specimens in different preservation stages, but others are very rare very and only known from a few records.

Remoras (Fam. Echeneidae) are represented by only three records that all belong to a new genus and species, Oligoremora rhenana (Micklich et al., in press), which resembles in its meristic data the extant representatives of the genus Remora, but also shares several morphological details with the genus Echeneis. It differs from all other echeneids by the combination of its meristic data, as well as in several morphological details, some of which are rather unusual, at least for today's representatives of this family.

Ponyfishes (Fam. Leiognathidae) are represented by one record only. It shows some morphological traits that are typical for that family, but also differs from all extant and fossil leiognathids in the combination of several other character states, some of which are reminiscent of extant representatives of the family Gerreidae. According to this fossil record, there are some similarities with Leiognathoides altapinna (Weiler, in Hess und Weiler, 1955), a species that has been reported from contemporary localities in Switzerland and Russia. But there also are some deviations, which could be indicative of a different species.

Based on two specimens, the tholichtys larval stage of butterfly fishes (Fam. Chaetodontidae) was first described from the Grube Unterfeld clay pit in 2009. Such larvae are extremely rare in regular tropical plankton and midwater trawlings (0.1% of all captured larvae on average). Yet, two more of them were discovered in 2012. They differ from the former records in important morphological details and might even represent another taxon.

185 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

UNRAVELLING THE HISTORY OF A UNIQUE TRICERATOPS GRAVEYARD FROM EASTERN WYOMING, USA P. Kaskes1*, L.A. Portanger1 and A.S. Schulp1,2 1Vrije Universiteit Amsterdam, Department of Earth and Life Sciences, De Boelelaan 1085, 1081 HV Amsterdam, the Netherlands 2Naturalis Biodiversity Center, Darwinweg 2, 2333 CR Leiden, the Netherlands

*[email protected]

Keywords: Triceratops, Ceratopsidae, Lance Formation, bonebed, taphonomy

In August 2015, Naturalis Biodiversity Center (Leiden, the Netherlands) continued its excavations at a unique bonebed of ceratopsian dinosaurs south of Newcastle in Weston County, eastern Wyoming, USA. Preliminary field data shows these Upper Maastrichtian terrestrial deposits of the Lance Formation to hold at least five fairly complete skeletons of the genus Triceratops, ranging from juveniles to subadults and adults. Discoveries of Triceratops bonebeds are extremely rare and two previously described localities, in the time-equivalent Hell Creek Formation in Montana, amounted only three individuals. Therefore, this site marks the largest monospecific bonebed of Triceratops found so far.

Detailed geological fieldwork at the dig site and adjacent exposures was carried out to investigate the sedimentology and to develop possible taphonomic scenarios responsible for the burial of these horned dinosaurs. The Triceratops skeletons are associated with abundant micro- and macrofloral and -vertebrate remains, all incorporated within an organic-rich siltstone matrix. The remains are associated, but also show clear disarticulation. The fifth skeleton is located at the same site, c. 4 m above the main bonebed, implying that this specimen was buried at the same location thousands of years later. The preservation of multiple, ontogenetically different individuals within the same sedimentary horizon and in close proximity of each other suggests that these animals died together, but better geological understanding is needed to verify this scenario.

High resolution grain-size and thermogravimetric analysis of the bonebed and surrounding rock units can reveal the hydrodynamic and palaeoenvironmental conditions prior, during and after deposition of the bone accumulation. This way, it may be possible to differentiate between two post-mortem scenarios. The first hypothesis considers a fluvial origin for the bonebed in which the dinosaurs possibly accumulated on an overgrown pointbar after drowning. The second hypothesis suggests burial linked to potential trapping of the animals inside a swampy depression. Hence, unravelling the complex sedimentary origin of this Triceratops graveyard might provide new insights about gregarious behaviour –if any– of these famous horned dinosaurs.

186 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

ANATOMY AND STATUS OF BREVICERATOPS KOZLOWSKII – PROTOCERATOPSID DINOSAURS FROM THE LATE CRETACEOUS OF GOBI DESERT Ł. Czepiński1*

1University of Warsaw, Department of Palaeobiology and Evolution, Faculty of Biology, Biological and Chemical Research Centre, Żwirki i Wigury 101, 02-089 Warsaw, Poland

* \[email protected]

Keywords: Dinosauria, Ceratopsia, Mongolia, Cretaceous

Several species of ceratopsian dinosaurs are known from the Late Cretaceous sediments of Gobi Desert, including Protoceratops andrewsi, Protoceratops hellenikorhinus, Bagaceratops rozhdestvenskyi, Udanoceratops tschizhovi and Breviceratops kozlowskii. The last one was originally described by Teresa Maryańska and Halszka Osmólska in 1975 as ?Protoceratops kozlowskii. In 1990 Sergei Kurzanov coined a new genus for this species, Breviceratops, and referred additional material to it. His material afterwards was transferred to Bagaceratops rozhdestvenskyi, and since that time Breviceratops (=?Protoceratops) kozlowskii is considered a junior synonym of Bagaceratops rozhdestvenskyi.

I re-examined the holotype (ZPAL MgD-I/117) and reffered material (ZPAL MgD- I/116) of ?Protoceratops kozlowskii. Although both the holotype and referred material are likely juvenile individuals, they represent a protoceratopsid dinosaur distinct from both Protoceratops and Bagaceratops. Hence I suggest to resurrect the name Breviceratops kozlowskii as a valid genus and species, and propose a revised diagnosis for this taxon. Breviceratops can be distinguished from other protoceratopsid dinosaurs by an unique combination of plesiomorphic and derived features of the skull. Its stratigraphic distribution and relations with other Protoceratopsidae may shed a new light on the paleobiogeography and evolutionary history of this group of ceratopsian dinosaurs as well as biostratigraphy of the Late Cretaceous formations of Gobi Desert.

187 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

THE APPENDICULAR SKELETON OF PROTOCERATOPS ANDREWSI – ITS ONTOGENETIC CHANGES AND IMPLICATIONS FOR LOCOMATION J. Slowiak1*

1University of Warsaw, Department of Palaeontology, Żwirki i Wigury 93, 02-089 Warszawa, Poland

*[email protected]

Keywords: Protoceratops, ontogeny, locomotion, Mongolia

Protoceratops andrewsi is a well-known ceratopsian dinosaur form the Djadokhta Formation (Upper Cretaceous, Mongolia). Discovery of numerous (including fully articulated) skeletons of this dinosaur provides an excellent opportunity to study the ontogenetic and intraspecific variation in this species. My work is focused on the appendicular skeleton of P. andrewsi. Morphological observations of changes in bones shape during ontogeny, limb measurements, and histological data bring new information about the lifestyle of this basal neoceratopsian. It seems that the juvenile individuals had been bipedal and further in life P. andrewsi becomes quadrupedal. The results are then compared with other known taxa of basal Neoceratopsia (of various locomotor adaptations) and Psittacosauridae (obvious bipeds) giving us insight into the evolution of postural change in ceratopsian dinosaurs which evolved from the early Cretaceous bipedal forms and gave rise to large, solely quadrupedal Ceratopsidae.

188 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

BRAIN SIZE EVOLUTION IN CAVE BEARS (URSUS SPELAEUS) K. Veitschegger1* and M.R. Sánchez-Villagra1

1University of Zurich, Paleontological Institute and Museum, Karl Schmid-Strasse 4, 8006 Zurich, Switzerland

*[email protected]

Keywords: EQ, Carnivora, Ursidae, fat storage, diet

Cave bears were a widespread Eurasian species until they went extinct close to the end of the Pleistocene. The basic anatomy of their brains has been described based on natural as well as virtual endocasts. However, their relative brain size has not been quantified and compared in a phylogenetic and temporal context. Cave bear skulls are often recovered complete, so the volume of the brain can be accurately measured with glass beads. Here, we present a comprehensive dataset of encephalization quotients (EQ) from 99 cave bear skulls representing 19 localities. Additionally, the ancestral cave bear species Ursus deningeri is represented in the dataset. For a phylogenetic comparison, 185 skulls from all 8 extant bear species (Ailuropodinae, Ursinae, Tremarctinae) were measured. Cave bears secondarily decreased their relative brain size early on in their evolutionary lineage. Already U. deningeri had a considerably smaller EQ than all extant bear species. The American black bear, U. americanus, has the smallest EQ among all extant species. Cave bears and U. americanus exhibit two distinct characteristics, which have been shown to negatively influence brain size. Both species hibernate and share a more herbivorous (U. americanus) or very likely completely herbivorous (U. spelaeus) diet. Based on the study of extant mammalian species, it has been shown that a reduced quality of food and a high amount of fat storage have a negative trade-off with brain size. In cave bears the combination of these characteristics could have had considerable effect on the brain size.

189 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

LATE PLEISTOCENE GROUND SLOTHS OF THE YUCATAN PENINSULA, MEXICO S.R. Stinnesbeck1, E. Frey1, J. Aviles Olguín2, W. Stinnesbeck3, A.González González4, E. Aceves Núñez2 and C. Rojas Sandoval5

1Department of Palaeontology and Evolution, State Museum of Natural History Karlsruhe, Erbprinzenstr. 13, 76133 Karlsruhe, Germany 2Instituto de la Prehistoria de América, Puerto Juarez, Carretera Cancún -Tulum Km 282, Solidaridad, 77710 Playa del Carmen, Q.R. Playa del Carmen, Mexico 3Institut für Geowissenschaften, University of Heidelberg, Im Neuenheimer Feld 234, 69120 Heidelberg, Germany 4Museo del Desierto, Carlos Abedrop Dávila 3745, Nuevo Centro Metropolitano de Saltillo, 25022 Saltillo, Coahuila, Mexico 5Instituto Nacional de Antropología e Historia (INAH), carretera federal 307, km 128, 77710 Tulum, Quintana Roo, Mexico

*[email protected]

Keywords: Ground sloth, Pleistocene, Yucatan peninsula

Ground sloths are characteristic and enigmatic faunal elements of the Americas with a wide dispersal during most of the Neogene. In contrast to the extensive fossil record from the Caribbean and North- and South American localities, only few remains, majorly isolated single bones, have been described from Mexico.

The submerged cave system of the northern Yucatan Peninsula (YP) reveals a diverse megafaunal assemblage from the Pleistocene-Holocene transition. One part of this assemblage are giant ground sloths of a partially supreme preservation. Several skeletons of Nothrotheriops shastensis, which was widespread throughout Mexico, have been found in the caves. In the El Zapote sinkhole near Puerto Morelos in the Mexican state of Quintana Roo, Mexico a large megalonychid was discovered that was endemic to Mexico. It differs significantly from all known Megalonychidae, including those from the Greater Antilles and South America. The new ground sloth findings from the YP also reveal a coexistence with humans for the latest Pleistocene and, possibly, the early Holocene, as indicated by cut markings in several bones.

190 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

PREDOMINANCE OF PENTADACTYLY IN SPRAWLERS S.B. Kümmell1*

1Institute of Evolutionary Biology and Morphology, Center for Biomedical Education and Research, Faculty of Health, School of Medicine, University Witten/Herdecke, Stockumer Straße 10, 58454 Witten, Germany

*[email protected]

Keywords: Pentadactyly, Synapsida, sprawlers, autopodial rotation

Most extant animals that move as sprawlers have five or four digits, e.g. most lacertid, testudine and amphibian sprawlers and Crocodylia. In extant parasagittal animals, reduction to less than five digits is commonly seen. Artiodactyla often show only two digits, Perissodactyla four to one. Among fossil Synapsida, the time at which parasagittality came about is still under debate, but all authors agree that at least Theria of the Late Cretaceous were parasagittal. All known Synapsida until the end of the Cretaceous were pentadactyl in manus and pes with two exceptions: Cistecephalus and Fruitafossor, both of which were highly fossorial and not fast runners.

