Island Life in the Cretaceous
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A peer-reviewed open-access journal ZooKeys 469: 1–161Island (2015) life in the Cretaceous - faunal composition, biogeography, evolution... 1 doi: 10.3897/zookeys.469.8439 REVIEW ARTICLE http://zookeys.pensoft.net Launched to accelerate biodiversity research Island life in the Cretaceous - faunal composition, biogeography, evolution, and extinction of land- living vertebrates on the Late Cretaceous European archipelago Zoltán Csiki-Sava1, Eric Buffetaut2, Attila Ősi3, Xabier Pereda-Suberbiola4, Stephen L. Brusatte5 1 Department of Geology, Faculty of Geology and Geophysics, University of Bucharest, 1 N. Bălcescu Blvd, 010041 Bucharest, Romania 2 Centre National de la Recherche Scientifique, UMR 8538, Laboratoire de Géologie de l’Ecole Normale Supérieure, 24 rue Lhomond, 75231 Paris Cedex 05, France 3 MTA-ELTE Lendület Dinosaur Research Group, Pázmány Péter sétány 1/c, Budapest, 1117, Hungary 4 Universidad del País Vasco/Euskal Herriko Unibertsitatea, Facultad de Ciencia y Tecnología, Departamento de Estratigrafía y Paleontología, Apartado 644, 48080 Bilbao, Spain 5 School of GeoSciences, University of Edinburgh, Grant Institute, King’s Buildings, West Mains Road, Edinburgh EH9 3JW, UK Corresponding author: Zoltán Csiki-Sava ([email protected]) Academic editor: Hans-Dieter Sues | Received 19 August 2014 | Accepted 20 November 2014 | Published 8 January 2015 http://zoobank.org/470FC0A3-90EB-4581-8064-D6F78048429C Citation: Csiki-Sava Z, Buffetaut E, Ősi A, Pereda-Suberbiola X, Brusatte SL (2015) Island life in the Cretaceous - faunal composition, biogeography, evolution, and extinction of land-living vertebrates on the Late Cretaceous European archipelago. ZooKeys 469: 1–161. doi: 10.3897/zookeys.469.8439 Abstract The Late Cretaceous was a time of tremendous global change, as the final stages of the Age of Dinosaurs were shaped by climate and sea level fluctuations and witness to marked paleogeographic and faunal changes, before the end-Cretaceous bolide impact. The terrestrial fossil record of Late Cretaceous Europe is becoming increasingly better understood, based largely on intensive fieldwork over the past two decades, promising new insights into latest Cretaceous faunal evolution. We review the terrestrial Late Cretaceous record from Europe and discuss its importance for understanding the paleogeography, ecology, evolution, and extinction of land-dwelling vertebrates. We review the major Late Cretaceous faunas from Austria, Hungary, France, Spain, Portugal, and Romania, as well as more fragmentary records from elsewhere in Europe. We discuss the paleogeographic background and history of assembly of these faunas, and argue that they are comprised of an endemic ‘core’ supplemented with various immigration waves. These faunas lived on an island archipelago, and we describe how this insular setting led to ecological peculiarities such as low diversity, a preponderance of primitive taxa, and marked changes in morphology (particularly body Copyright Zoltán Csiki-Sava et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 2 Zoltán Csiki-Sava et al. / ZooKeys 469: 1–161 (2015) size dwarfing). We conclude by discussing the importance of the European record in understanding the end-Cretaceous extinction and show that there is no clear evidence that dinosaurs or other groups were undergoing long-term declines in Europe prior to the bolide impact. Keywords Late Cretaceous, Europe, island, faunal evolution, paleobiogeography, extinction Table of contents Introduction ............................................................................................................. 3 Europe in the Late Cretaceous: Paleogeography and Paleotectonics of an Ancient Island Archipelago ............................................................................................... 6 The Late Cretaceous Continental Vertebrate Faunas of Europe ............................... 10 A. General Overview .......................................................................................... 10 Cenomanian .................................................................................................. 11 Turonian ........................................................................................................ 19 Coniacian ...................................................................................................... 20 Santonian ....................................................................................................... 21 Campanian .................................................................................................... 23 Maastrichtian ................................................................................................. 25 B. Santonian, Hungary (Iharkút and Ajka) ......................................................... 29 History of research ......................................................................................... 29 Geological setting ........................................................................................... 29 Faunal overview ............................................................................................. 31 C. Lower Campanian, eastern Austria (Muthmannsdorf) ................................... 36 History of research ......................................................................................... 36 Geological setting ........................................................................................... 36 Faunal overview ............................................................................................. 37 D. Santonian–Maastrichtian, Southern France ................................................... 39 History of research ......................................................................................... 39 Geological setting ........................................................................................... 40 Faunal overview ............................................................................................. 42 E. Campanian–Maastrichtian, Spain and Portugal ............................................. 46 History of research ......................................................................................... 46 Geological setting ........................................................................................... 47 Faunal overview ............................................................................................. 51 F. Coniacian and Campanian–Maastrichtian, Romania ...................................... 57 History of research ......................................................................................... 57 Geological setting ........................................................................................... 58 Faunal overview ............................................................................................. 61 Discussion and Conclusions ................................................................................... 69 Late Cretaceous faunal composition, evolution and paleobiogeography .............. 69 Faunal composition: distribution, endemism and provinciality .................. 71 Island life in the Cretaceous - faunal composition, biogeography, evolution... 3 Faunal composition and evolution: the history of research ......................... 75 Faunal composition and evolution: the old European core ......................... 76 Faunal composition and evolution: the Asiamerican kinship ...................... 77 Faunal composition and evolution: the Gondwanan immigrants ................ 80 Interactions between Late Cretaceous Europe and other bioprovinces – origin, timing and route of faunal connections ...............................................................82 Late Cretaceous faunal evolution in continental Europe ............................. 85 Late Cretaceous island life .............................................................................. 92 Insularity-related features of the European Late Cretaceous vertebrate assem- blages ......................................................................................................... 94 Insularity-related adaptations of the European Late Cretaceous island-dwelling taxa ............................................................................................................ 96 Events at the Cretaceous-Tertiary boundary in the European Archipelago ........100 Dinosaurs ................................................................................................ 100 Other vertebrates ..................................................................................... 102 Patterns of extinction and survival near the Cretaceous–Paleogene boundary ...105 Acknowledgements ............................................................................................... 108 References ............................................................................................................ 109 Introduction The most iconic picture of a Late Cretaceous terrestrial ecosystem is probably Tyran- nosaurus attacking Triceratops on the vast, fertile floodplains of North America, as a suite of smaller dinosaurs, mammals, crocodyliforms, turtles, and pterosaurs look on. This vignette has been repeated often in movies and museum exhibits, and for good reason: the terrestrial fossil record