When walking with abducted limbs, the trunk undulates around the fixed autopodium inducing rotation in the zeugopodia and autopodia. This rotation can be accommodated in the autopodium through certain movements in the autopodial joints, during rolling or through horizontal rotation on the substrate. This is called autopodial rotation.

Autopodial rotation differs in different groups of sprawlers. In the sprawling gait of Crocodylia, it is done mainly by the metapodium, in which each of the more lateral metatarsale is flexed relative to the more medial during the propulsion phase. In fossil Synapsida, autopodial rotation takes place through medial abduction and rotation in the autopodial joints and through rolling around the oblique axis through the distal heads of the metapodialia I-IV. After the lift-off of the metapodium, the lateral digits extend before the medial. These forms of autopodial rotation are not possible with less than four digits.

Many digits and high manipulative abilities require a lot of muscles, making the autopodium heavy. Small animals have high muscle forces in relation to their body mass, whereas in large terrestrial parasagittal animals, reduction of digits and manipulative behavior facilitates a fast and sustainable locomotion. In terrestrial sprawlers, this evolutionary pathway of digit reduction to less than four is not possible, because of the constraints of autopodial rotation. In Mesozoic Mammaliamorpha, with their fully parasagittal or only partly abducted limbs, reduction in digit number was not likely, because of their small size and their often scansorial or arboreal lifestyle.

191 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

FIRST RESULTS OF THE SCIENTIFIC EXCAVATION NETWORK IN THE CLAY PIT "UNTERFELD" (OLIGOCENE) AT RAUENBERG (GERMANY) - ABUNDANCE AND DIVERSITY K. Eck1*, K. Hatsukano1, E. Frey1, W. Stinnesbeck2 and N. Micklich3

1State Museum of Natural History, Erbprinzenstraße 13, 76133 Karlsruhe 2Institute of Earth Sciences, Heidelberg University, Im Neuenheimer Feld 234, 69120 Heidelberg, Germany 3Hesse State Museum Darmstadt, Friedensplatz 1, 64283 Darmstadt

*[email protected]

Keywords: Diversity, Germany, Paleoecology, , Unterfeld

In addition to a variety of invertebrates and fossil plants, the clay pit "Unterfeld" provides a broad diversity of vertebrate taxa. Hitherto more than 70 species of fishes, nine species of birds, two species of sea turtles, a sirenian and a creodont have been described. In 2014 we started a project financed by the Klaus Tschira Foundation to compile the information given and to conduct a scientific excavation to finally carry out an investigation on the paleoenvironment of this site.

During the first two years of the project we registered more than 9500 specimens that are housed in the collections of the five institutions: Institute of Earth Science/Heidelberg University, State Museum of Natural History Karlsruhe, Hesse State Museum Darmstadt, Senckenberg Research Institute Frankfurt and State Museum of Natural History Stuttgart. The registration of the specimens in a commented inventory catalogue is finished. Here, we present an overview of the abundance and diversity of the taxa from the clay pit "Unterfeld" in the collections. Additionally we provide first results of the excavation campaign 2015 in terms of quantity and diversity of the taxa collected in the field in comparison with the collection material. Drill core analyses yield first information on environmental proxies. These new data allow a first reconstruction of the palaeoecological environment of the "Unterfeld" during the Rupelian.

192 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

North Sea Basin

THE NORTH SEA: A TREASURE TROVE FOR PLEISTOCENE VERTEBRATE PALAEONTOLOGY AND ARCHEOLOGY D. Mol1*

1Natural History Museum Rotterdam, Westzeedijk 345, 3015 AA Rotterdam, The Netherlands

*[email protected]

Although the woolly mammoth is certainly the most well-known icon from the Ice Age, he was not the only inhabitant of that era. The woolly rhino and the saber-toothed cat roamed Northern Europe as well, in the shade of their mammoth companions. These species lived in the lowlands between the British Isles and the Netherlands, and their fossil remains are uncovered on a daily basis. During the Ice Age, the bottom of the North Sea used to be a vast plain, a delta of the River Meuse and the Thames, which we recognize today as the Mammoth Steppe.

Over the last few decades, close collaboration between the fishermen of the North Sea and scientists has resulted in a complete overview of what used to be the vast plains of the Ice Age. The flora and fauna of that time are now fully mapped and still educate us today.

Apart from the animal remains, it is evidently clear that man used to be part of the Mammoth Steppe. Until about 8,000 years ago, when the North Sea reached its current sea-level, man lived in these lowlands.

In this report I will not only discuss the paleontological facts, but will also deal with the archeological value of this part of Northern Europe, in which man was an important player. Middle-Paleolithic artifacts from several sites of the plains of the North Sea, such as remarkable axes, will be on display for the first time in history. These sites can now be identified for certain as the home of the Neanderthals.

Furthermore, this account will not only deal with the Middle-Paleolithic component of the North Sea. Artifacts from the Mesolithic Age will also be discussed, as well as Mesolithic human remains. Moreover, several Neolithic axes provide proof of transport routes between Great Britain and the European mainland during the Ice Age.

193 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

AN UPDATE ON MOSASAUR ‘LARS’ FROM THE TYPE AREA OF THE MAASTRICHTIAN STAGE, SOUTEAST NETHERLANDS J.W.M. Jagt1*, L. Barten and J. Barten

1Natuurhistorisch Museum Maastricht, de Bosquetplein 6-7, 6211 KJ Maastricht, the Netherlands

*[email protected]

Keywords: Mosasauridae, Late Cretaceous, preparation, subadult individual

Currently, the partial mosasaur skeleton recovered from Emael Member (Maastricht Formation; late Maastrichtian) at ENCI-Heidelberg Cement Group quarry (Maastricht), is the main attraction in the new Sciencelab at the museum. Since the opening of this permanent laboratory (January 16, 2016), volunteers and museum staff have been preparing two principal pieces. One is large chunk of matrix, of around 90 kg, that has yielded fairly numerous flipper bones and two pieces of a pterygoid jaw (with two teeth, one preserved in situ). Of note is that at least four of these flipper bones show healed lesions, with scar tissue, indicative of some kind of encounter during life. The main piece (‘skull block’) has now been completely freed from its protective plaster jacket and framed in so as to avoid diagonal cracks to develop further and lead to bone damage. A partial upper jaw (left) is exposed, the right jaw apparently being displaced and situated underneath; the premaxilla has not yet been recognised. The lower jaws are also documented, but these have suffered considerable damage during lifting and transportation, although all broken pieces have been salvaged. More of the skull appears to be sitting in this block; on one end, at least two cervical vertebrae have now been exposed. To date, we have not seen any elements, or quadrate(s), but all bones now freed from the matrix clearly show this to be a subadult individual of hoffmanni, the commonest mosasaur taxon in the area. Volunteers and museum staff working at the Sciencelab (on regular working days [except for Mondays] and during weekends) wear headsets to allow them to communicate with the public. Visitors can use four cameras to zoom in on specific parts of the blocks under preparation, and on another screen videos are shown of the quarry, highlighting several issues of fossil content, the history of the name ‘Maastrichtian’ and mosasaurs in general. At a later date, more detailed information on the mode of life of mosasaurs, their skeletal structure and other anatomical details, will be made available to the public.

194 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

A DUTCH 'CRYSTAL PALACE' - MOSASAUR, TURTLE AND DINOSAUR IMAGERY IN SUBTERRANEAN GALLERIES, SOUTHERN LIMBURG (THE NETHERLANDS) E.W.A. Mulder1, E.A. Jagt-Yazykova2, J.W.M. Jagt3 and G.J. Boekschoten

1Museum Natura Docet/Wonderryck Twente, Oldenzaalsestraat 39, 6211 KJ Denekamp, the Netherlands 2University of Opole, Laboratory of Palaeobiology, Oleska 22, 45-052 Opole, Poland 3Natuurhistorisch Museum Maastricht, de Bosquetplein 6-7, 6211 KJ Maastricht, the Netherlands

*[email protected]

Keywords: Reptiles, sculptures, charcoal

At the municipal subterranean galleries of the Gemeentegrot Valkenburg (Cauberg, Valkenburg aan de Geul, the Netherlands) which we visited last in February 2014, there are some marvellous charcoal drawings and sculptures (marked* below) of several Late Cretaceous tetrapods, namely the dinosaurs Triceratops flabellatus* (by Jos. Eijmael, 1921) and Megalosaurus* (by Jean Caelen, 1895), the plesiosaur Elasmosaurus platyurus, the pterosaur Pteranodon ingens and the mosasaurs Tylosaurus dyspelor and Mosasaurus (by Tielens, no date) and Plioplatecarpus* (by Jean Caelen, 1896), as well as a marine turtle*. On the whole, the sculptures are both overly massive, with enormous scales on backs, bellies and extremities and too many teeth in upper and lower jaws, but also show remarkably much detail. The rock, a rather fine-grained biocalcarenite of late Maastrichtian age, in which these sculptures were hewn, is conducive to such detail being pictured. All in all, they are reminiscent of dinosaurs at Dinosaur Court, London (Crystal Palace). At some point, Benjamin Waterhouse Hawkins must have been a source of inspiration for the local artists, but how exactly they learnt of his work is unknown. A picture postcard, stamped in 1921, illustrates a closely comparable turtle, as well as a . Undoubtedly, the latter was intended to represent a mosasaur of some sort, clearly demonstrating that the true nature of this extinct marine monitor lizard was not familiar to all artists at the time. The sculptures featuring on this postcard were made by Jules Sonderijker (1865-1950) of Maastricht. The turtle (at the so-called Zonneberg Museum), damaged beyond repair by vandals, has now been excavated completely. Of the crocodile it appears that only photographs survive (T. Breuls, pers. comm.). At the northwesterly edge of the Sint-Pietersberg is the Schark War Monument (www.maastrichtunderground.nl), which is accessible via the ENCI- Heidelberg Cement Group quarry. In these galleries another sculpted and painted mosasaur image is currently accessible. This will be visited during the EAVP field trip on Sunday, July 10, 2016.

Acknowledgements – Kevin Amendt, Ton Breuls, Harry Habets, John Hageman, Peter Jennekens and John Vincken for either accompanying us on site or for supplying additional information.

195 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Hominins poster V1

SMALL BIRDS OF THE TD10 LEVEL OF GRAN DOLINA: NEW DATA ON THE MIDDLE PLEISTOCENE OF ATAPUERCA (SPAIN) C. Núñez-Lahuerta1*, J. Galán1 and G. Cuenca-Bescós1

1University of Zaragoza, Department of Earth Sciences, Pedro Cerbuna 12, 50018 Zaragoza, Spain

*[email protected]

Keywords: Birds, Middle Pleistocene, Atapuerca sites

The Gran Dolina (TD) is one of the numerous paleontological sites of the Sierra de Atapuerca karst complex (Burgos, northwestern Spain), which are known by their exceptional Quaternary stratigraphic and paleontological record. It has a 19m thick Lower and Middle Pleistocene infilling, which is divided in eleven lithostratigraphic units (from TD1 at the bottom to TD11 at top). This work is focused on the Level TD10, which has approximately 4m thick and is also divided in four sub units (from TD10.1 to TD10.4). The level is a succession of debris flow facies from the main entry of the cave with little inputs from lateral secondary entries. The dates range from 458±39ka to 244±26ka. The level shows a high diversity of macro and micromammals, also, anthropogenic damage affecting to ungulates, carnivores, rabbits and birds (both cut marks and bone breakage) were found.

In this work the avian remains of the level TD10 are studied. The specimens have been recovered only by concentrating the sediment with by washing and sieving the sedimentary materials acquired from the excavations. Thus, the analysed remains correspond mostly to the smallest of all birds of the ecosystem. Only the bird remains from the TD6 level (Homo antecessor level) have been studied in the Gran Dolina, reporting 34 different taxa. The main objective of this work is to complement this information with the bird data of the level TD10.

The first analysis of the TD10 level bird remains reports 345 remains corresponding with at least seven different taxa: Galliformes indet., Falco sp., Passeriformes indet., Turdus sp., Sturnus sp., Prunella collaris, Corvidae indet.

The taphonomic analysis reveals a high level of fragmentation of the bones, and points to the action of a diurnal bird of prey as accumulation agent, as described in other analysis of TD10. Nevertheless the high fragmentation of the remains can be also due to trampling processes.

The results must be taken carefully because the analysis is currently keeping going, more samples will be analyzed. Also is important to note that only the small bird remains are included for now.

196 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Hominins poster V2 MIDDLE PLEISTOCENE BATS FROM THE TD10 LEVEL OF GRAN DOLINA (SIERRA DE ATAPUERCA, BURGOS, SPAIN): NEW INSIGHTS AFTER UPDATING DATA J. Galán1*, C. Núñez-Lahuerta1, J.M. López-García2 and G. Cuenca-Bescós1

1University of Zaragoza, Department of Earth Sciences, Pedro Cerbuna 12, 50018 Zaragoza, Spain 2IPHES, Institut Català de Paleoecologia Humana i Evolució Social, Campus Sescelades URV edifici W3, 43007 Tarragona, Spain

*[email protected]

Keywords: Chiroptera, Middle Pleistocene, mouse-eared bats, horseshoe bats, Atapuerca sites

The sequence of Gran Dolina site (TD), Lower to Middle Pleistocene karst-filling deposits, is divided into eleven lithostratigraphic units (TD1-TD11). Here we present the first results regarding the study of fossil bats from level TD10, a 4m succession of debris flow deposits coming from the outside of the ancient cave (dates from 458±39 ka to 244±26 ka BP). The prolific literature regarding TD10 lithic and faunal assemblages suggests an intensive human occupation. We mean to contribute to the palaeoenvironmental approach of the site and also to the global knowledge of long- term occurrence of chiropters in Iberia.

We analyzed 353 samples obtained by the systematic excavation of sublevels TD10- 3, TD10-2 and TD10-1. Only 25 contained bat remains. Our assemblage comprises 114 specimens assigned to two taxa and consists on teeth and disarticulated quite fragmented bones (mainly cranial elements, sometimes showing dissolution). Myotis myotis appears constantly through the sequence: MNI=10 in TD10-3, MNI=6 in TD10-2, MNI=10 in TD10-1, all groups of age (from yearlings to elders) represented. The size ranges of upper and lower molars were compared with extant Iberian M. myotis, our specimens fitting well but being somehow more elongated antero- posteriorly. One individual of Rhinolophus ferrumequinum appeared in TD10-3.

They are recurrent taxa in the Atapuerca sites, and frequently occur throughout the Iberian Quaternary record as well as along the current Iberian bat communities, where they show preference for woodland or transitional areas. The strong dominance of M. myotis, a cave-roosting taxon, suggests the presence of a nearby colony by the time even if they unlikely inhabited the very cave-site judging by human occupation and taphonomic evidences. The age mortality profile does not fit with theoretical natural-death profiles, thus the accumulation could be originated by a predator, or be the result of catastrophic, rapid events affecting bats roosting in a connected karst cavity and then transported.

The analysis is keep going, and as further researches we mean to: firstly, amply the sampling for sublevels TD10-2 and TD10-3 (only few squares were studied); secondly, compare our assemblage with those from other Middle Pleistocene sites in Atapuerca.

197 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Hominins poster V3 THE LATE MIOCENE (EARLY TUROLIAN, MN11) FAUNA OF CORAKYERLER (TURKEY): NEW COLLECTION OF LARGE MAMMALS A. S. Erol1*, T.T. Kaya2, A.Y.Yavuz3, S. Mayda4 M.Cihat Alçiçek2 and L.W. van den Hoek Ostende5

1Ankara University, Faculty of Languages, History and Geography, Anthropology Department, Sihhiye-Ankara, Turkey. 2 Ege Univ., Natural History Research & Application Center, Bornova, 35100 Izmir, Turkey 3Mehmet Akif Ersoy Univ., Faculty of Art & Sciences, Dept of Anthropology, Burdur-Turkey 4*Ege University, Faculty of Science, Department of Biology, Bornova, 35100 Izmir, Turkey 5Department of Geology, Pamukkale University, 20070 Denizli, Turkey 6Naturalis Biodiversity Center, PO Box 9517, 2300 RA, Leiden, The Netherlands

*[email protected]

Keywords: Corakyerler, Late Miocene, Turolian, Large mammal, Turkey

The Corakyerler Hominoid Site has produced one of the largest and most diverse faunas from the late Miocene of Anatolia since the beginning of 2000. Besides its unique Hominoid record, the characterizing taxa for Corakyerler include also a large component of bovids, as well as diverse carnivores and other ungulates. However, over 3000 fossil specimen awaits taxonomic revision and only some preliminary results are available at this stage of the study. In the framework of the joint project of Ankara and Ege team, we are able to reconstruct the faunal list as well as revising previous records which also includes subsequent records stored in Ege and German Museums. In the first steps of our study, we distinguished new taxa belonging to proboscideans, giraffids, equids and carnivores.

In addition to the proboscidean record represented solely by Choerolophodon, a gomphothere upper molar is now assigned to Tetralophodon longirostris. Hyaenids are characterized by two large sized forms, Lycyaena sp. and Adcrocuta eximia while the Ictitherium record of the former lists is now rejected. Mustelids are well represented in the latest collections which are now assigned to Promephitis, Parataxidea maraghana and the first Turolian Lutrinae record of Turkey which will be assigned to a new species of Sivaonyx. We have also recognized a third giraffid taxon, Samotherium boissieri. The most important finding of the assemblage is a single phalanx which exhibits similarities to the common Turolian chalicothere taxon of Ancylotherium pentelicum. This specimen is also the first Chalicothere record of the fauna while it gives an important paleoecological signal. Last but not least, the second Hipparion form of the fauna is here referred to H. depereti. Both Tetralophodon and H. depereti make their first appearance in the Turolian of Turkey in Corakyerler.

In conclusion, the first results of this systematic revision have revealed 8 new taxa. In the light of the newest additions, we may assign the Corakyerler mammalian fauna to the early Turolian (early MN11). The further study of the fauna will elucidate the position of the fauna in biochronological and chronostratigraphic scales.

SM was supported by “Ege University 2015 FEN 17” Research Grant. This research was supported by grants of Turkish Ministry of Culture and Tourism and Ankara University Anthropology Department. 198 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Hominins poster V4 PILTDOWN MAN AND THE TRIUMPH OF THE DAWSONIAN METHOD S.K. Donovan1*

1Naturalis Biodiversity Center, Darwinweg 2, 2333 CR Leiden, The Netherlands

*[email protected]

Keywords: Eoanthropus dawsoni, forgery, Charles Dawson, Sussex, Plio-Pleistocene

Piltdown Man (Eoanthropus dawsoni, Woodward) was the most notable forgery in 20th Century science. It was published in 1913 and the falsification was not uncovered until 1953. The forger was the amateur archaeologist Charles Dawson (1864-1916). Dawson ‘found’ or was present at the discovery of every fragment of Piltdown Man. Few have appreciated Dawson’s skill; he did much more than merely provide material that fitted with prevalent theories of human evolution. He chose the type locality on private land, not generally accessible. Dawson was ostracized from the local amateur archaeological community, the group most likely to collect the Piltdown site without supervision. Finds were described by the leading palaeontologist in Britain, A.S. Woodward, whose expertise was in lower vertebrates. The leading palaeoanthropologists in Britain, Arthur Keith and G.E. Smith, mainly worked with casts and wasted energies debating theoretical issues. Yet Dawson’s true genius was in presenting British palaeoanthropology with just what it wanted, a large-brained, Pliocene ‘missing link’.

199 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

North Sea Basin poster V5

FIRST RECORD OF WILLOW GROUSE LAGOPUS LAGOPUS (LINNAEUS, 1758) FROM THE LATE PLEISTOCENE OF THE NORTH SEA B.W. Langeveld1*

1 Natural History Museum Rotterdam, Westzeedijk 345, 3015 AA Rotterdam, The Netherlands.

*[email protected]

Keywords: Aves, birds, Eurogeul area, Zandmotor

Two Late Pleistocene humeri fragments represent the first record of willow grouse Lagopus lagopus (Linnaeus, 1758) retrieved from sediments from the North Sea. They were collected from sediments dredged from the Eurogeul area (North Sea close to the Port of Rotterdam, The Netherlands) deposited on the artificial ‘Zandmotor’ coastal reinforcement near The Hague and are kept in the collection of the Natural History Museum Rotterdam. The geology of the sand source area and the material’s taphonomy demonstrate it is Late Pleistocene (Weichselian; last glacial) in age, but inadequate preservation of collagen made it impossible to obtain a 14C-date as independent confirmation. Lagopus lagopus is a common component of the Mammoth Steppe ecosystem and the record presented here contributes to our knowledge of that ecosystem on what is now the bottom of the North Sea.

200 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

North Sea Basin poster V6

MAASVLAKTE 2: A PLEISTOCENE TREASURE TROVE B.W. Langeveld1*and D. Mol1

1Natural History Museum Rotterdam, Westzeedijk 345, 3015 AA Rotterdam, The Netherlands

*[email protected]

Keywords: Eurogeul area, extensive collections, North Sea, Pleistocene mammal remains, private collectors

Maasvlakte 2 is the most recent artificial westerly extension of the Port of Rotterdam (The Netherlands). It has been created from over 240 million m3 suction-dredged, coarse fluvial fossiliferous sediments. These sediments were dredged from the North Sea bed in the Eurogeul area, just offshore the Port of Rotterdam; an area famous for its Pleistocene fossils in great abundance and exquisite preservation. Since its opening in mid-2012, many private collectors regularly have been collecting fossils from the Maasvlakte 2 beach. Our studies show their finds range from unidentifiable bone scraps to fossils that represent species new to The Netherlands or the North Sea. We estimate the total available material to be over 10,000 identifiable specimens. Their age ranges from the Early Pleistocene right up to the Holocene. This provides ample opportunity to add to our still relatively limited knowledge of the Early Pleistocene terrestrial and marine faunas of The Netherlands and the North Sea. Here we argue this rich source of information, both Maasvlakte 2 and the private collections, should not be ignored and demonstrate its value by presenting two intriguing finds: a canine of sabretooth cat Homotherium cf. latidens Owen, 1846 and a humerus of great auk Pinguinus impennis (Linnaeus, 1758).

201 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary poster V7

ASSESSING THE COMPLETENESS OF A “DUTCH” ALLOSAURUS FRAGILIS FROM HISTORICAL PHOTOGRAPHS AND COMPUTED TOMOGRAPHY J. Heijne1, 2*, S. Bijl1,7, J. Ponstein1,3,4, J.H.J.L. van der Lubbe1 , G.A. van Oord4, D.F.A.E. Voeten5, H.B. Vonhof1,6 and A.S. Schulp1,7 1 Faculty of Earth and Life Sciences, Vrije Universiteit Amsterdam, De Boelelaan 1085, 1081 HV Amsterdam, The Netherlands; 2 Division of Paleontology, Steinmann Institute, University of Bonn, Nußallee 8, 53115 Bonn, Germany; 3 Department of Earth Science, Palaeobiology Programme, Uppsala University, Villavägen 16, SE-752 36 Uppsala, Sweden; 4 Museon, Stadhouderslaan 37, 2517 HV Den Haag, The Netherlands 5 Department of Zoology and Laboratory of Ornithology, Palacký University, 17. listopadu 50, 771 46 Olomouc, Czech Republic; 6 Department of Climate Geochemistry, Max Planck Institute for Chemistry, Hahn-Meitner- Weg 1, 55128 Mainz, Germany 7 Naturalis Biodiversity Center, Darwinweg 2, 2333 CR Leiden, The Netherlands

*[email protected]

Keywords: Theropoda, Morrison Formation, quarry documentation, exhibition

In situ specimen documentation represents an important aspect of fossil collecting. However, commercial trade may result in museum collections acquiring fossils that are accompanied by limited or no supporting scientific documentation. Such specimens lack the valuable context information usually required for subsequent scientific analysis and interpretation. An example of a commercially-sourced fossil is the Allosaurus fragilis of Museon (The Hague). Until recently, this individual (M200000) represented the sole large non-avian theropod on exhibit in The Netherlands. The specimen was discovered near the town of Jensen (Utah, United States) at the end of the 20th century and was acquired by the museum in 1999. Since additional documentation for the specimen is not available, its exact provenance and geological context remain unknown. To gain more insight in the specimen itself, we assessed preservation state, skeletal completeness and the mode of restoration for this specimen. Our analysis of the skeleton in its present form is based on morphological observations on the specimen itself, analysis of the scant photographic documentation that presently accompanies the specimen, and CT imaging of the highly fractured skull and mandible. Reconstruction of the skeleton was undertaken before its transport to The Netherlands and involved extensive (re)modelling of (partially) missing skeletal elements with varying degree of accuracy. Approximately 37 percent of the original postcranial elements is present. The small distal elements of the appendicular skeleton and most of the ribs and haemapophyses are lacking. Three-dimensional computed tomography revealed that approximately 75 cranial fragments and 41 teeth (of which 2 had been temporarily removed for isotopic analysis) are preserved. Interestingly, photographic documentation suggests that only four teeth were in situ at the time of collection, which indicates that the other teeth represent isolated fragments that have been added to the skull during restoration. In the mandible, only the fairly complete right ramus consists of genuine fossil bone. The higher degree of completeness of dextral aspects of the cranium and mandible is less evident for the postcranium. Although the skeleton is quite complete and well-suited for educational purposes, the lack of additional information limits its value for future scientific endeavors. 202 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary poster V8

NEW POSTCRANIAL ELEMENTS OF MEGALOSAURIDS (DINOSAURIA, TETANURAE) FROM THE UPPER JURASSIC OF PORTUGAL E. Malafaia1,2, P. Mocho2,3, F. Escaso2,3 and F. Ortega2,3

1Instituto Dom Luiz, Universidade de Lisboa 2Laboratório de Paleontologia e Paleoecologia, Sociedade de História Natural 3Grupo de Biología Evolutiva, Facultad de Ciencias, UNED

*[email protected]

Keywords: Megalosauridae, Torvosaurus, Late Jurassic, Lusitanian Basin

The Portuguese record of Late Jurassic megalosaurids includes a set of cranial and postcranial elements traditionally interpreted as closely related to the North American species Torvosaurus tanneri, but more recently reinterpreted as a species so far exclusive of the Lusitanian Basin: T. gurneyi. Herein it is reported a set of unpublished axial and appendicular elements collected in Praia da Corva (Torres Vedras) that present a combination of shared characters with megalosaurid theropods including Torvosaurus and Megalosaurus. These remains, which show evidences of long weather exposure, were collected from surface along the cliff, in an area composed by upper Kimmeridgian sediments. Thus, and despite these elements present compatible morphology and size, it is not possible to determine if they belong to the same individual. The axial elements include three centra of posterior dorsal vertebrae, a fragment of the neural arch of a dorsal vertebra, and eight partial caudal vertebrae. Appendicular skeleton is represented by an almost complete left fibula and fragments of the tibial diaphysis.

The general morphology of the dorsal centra is similar to that of Megalosaurus sharing a flat and transversely broad ventral surface. The great development of the pleurocoels is comparable with that of Torvosaurus, occupying almost the entire anteroposterior length of the lateral surface the centra. The caudal vertebrae present broad ventral grooves, a character sometimes considered as an autapomorphy of Ceratosaurus, but having a much wider distribution among theropods. These vertebrae share with Torvosaurus the wider than high articular facets and the presence of well-marked neurocentral depressions. The fibula is significantly more robust than those of any other known theropod from the Lusitanian Basin and more similar to Torvosaurus and to an indeterminate megalosauroid from the .

In summary, the specimens of Praia da Corva are assigned to Megalosauridae, with several elements suggesting a close relationship with Torvosaurus. Some distinct characters would be interpreted as autapormorphies of the skull-based species T. gurneyi or belonging to an unknown non-Torvosaurus megalosaurid taxon.

203 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary poster V9

TRIDACTYL TRACKS FROM THE JURASSIC-CRETACEOUS TRANSITION: THE CORCOLILLA TRACKSITE (ALPUENTE, SPAIN) 1 2 3 4 1,5 N.L. Razzolini *, M. Suñer , B. Holgado , C. de Santisteban and À. Galobart

1Mesozoic Research Group, Institut Català de Paleontologia 'Miquel Crusafont' (ICP), C/ Escola Industrial 23, ES-08201 Sabadell, Catalonia, Spain 2Museo de Paleontología de Alpuente, Avda. José Antonio, 17, ES-46178 Alpuente, Valencia, Spain 3Laboratório de Sistemática e Tafonomia de Vertebrados Fósseis, Departamento de Geologia e Paleontologia, Museu Nacional/Universidade Federal do Rio de Janeiro (UFRJ), Quinta da Boa Vista, s/n, São Cristóvão, BR-20940-040 Rio de Janeiro, RJ, Brazil 4Departament de Geologia, Universitat de València, C/ Dr. Moliner, 50, ES-46100 Burjassot, Valencia, Spain 5Museu de la Conca Dellà, carrer del Museu, 4, ES-25650 Isona, Lleida, Spain

*[email protected]

Keywords: Jurassic-Cretaceous transition, tridactyl tracks, photogrammetry, 3-D models, osteological convergence

The Corcolilla tracksite is located in the Villar del Arzobispo Formation (Tithonian- Berriasian, Alpuente, Valencia, Spain) and presents more than a hundred tridactyl footprints of different sizes, previously characterized by four morphotypes attributed to both "small and large theropods and ornithopods" dinosaurs. After more than a decade, excavations of the tracksite have uncovered a new surface and a new trackway was revealed displaying a different morphotype characterized by a very small size (<10 cm), high interdigital divarication angle (II^III and III^IV >50º) and high pace angulation (170-180º). Photogrammetric analyses allowed a reinterpretation of the previously studied tracks as follows: - the "small ornithopod and small theropod morphotypes" correspond to a medium-sized (20-30 cm) tridactyl, symmetrical track, slightly longer than wide, with quadrangular-rounded "heel" pad impressions, robust digits with no discrete phalangeal pads and a notch developing in connection of digit II and "heel" pad. This morphotype strongly recalls the recently emended ichnogenus Iguanodontipus? oncalensis. The "large ornithopod and large theropod morphotypes" correspond to medium-large tridactyl tracks (>40 cm) recognized in at least two trackways featured by rounded "heel" pad with fleshy, subparallel digits impressions with claw marks and sometimes by a very shallow to absent "heel" pad impression, faint constrictions in the digits, claw marks and interdigital angles of 20-30º.We propose that: 1) the newly uncovered <10 cm length tridactyl tracks might correspond to a small-sized avian-like trackmaker, 2) the medium- sized tridactyl tracks might be produced by a non-Iguanodontia ornithopod trackmaker and 3) the medium-large tridactyl tracks do not have a clear origin due to morphological bias and to the osteological convergence during the Jurassic- Cretaceous transition entangling the already dichotomic recognition. Nevertheless, two solutions are proposed for this medium-large tridactyl morphotype: a) medium- sized theropod trackmaker, supported by teeth and large tracks found in the same Formation; b) medium-sized ornithopod trackmaker, supported by osteological remains attributed to basal Ankylopollexia found in the same Formation and by other large ornithopod tracks described for the same age in Portugal (Lourinhã Formation).

204 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary poster H10

CARCHARONDONTOSAURID THEROPOD TEETH FROM BARRANCO DEL HOCINO 1, A BARREMIAN (EARLY CRETACEOUS) SITE OF NE SPAIN 1 1 1,2 2 1 A. Alonso *, J.M. Gasca , P. Navarro-Lorbés , C. Rubio and J.I. Canudo

1University of Zaragoza, Department of Earth Sciences, Pedro Cerbuna 12, 50009 Zaragoza, Spain 2Paleoymás S.L. Pol. Empresarium, C/Retama 17, Nave 24C, 50720 La Cartuja Baja, Zaragoza, Spain

*[email protected]

Keywords: Dinosauria, tetanurae, isolated teeth, Blesa Fm, Iberian Range

The material studied here come from the new vertebrate fossil locality of Barranco del Hocino 1, located within the municipality of Estercuel, Teruel province, Spain. Geologically, outcrops are located in the southwestern area of the Oliete sub-basin, within the northwesternmost part of the Cretaceous Maestrazgo Basin in the Iberian Range. This fossil locality is placed in the middle part of the Blesa Formation, Barremian (Early Cretaceous) in age.

Lithologically the Barranco del Hocino 1 section is comprised by a succession of marly/lutitic levels with evidence of paleosoils alternating with burrowed grey limestone beds. The fossiliferous bed consists of grey lutites with presence of bioturbation, carbonate nodules and calcretes. The fossil content consists of bivalve molluscs, gastropods, charophytes, ostracods and vertebrates. The palaeoenvironment is interpreted as an alluvial plain with evidence of shallow freshwater palustrine episodes and development of palaeosoils.

One campaign of fieldwork has enabled us to recover roughly one hundred bone remains and to identify a moderately diverse assemblage of vertebrates. The fossil locality Barranco del Hocino 1 is a bonebed composed of disarticulated elements with a notable degree of breakage and incompleteness. The fossil association is dominated by ornithopod dinosaurs. In addition, ankylosaur bones, isolated theropod teeth and microvertebrate remains (Crocodylomorpha and Osteichthyes teeth) are also present. The tetanuran carcharodontosaurids are the most abundant isolated theropod teeth, showing the presence of this group in new areas of the Lower Cretaceous of Spain.

Carcharodontosaurid teeth from Barranco del Hocino 1 are ziphodont teeth, with labiolingually compressed and elongated crowns and a lanceolate cross-section. Crowns bear serrated mesial and distal carinae. The enamel surface is ornamented with both marginal and transversal undulations. Two different morphotypes can be recognized: the first one shows more elongation and a slightly narrower cross section. The second has a low profile; the differences between morphotypes may be explained by different tooth positions, ontogenetic variation or the presence of two taxa of this family in Barranco del Hocino 1.

205 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary poster R14

METRIACANTHOSAURIDS (DINOSAURIA: THEROPODA) FROM THAILAND AND PALEOBIOGEOGRAPHY OF METRIACANTHOSAURIDAE A. Samathi1*

1Steinmann-Institut für Geologie, Mineralogie und Paläontologie, Universität Bonn, Nussallee 8, 53115 Bonn, Germany

*[email protected]

Keywords: endemism, regional extinction, paleobiogeography, Early Cretaceous, Thailand

Metriacanthosauridae (= Sinraptoridae) is a group of large-bodied, basal allosauroid theropods from the Middle to Late Jurassic and possibly survived to the Early Cretaceous. They were previously thought to be endemic to China, but were found later in Europe and Thailand. Metriacanthosauridae consists of subgroup Metriacanthosaurinae, ‘Yangchuanosaurus’ group, possibly basal forms: Xuanhanosaurus and Shidaisaurus, and some problematic taxa: Lourinhanosaurus, Poekilopleuron, and Siamotyrannus.

In Thailand, two metriacanthosaurids (Kham Phok and Phu Noi specimens) have been reported from the Late Jurassic – Early Cretaceous Phu Kradung Formation and one problematic taxon (Siamotyrannus) from the Early Cretaceous Sao Khua Formation. The Kham Phok specimen (SM 10) was assigned here to belong to subclade Metriacanthosaurinae based on the bulbous fibular crest on the tibia. The Phu Noi specimens which include skull elements and postcranial materials are still under studied. Preliminary analysis suggested the Phu Noi specimens are closer to Sinraptor dongi from the Upper Jurassic of northwestern China than Yangchuanosaurus from the Middle Jurassic of southwestern China. Siamotyrannus was found to belong to Metriacanthosauridae by Carrano et al (2012), and found to possibly be a basal coelurosaur by Samathi et al (2015) but the metriacanthosaurid hypothesis cannot be rejected yet.

Metriacanthosaurids more probably originated in China at least before the Middle Jurassic and spread to Europe and Southeast Asia during the Middle to Late Jurassic and might survive to the Early Cretaceous. The taxonomic position of Siamotyrannus plus the exact age of Phu Noi locality, Phu Kradung Formation of Thailand will confirm whether metriacanthosaurids could have survived into the Early Cretaceous or not.

206 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary poster R15

A NEW SPECIMEN OF OTHNIELOSAURUS CONSORS 1,2 1 A. Elsler * and J. Kriwet

1University of Vienna, Department of Paleontology, Geozentrum, Althanstrasse 14, 1090 Vienna, Austria 2University of Bristol, School of Earth Sciences, Queens Road, BS8 1RJ Bristol, UK

*[email protected]

Keywords: Othnielosaurus, Neornithischia, Jurassic, Morrison, USA

Here we describe a new specimen of the ornithischian dinosaur Othnielosaurus consors from the Upper Jurassic Morrison Formation. The specimen, nicknamed “Barbara”, was found in fluvial sandstones of the Howe Stephens Quarry (Shell, Wyoming, USA). It represents the most complete and, with a total body length of approximately 2.2 m, largest single skeleton currently known for this taxon. The postcranial skeleton is virtually complete except for the partially preserved forelimbs. The cranial material consists of a relatively well preserved lower jaw and a right maxilla (including dentary and maxillary teeth). The specimen exhibits the most complete cranial material found in a single individual of the taxon. The preservation quality of the bones is variable, ranging from relatively well preserved to heavily crushed.

The specimen can clearly be distinguished from similar ornithischians of the Morrison Formation like Drinker nisti, Dryosaurus altus and Camptosaurus or the heterodontosaurid Fruitadens haagarorum. Slight differences between this skeleton and other specimens referred to Othnielosaurus consors can be noted, but are explained by either individual or ontogenetic variation or taphonomic processes and the preservation quality of the specimen. Whether the size difference in comparison with other specimens is related to ontogeny or sexual dimorphism cannot be determined currently.

A phylogenetic analysis recovers Othnielosaurus consors with the new character scorings as a member of Neornithischia and sister taxon to Cerapoda. This result differs from earlier assessments, which referred to the taxon as a ‘hypsilophodontid’ or basal ornithopod, and is in agreement with relatively recent phylogenetic studies. The position of Othnielosaurus consors, however, remains unstable as reflected by low bootstrap support values. Indeed, additional phylogenetic analyses that include both Kulindadromeus zabaikalicus and Othnielosaurus consors place the latter among ornithopods. Additional cranial material is needed to better establish the relationships of the taxon.

207 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary poster V13

AN UPDATE FOR UDANOCERATOPS TSCHIZHOVI SKULL - (EVEN MORE) UNIQUE CERATOPSIAN DINOSAUR FROM THE LATE CRETACEOUS OF THE GOBI DESERT, MONGOLIA Ł. Czepiński1* and L.Panzarin2

1University of Warsaw, Department of Palaeobiology and Evolution, Faculty of Biology, Biological and Chemical Research Centre, Żwirki i Wigury 101, 02-089 Warsaw, Poland; 2Museo Paleontologico Cittadino Via Valentinis, 134 34074 Monfalcone, Italy

*[email protected]

Keywords: Dinosauria, Ceratopsia, Cretaceous, Mongolia

Leptoceratopsidae is a clade of ceratopsian dinosaurs best known from the Late Cretaceous deposits of North America (i.e. Leptoceratops, Montanoceratops, Gryphoceratops, Unescoceratops) and Asia (i.e. Zhuchengceratops, Ischioceratops and Udanoceratops). The largest member of this group is Udanoceratops tschizhovi which remains were found in Udyn Sayr (Djadokhta Formation, Gobi Desert) and described by Sergei Kurzanov in 1992. The holotype contains many, mostly disarticulated bones of the skull and postcranial skeleton. The cranium is the only material of this species described in literature so far, thus its interpretation is crucial for understanding the biology of this still enigmatic ceratopsian.

Here we report results of our re-examination of the holotypic skull (PIN 3907/11) and found that the previously presented reconstruction was erroneous in several ways. In addition we recognized few cranial bones not mentioned in the original description, which throws new light on its cranial anatomy. Accordingly we provide a new reconstruction of the Udanoceratops cranium, suggesting possible evidence for extreme adaptation in this genus, with unique anatomy among all known ceratopsians. Our investigations may shed new light on evolution and different morphological adaptations patterns of this group of ceratopsian dinosaurs.

208 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary poster V14

ONTOGENETIC TRENDS IN TRICERATOPS - AN ANALYSIS OF CRANIAL AND POST-CRANIAL MORPHOLOGICAL DEVELOPMENT IN AN ASSEMBLAGE FROM WYOMING, USA D. Bastiaans1,2*, T.H. Trapman1,2, M. Guliker2 and A.S. Schulp2,3

1Utrecht University, Faculty of Geosciences, Heidelberglaan 2, 3584 CS Utrecht, the Netherlands 2Naturalis Biodiversity Center, Darwinweg 2, 2333 CR Leiden, the Netherlands 3Vrije Universiteit, Department of Earth Sciences, De Boelelaan 1085, 1081 HV Amsterdam, the Netherlands

*[email protected]

Keywords: Triceratops, Ceratopsidae, Ontogeny, Post-cranial, Assemblage

The ontogeny of Triceratops is a heavily debated topic, and the first qualitative studies in this area have only been conducted since the past few decades, mostly due to the dearth of representative specimens. These ontogenetic studies have predominantly been limited to cranial material, and caused some controversy with relation to the validity of certain ceratopsid taxa (e.g. Torosaurus, Nedoceratops). Ongoing fieldwork in the Lance Formation near Newcastle, Wyoming, by Naturalis Biodiversity Center has uncovered a multigenerational assemblage of at least five individuals of Triceratops and in excess of five hundred bones and bony elements. The site is quite unique as it already yielded a rich assemblage of both cranial and post-cranial material, the latter being relatively scarce. The discovery is particularly remarkable since it includes post-cranial material of several individuals of different sizes, and therefore, presumably, different ontogenetic stages. This provides more resolution in the ontogenetic development in Triceratops post-cranial morphology, especially from late-juvenile or early sub-adult to adult stage. In dinosaurs, a major source of ontogenetic indicators comes from co-ossification regimes (i.e. suture closure). This most likely occurs in a relatively set pattern in dinosaurs, much like it does in mammals. By employing a combination of histology, allometric measurements, and analyses of these and other morphological features (e.g. timing of suture closure and development and loss of cranial ornamentation), the specific mid- to late-stage ontogenetic changes in Triceratops are described. The Triceratops material from Wyoming suggests that individuals noticeably slowed in growth during the sub-adult stage, significantly earlier in ontogeny than the closure of most of the cranial sutures. Histological data and comparison of overall morphology and allometric measurements with existing datasets of Triceratops material seems to suggest multiple sub-adults of slightly variable ages, which would be ideal for describing the changes later in skeletal maturation. Additionally, we postulate that the morphology of certain post-cranial elements (e.g. pelvic region) experiences major remodelling during ontogeny. More intense histological sampling and morphometric analyses combined with overall three-dimensional renderings will aid in determining the exact timing of expression of certain morphological features.

209 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary poster V15

FIRST EVIDENCE FOR HADROSAURIDAE (ORNITHOPODA: HADROSAUROIDEA) FROM CHILEAN PATAGONIA T. Jujihara1, 3*, S. Soto-Acuña2, A. Vargas2, E. Frey3, W. Stinnesbeck4, H. Mansilla5, M. Vogt4 and M. Leppe5

1Karlsruhe Institute of Technology, Department of Civil Engineering, Geo and Environmental Science, Kaiserstrasse 12, 76131 Karlsruhe, Germany 2University of Chile, Department of Biology, Laboratory of Ontogeny and Phylogeny, Las Palmeras 3425, Ñuñoa, Santiago, Chile 3State Museum of Natural History Karlsruhe, Institute of Bio- and Geosciences, Erbprinzenstrasse 13, D-76133 Karlsruhe, Germany 4Heidelberg University, Institute of Geological Sciences, Im Neunheimer Feld 234, 69120 Heidelberg, Germany 5Chilean Antarctic Institute, Laboratory of Paleobiology, Lautaro Navarro 1245, Punta arenas, Chile

*[email protected]

Keywords: Hadrosauridae, Late Cretaceous, South America, Patagonia.

Hadrosaur remains are here recorded for the first time from Chile. The material was collected in the Última Esperanza Province of the Region of Magallanes and Chilean Antarctica in Maastrichtian deposits of the Dorotea Formation and consists of mandibular fragments, cervical and caudal vertebrae, as well as remains of the appendicular skeleton. The presence of hadrosaurs is now identified based on the following characteristics: (1) a dentary with deep alveolar grooves and parallel in lingual surface. (2) cervical vertebral centra heart-shaped anterior and posterior articular facets. (3) caudal vertebral centra with hexagonal outline. (4) humerus with a deltopectoral crest that extends over the proximal half the bone. (5) scapula with a convex dorsal margin. The curvature of the scapula originates level with the dorsal margin of the pseudoacromion process; the outline of the scapula is dorsally most pronounced level with the dorsoventral constriction. (6) straight femoral shaft with a prominent fourth trochanter and distal articular condyles caudally expanded with deep extensor and flexor grooves. (7) straight tibia, which is widely expanded both proximally and distally; distal end with marked asymmetry between the medial and lateral malleolus. (8) spade-shaped, flattened pedal ungual phalanges, which are proximo-laterally expanded beyond the articular facet. The fragmentary material from the Valle Río de Las Chinas area northeast of Torres del Paine National Park conform the southernmost occurrences of dinosaurs in South America and extends the occurrence of South American hadrosaurids by some 570 km to the south. Our discovery together to the previous record of hadrosaur remains in the Antarctic continent support the hypothesis of a late dispersal event of Hadrosauridae from South America into Antarctica, which infers an end-Cretaceous terrestrial connection between Patagonia and the Antarctic Peninsula.

210 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary poster V16

JOURNAL OF PALEONTOLOGICAL TECHNIQUES: A FREE, OPEN ACCESS JOURNAL EXCHANGING KNOWLEDGE BETWEEN TECHNICIANS, PREPARATORS AND RESEARCHERS F.M. Holwerda 1,2,3* and E. Tschopp 1,2,4

1GeoBioTec, Faculdade de Ciencias e Tecnologia (FCT), Universidade Nova de Lisboa, Caparica, Portugal 2Museu da Lourinhã, Rua João Luís de Moura 95, 2530-157 Lourinhã, Portugal 3Staatliche Naturwissenschaftliche Sammlungen Bayerns (SNSB), Bayerische Staatssammlung für Paläontologie und Geologie, Richard-Wagner-Strasse 10, 80333 München, Germany 4Dipartimento di Scienze della Terra, Università di Torino, Via Valperga Caluso 35, 10125 Torino, Italy

*[email protected]

Keywords: publication, open access, collection reports, natural history, conservation

The Journal of Paleontological Techniques (JPT) was established in 2006 in the Museu da Lourinhã (ML), Portugal, as a means to provide a platform for preparators of the ML to share ideas and knowledge with their peers. After this, it grew to be an open access, free journal, publishing mostly on the collecting, preparation, conservation, and exhibition of natural history objects, such as holotypes of extant species, fossils, and historical museum specimens. These natural history objects provide a wealth of information on past and present biodiversity. Because collection and/or conservation techniques might alter the objects in ways that could negatively influence the outcomes of future research, a detailed report of the methodologies used from acquisition to conservation of specimens is crucial. Despite the importance of such reports, until recently, no specific, scientific publications existed for museum technicians and scientists to share knowledge between each other.

The Journal of Paleontological Techniques publishes a wide variety of articles, ranging from excavation reports and papers on preparation techniques to new methodologies in collection management and scientific study, among others. Manuscripts are subjected to peer-review to ensure scientific standards. Papers are published as single-paper volumes upon final approval of the proofs, and are available as .pdf under a CC-BY license.

The editorial board currently consists of an international group of early-stage scientists based in/from Argentina, Belgium, France, Germany, Italy, The Netherlands, Portugal, South Africa, Switzerland and the UK. Most editors have a palaeontological background, all with a unique expertise within that field (e.g. miscroscopy, 3D model production, phylogeny, morphometrics, preparation, and microbiology). JPT welcomes submissions!

211 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary poster V17

NEW DATA ON LATE JURASSIC SAUROPODS OF THE TURCIFAL SUB-BASIN (PORTUGAL) P. Mocho1,2*, R. Royo-Torres3, F. Escaso1,2, E. Malafaia4,2, C. de Miguel Chaves1, I. Narváez1, A. Pérez-García1,2 and F. Ortega1,2

1Grupo de Biología Evolutiva, Facultad de Ciencias, UNED, c/ Senda del Rey, 9, 28040 Madrid, Spain 2Laboratório de Paleontologia e Paleoecologia, Sociedade de História Natural, Polígono Industrial do Alto do Ameal, Pav.H02 e H06, 2565-641, Torres Vedras, Portugal 3Fundación Conjunto Paleontológico de Teruel-Dinópolis/Museo Aragonés de Paleontología, av. Sagunto s/n. E-44002 Teruel, Spain 4Instituto Dom Luiz, Universidade de Lisboa. Edifício C6, Campo Grande, 1749-016 Lisboa, Portugal

*[email protected]

Keywords: Sauropoda, Late Jurassic, Lusitanian Basin

The Turcifal Sub-basin is located at the SW part of the Lusitanian Basin Central sector (Portugal), and is particularly rich in dinosaur fossil-sites, revealing a diverse dinosaur fauna composed by sauropods, theropods, thyreophorans and ornithopods. The sauropods are well represented in this sub-basin. However, relatively scarce information is so far available because most of these specimens are currently under study and/or in preparation. Most of the sauropod discoveries at the Turcifal Sub- basin come from the Sobral and Freixial Formations.

Many sauropod fossil-sites were located in the Torres Vedras municipality, highlighting the localities of Praia Azul, Cambelas, Assenta and Gentias. One of the most important sites is located in Casal da Costa (Cambelas). Appendicular and axial sauropod material, including dorsal vertebrae, was recovered there. The dorsal vertebrae share a similar morphology to those of Camarasaurus supremus, sharing the presence of bifurcated neural spines with camarasaurids. The presence of a circular spinoprezygapophyseal fossa suggests that this sauropod could be attributed to Lourinhasaurus alenquerensis. Another specimen found in Cambelas, known by axial and appendicular remains, was previously related to Diplodocidae based on the presence of a marked expansion of the ischiatic distal end. The presence of a pronounced lateral bulge, observed in the femur of this specimen, is generally considered as a synapomorphy of Titanosauriformes. However, it also occurs in . Other partial skeleton, also found in Cambelas, has an anterior caudal neural spine with a delta-shaped distal end, a feature common in camarasaurids. Other sauropod occurrences in this area include several teeth (heart-, spatulate- and compressed cone-chisel-shaped teeth), axial and appendicular elements. The Upper Jurassic sauropod fossil record is abundant in the Turcifal Sub-basin, and is still relatively unknown. A preliminary analysis of this record notes the presence of basal eusauropods (probably turiasaurs), diplodocids (a specimen with diplodocine affinities), and basal macronarians (including camarasaurids and titanosauriforms).

212 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary poster V18

HISTOLOGY OF THE SAUROPOD LONG BONES FROM THE HOWE-STEPHENS QUARRY (MORRISON FORMATION, WYOMING): TESTING HYPOTHESES OF SKELETAL UNITY K. Wiersma 1*, A. Canoville1 and P.M. Sander1

1Steinmann Institute for Geology, Mineralogy, and Paleontology, University of Bonn, Bonn, 53115, Germany

*[email protected]

Keywords: Histology, Sauropods, Morrison Formation

Sauropod bone histology has emerged as the major source of information on life history of these giant extinct animals. It provides an insight into the growth record and ontogenetic age of an individual. Sauropod specimens from the Morrison Formation are mostly found as partially articulated skeletons or as isolated bones, as seen in, e.g., the Dinosaur National Monument quarry, which makes an assignment of specific bones to individuals difficult. In this study, a detailed assessment of skeletal unity is provided by means of palaeohistology for several Morrison Formation sauropod partial skeletons and isolated long bones from the Howe-Stephens Quarry, Wyoming, USA. Using histological characters such as Histological Ontogenetic Stages, growth marks, annual cyclicity, remodeling rate, and the number of generations of secondary osteons, an assignment of bones to individuals was made and compared with assignments that were made based on field observations. It is possible to histologically assign isolated bones to an existing individual, match isolated bones to comprise a new individual, and to test whether a specific bone belongs to the assigned individual. The hypothesis of skeletal unity was also tested for select other Morrison Formation and Tendaguru specimens, and the method can also be applied here. The findings of this study thus establish a method for testing skeletal unity in fossil tetrapod skeletons.

213 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary poster V19

WHY SOME TITANOSAUR OSTEODERMS HAVE INTERNAL VOIDS AND SOME DON’T? NEW INFORMATION FROM LO HUECO (SPAIN) D. Vidal1*, F. Ortega1, F. Gascó1, A. Serrano-Martínez1 and J.L. Sanz2

1Grupo de Biología Evolutiva, Facultad de Ciencias, UNED, Paseo Senda del Rey 9, 28040 Madrid, Spain 2Unidad de Paleontología, Facultad de Ciencias, Universidad Autónoma de Madrid, Calle Darwin 2, 28049, Madrid, Spain

*[email protected]

Keywords: Titanosaur, Osteoderms, Cretaceous, Lo Hueco, CT-Scan

Titanosaurs were the only sauropods known to have evolved a dermal armor. This dermal armor is made up by i) millimeter-sized dermal ossicles and ii) larger osteoderms referable to at least two morphotypes: "scutes" and "bulb and root". Among the latter, at least two specimens had previously been found to have large voids inside, one in Madagascar (associated with an "adult-sized" specimen of Rapetosaurus) and other in Spain (from the large titanosaur assemblage of Lo Hueco).

A more thorough analysis of the osteoderms found at Lo Hueco (Upper Cretaceous, Spain) by CT scanning and studying natural breakages have revealed that the presence of voids is variable among the sample. CT scanning revealed some osteoderms have small to large lower density areas, interpreted as cavities. All the cavities occur in the core of the bulb region, and develop toward the root end in those osteoderms with the largest voids. Some osteoderms show no evidence of voids, having longitudinal and transverse internal channels instead.

Both channels and cavities are connected with the large foramen located under the bulb in the deep side of the , which suggest that titanosaur osteoderms, like those of modern crocodiles, were extensively perfused by blood vessels.

Osteoclastic remodeling is the most widely accepted hypothesis for the voids. Three main hypothesis have been formulated to explain the mobilization of so much bone from the core of the osteoderm: i) aging, ii) adaptations to extremely seasonal environments and iii) a calcium reservoir for oogenesis.

Of the 16 osteoderms studied, 4 medium to large presented voids, 6 small to large had very dense bone and channels and 6 could not be assessed because of their fragmentary condition: they either could have been entirely compact or have voids of variable sizes. This distribution, and the fact that crocodiles also mobilize calcium from their osteoderms during oogenesis, would support this process to explain voids in titanosaur osteoderm, rather than the other two hypotheses.

214 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary poster V20

NEW DINOSAUR REMAINS FROM THE UPPER MAASTRICHTIAN OF HUESCA, SPAIN (TREMP BASIN, SOUTHEASTERN PYRENEES) E. Puértolas-Pascual1*, J.I. Canudo1, P. Cruzado-Caballero2, X. Pereda- Suberbiola3, J. Larrañaga4 and G. Martín4

1Grupo Aragosaurus-IUCA, Área de Paleontología, Dpto. Ciencias de la Tierra, Universidad de Zaragoza, 50009 Zaragoza, Spain. 2CONICET, Instituto de Investigación en Paleobiología y Geología, Universidad Nacional de Río Negro, Av. Roca 1242, General Roca, 8332, Río Negro, Argentina. 3Universidad del País Vasco/EHU, Dpto. Estratigrafía y Paleontología, Apdo. 644, 48080 Bilbao, Spain. 4Luberri, Oiartzungo Ikasgune Geologikoa Museoa, Pagoaldea pol. 41-42, 20180 Oiartzun, Spain

*[email protected]

Keywords: Dinosauria, Late Cretaceous, K-Pg boundary, Pyrenees

The continental facies of the Tremp Basin (northeastern Spain), together with other European regions such as Provence (France) and the Haţeg Basin (Romania), contain one of the best continental vertebrate record from the Upper Cretaceous of Europe. Therefore, this basin of the southeastern Pyrenees is an exceptional place to study the extinction of continental vertebrates across the Cretaceous-Palaeogene boundary (K-Pg). In this study we present the vertebrate assemblage of the new palaeontological sites discovered in the Serraduy area (Huesca, Spain), located in the distal margin of the Tremp Basin in the northwestern flank of the Tremp syncline. During the last palaeontological prospection campaigns (2014 and 2015), abundant fossil remains in previously known vertebrate sites were recovered, mainly dinosaurs, crocodylomorphs and chelonians. In addition, a magnetostratigraphic campaign was carried out and the westernmost outcrops of the Serraduy area were prospected for the first time allowing the discovery of new palaeontological sites (Larra 1 to 6). The preliminary study of these remains confirms that most of the recovered bones correspond to hadrosaurid remains, highlighting abundant isolated vertebrae (over 40, mostly small in size), several long bones fragments (mainly femora, tibiae, and phalanges), ribs and probably some girdle fragments. In addition, one isolated theropod tooth, chelonian plates and several vertebral centra, osteoderms and isolated teeth of eusuchian crocodylomorphs were also recovered.

The preliminary magnetostratigraphic datings in the area of Serraduy have allowed us to assign these palaeontological sites to the polarity chrons C30n and C29r (latest 2 My of the Mesozoic), as in other nearby fossiliferous areas such as Arén (Huesca, Spain). Therefore, the magnetostratigraphy places these remains within a sedimentary succession which contains the uppermost Maastrichtian, confirming these sites among the most modern in the Upper Cretaceous of Europe, being very close to the K-Pg boundary. This contrasts with more proximal sections of the Tremp Basin (Orcau-Isona area), where recent studies have revealed the presence of a hiatus without record of the chrons C31n to C30n, with the subsequent conversion of many of these sites to an older Maastrichtian age.

215 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary poster V21

TAPHONOMIC MODES IN THE DINOSAUR-BEARING TREMP FM (MAASTRICHTIAN, SOUTHERN PYRENEES) V. Fondevilla1*, B. Vila2, À. Galobart2,3 and O. Oms1

1Departament de Geologia (Estratigrafia), Facultat de Ciències, Universitat Autònoma de Barcelona, Carrer de l’Eix central, 08193, Cerdanyola del Vallès, Barcelona, Spain. 2Institut Català de Paleontologia Miquel Crusafont, Carrer de l’Escola Industrial, 23, E- 08201, Sabadell, Spain. 3Museu de la Conca Dellà, Carrer del Museu, 4, 25650, Isona, Lleida, Spain.

*[email protected]

Keywords: taphonomy, dinosaurs, palaeoenvironments, Maastrichtian, Pyrenees

To date, more than 200 fossil sites have been documented in the Maastrichtian South-Pyrenean basins, which include bones, eggs and clutches, and footprints. The available palaeontological and geological data allow us to focus new studies on the preservation patterns of the bone remains and their relationship with each palaeoenvironment.

At this current stage, we identified 5 taphonomic modes in the dinosaur bone record of the Tremp Formation. The two main taphonomic modes present in the fluvial- deltaic fine-grained channelized sandstones (with possible tidal influence) that characterize the ‘lower red unit’ are the channel-fill and the channel-lag bone accumulations (modes 1 and 2, respectively). These two modes result in fragmentary and very biased bone accumulations. A third taphonomic mode occurs in oxbow lakes and channel margins. This category is featured by better-preserved bones, sometimes partially articulated skeletons, and often includes both macro- and microfossils. Much less frequently, a fourth mode is represented by large accumulations of skeletal remains (bone beds) produced in muddy-silty floodplains (and palaeosols) or in mass flow deposits. This mode 4 has provided large amounts of disarticulated skeletons, and the bias seems to be less than in modes 1 and 2. Variable stages of bone preservation (fracturing, abrasion…) are found among these assemblages. Finally, the bones recovered in mudstones and marly limestones from the lagoonal environments of the 'grey unit' of the Tremp Formation (mode 5) represent the best in terms of quality of preservation and completeness, including single articulated skeletons. This mode, together with mode 3, probably represents autochthonous or parautochthonous assemblages rather than transported accumulations.

Regarding footprints, the fluvial-deltaic fine-grained sandstones of the ‘lower red unit’ preserve excellent examples of dinosaur tracks and skin impressions as natural casts. In the case of dinosaur eggs, they are found as complete eggs and clutches in muddy floodplains and eggshell fragments within sandstone channels.

Future research in the sedimentary processes that generated the bone and eggshell accumulations will enhance the understanding of the current taphonomic modes. This represents a key element to understand the stratigraphic distribution of vertebrate fossils in the Maastrichtian deposits of the southern Pyrenees. 216 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary poster V22

LATE MIOCENE ANURAN FAUNA FROM VOLCHAYA BALKA LOCALITY (NORTH CAUCASUS, RUSSIA) E. Syromyatnikova1,2*

1Borissiak Paleontological Institute of the RAS, Profsoyuznaya 123, 117997 Moscow, Russia 2 Zoological Institute of the RAS, Universitetskaya emb. 1, 199034 St. Petersburg, Russia

*[email protected], [email protected]

Keywords: Late Miocene, Turolian, Russia, Anura, Volchiya Balka

The Late Miocene Volchaya Balka (=Fortepianka) locality is situated near the settlement Gaverdovsky, Maykop District, Republic of Adygea, Russia. It was discovered in 2009 and sampled in 2011–2013 by a joint field team of the Geological Institute of RAS (Moscow) and the Institute of Arid Zones SSC RAS (Rostov-on- Don). The Volchaya Balka locality produced a diverse vertebrate and mollusk fauna of the Late Miocene (early Turolian, MN 11). The list of vertebrates (including anurans) from this site (referred to as Fortepianka) was only preliminary reported. The Volchaya Balka locality yielded numerous bones of Anura, belonging to at least eight taxa: Latonia sp. (Discoglossidae), Bombina sp. (Bombinatoridae), Hyla sp. (Hylidae), Palaeobatrachus sp. (Palaeobatrachidae), Pelobates sp. (Pelobatidae), Bufo sp. (Bufonidae), Rana sp., and Pelophylax sp. (Ranidae). This anuran association is generally common for the Late Miocene European faunas. Latonia sp. clearly differs from the most widely distributed L. gigantea, but is similar to L. ragei and L. vertaizoni in absence of sculpturing of the maxilla. Bombina sp. is similar to B. variegata in shape of junctura ilioischiadica, but the tuber superior varies from well developed (characteristic of B. bombina) to smooth (characteristic of B. variegata). Hyla sp. differs from the recent H. orientalis, which currently inhabits East Europe and West Asia, but is most similar to West European H. arborea in relatively well developed pars descendens of the ilium. Volchaya Balka is the first Late Miocene anuran fauna reported from Russia. It is among the most diverse and oldest anuran faunas of the Russian Neogene. The faunal composition (predominance of aquatic/semiaquatic taxa Latonia, Palaeobatrachus, Pelophylax) indicates a warm climate and presence of long-term permanent water bodies. Other rare taxa probably inhabited the surrounding wooded terrestrial habitats (Hyla), and were subterranean dwellers (Pelobates).

This study was supported by the Russian Foundation for Basic Research, project 15- 04-02079_a.

217 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary poster V23

AMPHIBIANS AND SQUAMATES FROM THE NEW MIOCENE LOCALITY OF MONEVA (SOUTH-CENTRAL EBRO BASIN, SPAIN) J. Benito1* and A. Bolet 2

1School of Geography, Earth and Environmental Sciences, University of Birmingham, Edgbaston, Birmingham; 2Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Barcelona, Spain

*[email protected]

Keywords: Amphibia, Squamata, Amphisbaenia, Serpentes, Miocene

The Middle-Upper Miocene (MN6-MN7) locality of Moneva has yielded a diverse herpetofaunal assemblage containing amphibians, lizards, amphisbaenians and snakes. Moneva is located in the South Ebro Basin, Spain, in a post regressive terrestrial environment, but associated invertebrate remains point to a saline water environment of difficult interpretation and with unclear relation in regard to the vertebrate remains. The fossil material is fragmentary but allows a precise identification of several taxa, as well as a tentative reconstruction of the palaeoenvironment they inhabited.

The identified amphibians include at least two indeterminate Salamandridae on the basis of jaw material, and the anurans Bufo sp., Pelophylax sp., and Rana sp., all represented by their ilia. Lizard material includes dentaries pertaining to Lacertidae indet. and jaws and osteoderms of the anguid Ophisaurus. Blanus sp. is the only amphisbaenian, and is represented by both jaw and vertebral remains. Snake vertebral material is poorly preserved and hence the identification is difficult, but one scolecophidian and several vertebrae with Natricinae affinities can be recognized.

The assemblage fills a gap in the record of Iberian herpetofaunas, as no other localities yielding diverse amphibian and reptile assemblages of this age have been described. The identified taxa follow the expected faunal distribution patterns according with previously known European Miocene climate changes, and are similar to those identified at contemporaneous western and central European localities. The composition of the herpetofauna points to an open and scarcely vegetated area with soils above groundwater level that supported seasonal streams and water bodies.

218 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Saturday plenary poster V24

UNUSUAL SKULL OF CYCLOTOSAURUS FROM THE LATE TRIASSIC OF KRASIEJÓW P. Janecki1,2*, J. Kowalski1,2, A. Bodzioch1 and M. Ruciński3

1Opole University, Department of Biosystematics, Laboratory of Palaeobiology, ul. Oleska 22, 45-052 Opole, Poland. 2Opole University, European Centre of Palaeontology, ul. Oleska 48, 45-052 Opole, Poland. 3Adam Mickiewicz University, Institute of Geology, ul. Maków Polnych 16, 61-606 Poznań, Poland

*[email protected]

Keywords: Cyclotosaurus, Capitosauria, Temnospondyli, Norian

The genus Cyclotosaurus is characterized by closed tinnitus openings, which is the most characteristic feature of Cyclotosauridae and, on the other hand, distinguishes this family from other families of the Capitosauria clade and from the all other Temnospondyli. This genus is well known from several localities in Germany, Poland, Thailand and eastern Greenland, where it occurs in Triassic continental deposits. Seven species have hitherto been included in the genus Cyclotosaurus (C. robustus Meyer and Pleninger 1844; C. mordax Fraas, 1913; C. pasthumus Fraas, 1913; C. hemprichi Kuhn 1942; C. ebrachensis Kuhn, 1932; C. intermedius Sulej and Majer, 2005; C. buechneri Witzmann and Schoch 2016) on the grounds of skull anatomy.

Last years, we found a few Cyclotosaurus skulls in the early Norian bone bed in Krasiejów. One of them (UOPB-1142) was nearly completely preserved together with its jaw arms, and shows some special anatomic features, which are different from all other species described till now. In comparison to the holotype of C. intermedius, which has been described from the same bone bed, the skull has round shaped ear openings (instead teardrop, as also in C. mordax and C. robustus connected with C. intermedius in one evolutionary line), and the biometric parameters are different. The jaw also shows few unique features, and the short row of small tooth on the coronate bones is most interesting because this was never recorded within Cyclotosauridae. All the observations may suggest either extremely high intraspecific variability of C. intermedius (including sexual dimorphism) or existence of at least two species of Cyclotosaurus in Silesia during the early Norian times.

219 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

List of Abstract Authors

Abels, H.A. 112 Aceves Núñez, E. 190 Adams, N. F. 47 Andrianasolo, T. H. 104 Agustí, J. 48 Alba, D.M. 43 Albersdörfer, R. 30 Alçiçek, M.C. 38, 39, 198 Alonso, A. 205 Alpagut, Berna 166 Altamura, S. 139 Andriamandimbiarisoa, L. 104 Angst, D. 108 Anquetin, J. 126 Antczak, M. 182 Arbez, T. 88 Ascarrunz, E. 127 Attard, M.R.G 103 Aurell, M. 148 Aviles Olguín, J. 190 Aytek, A.İ. 38, 162 Badiola, A. 148, 169 Báez, A.M. 19 Baigusheva, V.S. 42 Barendregt, R. W. 47 Barrett, P. M. 29, 155 Barten, J 194 Barten, L 194 Bartolini, S. 43 Bastiaans, D 209 Bauluz, B. 150 Beets, C.J. 112 Bellucci, L. 52 Belmaker, M. 40 Belvedere, M 114, 123 Benito, J. 218 Bennett, C.E. 85, 86 Benson, R. 74, 117 Benton, M.J. 67, 97 Berlioz, E. 45 Bethune, E 105 Bijl, S. 202 Bilgin, M 166 Billon-Bruyat, J.-P. 64 Blanco, A 65 Blondel, C. 45 Bodzioch, A. 90, 180, 181, 182, 219 Boekschoten, G J . 195 Bolet, A. 69, 133, 218 Bolotski, Ivan 119 Bolotski, Yuri 119 Bona, F. 52 Bonilla-Salomón, Isaac 147 Boscaini, A. 43, 122 220 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Botka, D. 39 Braun, B. 35 Bronzati, M. 117 Buccianti, A. 99 Buffetaut, E. 108 Bukhsianidze, M. 36 Bureš, S. 130 Butler, R.J. 157 Candy, Ian 47 Canoville, A. 213 Cantalapiedra, J. 122, 163 Canudo, J.I. 75, 116, 148, 169, 205, 215 Carnevale, G. 125 Castanera, Diego 115, 116 Cattin, Marielle 114 Chakravorti, S 92 Cerdeño, E. 41 Challands, T.J. 84, 85 Cherin, M. 52 Cincotta, A. 168 Clack, J.A. 85, 86, 87 Claessens, L.P.A.M. 102, 137 Claeys, Philippe 119 Claude, Juliën 126 Clausen, P. 103 Clement, A. 84 Clements, Christopher F. 131 Clemmensen, Lars B. 177 Codrea, V.A. 41, 109, 136 Colombero, S. 107 Croitor, R. 53 Cruzado-Caballero, P. 215 Cuenca-Bescós, G. 150, 196, 197 Cuny, G. 64 Czepiński, Ł. 176, 187, 208 D’Souza, L. 140 Dasch, Alexer 163 Davies, S.J. 86 De Boer, E.J. 102 De Esteban-Trivigno, S. 121, 122, 163 de Lapparent de Broin, F. 70 De Louw, P.G.B. 102, 137 de Miguel Chaves, C. 179, 212 de Santisteban, C. 204 de Vos, J. 102, 137, 158 de Wit, Maarten 101 Deeming, D.C. 167 Delfino, M. 125, 170 Demény, A. 59 Demirel, F.A. 38, 39, 166 den Ouden, Natasja 55 Díaz-Berenguer, E. 148, 169 Dick, D. 20 Díez Díaz, V. 118 Dirks, W. 154 Dodson, P. 111 221 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Donovan, Stephen K. 138, 199 dos Santos, Vanda F. 115 Dunhill, A. 97 During, M.A.D. 175 Dutour, Y. 120 Eck, Kristina 192 Elsler, A. 97, 207 Ernst, Scott 114 Erol, Ayla Sevim 198 Escaso, F. 203, 212 Escrig, Christian 121 Espigares, M.P. 43, 51 Evers, S.W. 74 Ezcurra, M.D. 157 Falk, D. 83 Fărcaș, C. 109, 136 Femenias-Gual, J. 146 Floore, R. 137 Flores, J. 67 Fondevilla, V. 216 Fortuny, J. 69, 90, 95, 135 Foth, C. 128 Frey, E. 23, 82, 111, 190, 192, 210 Furrer, H.F. 62 Galán, J. 196, 197 Galobart, À. 115, 204, 216 Ganzhorn, Jörg U. 104 Garbin, R. 127 García-aguilar, J.M. 51 Gasca, J.M. 205 Gascó, F. 214 Gasparik, M. 35 Gere, K. 35 Gil, Lluís 121 Giles, S. 84 Gill, A. 102 Godefroit, P. 119, 168 Göhlich, U.B. 129 Gómez, R.O. 19 González, Paula 163 González González, Arturo 190 Griffin, John N. 131 Gruntmejer, Kamil 90 Guerra-Merchán, A. 51 Güler, Gülşah 166 Guliker, Martijn 229 Guzmán-Gutiérrez, R. 111 Halaçlar, Kazım 39, 166 Handa, N. 153 Harvati, K. 37 Hatsukano, Kenji 192 Hayashi, Shoji 171 Heijne, J. 174, 202 Henderson, S. 84 Herrera, Y. 80 Herrera Flores, Jorge A. 67 222 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Hoch, E. 133 Holgado, B. 204 Holwerda, F.M. 62, 211 Houssaye, Alexra 171 Hume, J.P 102, 137 Iurino, D.A. 52 Jadwiszczak, P. 66 Jagt, John W.M. 76, 195 Jagt-Yazykova, Elena 76, 194, 195 Jalil, N.-E. 78 Janecki, P. 181, 219 Janoo, A. 102, 137 Jansen, M.N. 102, 137 Jaramillo, Carlos 131 Jiménez Fuentes, E. 71 Jiménez-Arenas, J. 51 Jordana, X. 152 Joyce, Walter G. 21, 127, 128 Kaiser, Thomas M. 104, 105, 121, 163 Kanık, Derya 39, 166 Karakütük, Seval 166 Kardynal, K. 180 Kaskes, P. 112, 186 Kaya, T.T. 38, 39, 166, 198 Kear, Benjamin P. 76 Kearsey, T.I. 86 Kędzierski, Mariusz 76 Kesici, Seda Deniz 166 Kierdorf, H. 154 Kierdorf, U. 154 Klein, C.G. 68 Klein, N. 174, 177 Kluiving, S.J. 102, 137 Kocsis, L. 59 Köhler, M. 152 Konidaris, G.E. 37 Konietzko-Meier, D. 90, 91 Kostopoulos, D. 37, 44, 45 Kotsakis, T. 41 Koufos, G.D. 37 Kovács, J. 59 Kowal-Linka, M. 180 Kowalski, J. 180, 181, 219 Krahl, A. 172 Kriwet, Jürgen 207 Kuiper, K.F. 112 Kümmell, Susanna B. 191 Langeveld, B.W. 200, 201 Larrañaga, J. 215 Larson, P.L. 112 Lecuona, A. 78 Ledogar, J.A. 103 Leuzinger, L. 64 Lister, A.M. 140, 161 Liston, J.J. 33, 134 Longrich, N.R. 68, 81 223 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

López-García, J.M. 197 Lovis, Christel 114 Lu, X. 153 Lubeek, J.K. 102, 137 Madurell-Malapeira, J. 43, 51, 152 Maidment, S.C.R. 156 Malafaia, E. 203, 212 Mallison, H. 118, 123 Maltese, A. 27 Maniakas, I. 44 Marcé-Nogué, J. 90, 121, 122, 163 Marchetti, L. 98 Marcos-Fernández, F. 63 Marigó, J. 146, 151 Marjanović, David 89 Markova, A.K. 61 Marshall, J.E.A. 85, 86 Martill, D.M. 25 Martin, J. 22 Martin, R.A. 48 Martin, T. 110 Martín, G. 215 Martinetto, E. 59 Martínez-Navarro, B. 43, 51 Marty, D. 113, 114 Marzola, Marco 177 Masini, F. 107, 141 Masters, Judith 101 Mateus, S 132, 177 Mäuser, M. 82 Maxwell, E. 20 Mayda, S. 38, 39, 166, 198 Mayr, G. 129 Mazza, P.P.A. 99, 107 Medin, T. 51 Meijer, H.J.M. 102, 137 Melis, R.T. 143 Merceron, G. 45 Mészáros, L. 35 Mészáros, R. 35 Meyer, Ch. A. 18, 113, 114 Micklich, N. 185, 192 Mìlan, J 177 Millward, D. 86 Minwer-Barakat, R. 146, 147, 151 Mocho, P. 203, 212 Mol, D. 193, 201 Monsch, K.A. 184 Moreno-Azanza, M 169 Moya-Costa, R. 150 Moyà-Solà, S. 43, 146, 147, 151 Muchagata, João 132 Mujal, E. 135 Mulder, E W.A. 195 Murelaga, X. 70 Muxworthy, A.R. 156 224 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Nacarino-Meneses, C. 152 Nakajima, Y 171 Narváez, I. 212 Navarro-Lorbés, P. 212 Neenan, J.M. 173 Niedźwiedzki, R 76 Nieuwland, I.J.J. 183 Núñez-Lahuerta, C. 196, 197 Obadă, T. 77 Oms, O. 115, 135, 216 Ortega, F. 63, 71, 179, 203, 212, 214 Padilla, S. 20 Palmqvist, P. 43, 51 Palombo, M.R. 50, 106, 139, 142,143, 144, 145 Pandolfi, L. 41, 59, 153 Panzarin, L 208 Paratte, G 114 Parra, M.L. 20 Parrilla-Bel, J. 75 Pawłowska, K. 54 Payros, A. 148 Pazonyi, P. 35 Pereda-Suberbiola, X. 215 Pérez Ben, C. 96 Pérez-García, A. 70, 71, 72, 73, 179, 212 Petculescu, A. 34 Petruso, D. 141 Pietersen, J 175 Pimiento,C 131 Pisani, D. 68 Ponstein, J. 202 Popescu, A 57 Popov, E. 64 Portanger, L.A. 186 Potapova, O. 154 Pouwer, Ronald 55 Prins, M.A. 112 Protopopov, A. 154 Puértolas-Pascual, E. 215 Rabi, M. 125 Rabinovich, R 161 Rakotondranary, S. J 104 Rauhut, O. 32, 62, 117 Razzolini, N.L. 114, 115, 204 Reich, T. 173, 178 Reijmer, John J.G. 175 Rijsdijk, K.F. 102, 137 Rivera-Sylva, H.E. 111 Rojas Sandoval, C 190 Rook, L. 43, 51, 52 Röper, M. 130 Ros-Montoya, S. 43, 51 Ross, P. 103 Royo-Torres, R. 212 Rozzi, R. 142 Rubio, C. 205 225 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Ruciński, M. 181, 219 Rufí, I. 43 Russell, P. 81 Ruta, M. 97 Sachs, Sven 76 Samathi, Adun 206 Sánchez-Villagra, M.R. 189 Sander, P.M. 171, 172, 174, 213 Sanz, J.L. 214 Şarbak, Ayşegül 166 Sardella, R. 52 Sato, Tamaki 171 Savorelli, A. 99, 107, 141 Scheyer, T.M. 173, 178 Schreve, Danielle C. 47 Schulp, A.S. 112, 186, 202, 209 Schulz-Kornas, Ellen 104 Schurian, Bernhard 123 Schwarz, D. 118 Schwermann, A.H. 110 Scofield, P. 103 Sengupta, D.P. 92, 93 Serrano-Martínez, A. 214 Sherwin, J.E. 86 Siber, Hans-Jakob 28 Sidor, Christian A. 98 Silva, Roi 169 Silva-Casal, R. 148 Slowiak, J. 188 Smit, J. 112 Smith, T. 136 Smithson, T.R. 85, 86, 87 Solomon, Al. 109, 136 Sookias, R.B. 157 Stead, Nicola 31 Stein, Koen H.W. 119 Stevens, Kent 114 Steyer, J.-S. 88, 94, 95 Stinnesbeck, Sarah R. 190 Stinnesbeck, Wolfgang 190, 192, 210 Striczky, L. 35 Suñer, M. 204 Surdi, G. 141 Sustaita, D. 103 Syromyatnikova, Elena 217 Szabó, B. 60 Szabó, Márton 65 Szabó, P. 59 Szentesi, Z. 35 Tafforeau, P. 130 Tan, Aytekin 166 Tanner, A. 68 Tesakov, A.S. 46, 48, 49 Teschner, E.M. 91 Thüring, B. 113 Thüring, S. 113 226 XIV EAVP Meeting, 6-10 July, 2016, Haarlem, The Netherlands

Timonina, G.I. 42 Tischlinger, H. 82 Titov, V.V. 42 Tong, H. 126 Tortosa, T. 120 Tourloukis, V. 37 Trapman, T. 79, 209 Tschopp, E. 211 Uhen, M. 131 Underwood, C.J. 56 Unwin, D.M. 24, 167 van den Hoek Ostende, L.W. 100, 166, 198 van der Lubbe, J.H.J.L. 175, 202 van Heteren, A.H. 103 van Kolfschoten, T. 49 van Oord, G.A. 202 Varela, S. 131 Vasile, Ş. 34, 58, 77 Veitschegger, K. 189 Velez-Juarbe, J. 131 Venczel, M. 34, 109, 136 Vennemann, T. 59 Vianey-Liaud, M. 147 Vidal, D. 214 Vila, B. 115, 216 Villa, A. 125 Vinther, J. 68 Vinuesa, V. 43 Virág, A. 35, 58, 60 Voeten, D.F.A.E. 130, 175, 178, 202 Voigt, S. 98 Vonhof, E.H.B 202 Wiersma, K. 213 Wills, M.A. 68, 124 Wings, O. 83, 110, 177 Winkler, D.E. 104, 105 Wintrich, T. 171 Witzel, C. 154 Witzel, U. 172 Wroe, S. 103 Yans, J. 168 Yavuz, A.Y. 198 Zedda, M. 143, 144

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