European Association of Vertebrate Palaeontologists

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th th

Annual Meeting of the of Meeting Annual

Royo-Torres, R., Gascó, F. and Alcalá, L., coord. (2012). 10th Annual Meeting of the European Association of Vertebrate Palaeontologists. ¡Fundamental! 20: 1–290.

EDITOR: © Fundación Conjunto Paleontológico de Teruel – Dinópolis COORDINATION: Rafael Royo-Torres, Francisco Gascó and Luis Alcalá. DISEÑO Y MAQUETA: © EKIX Soluciones Gráficas DL: TE–72–2012 ISBN–13: 978–84–938173–4–3

PROJECT: CGL 2009 06194-E/BTE Ministerio de Ciencia e Innovación

Queda rigurosamente prohibida, sin la autorización escrita de los autores y del editor, bajo las sanciones establecidas en la ley, la reproducción total o parcial de esta obra por cualquier medio o procedimiento, comprendidos la reprografía y el tratamiento informático. Todos los derechos reservados.

2 the accurate geological study has contributed to understand the The last dinosaurs of Europe: clade-specific succession of dinosaur faunas from the latest Campanian to the end heterogeneity in the dinosaur record of the of the Maastrichtian in the Ibero-armorican Island. southern Pyrenees Geological Setting

Àngel Galobart1, José Ignacio Canudo2, Oriol Oms3, The Arén Sandstone and Tremp Formations represent the coastal Bernat Vila2,1, Penélope Cruzado-Caballero2, and coastal to fully continental deposition, respectively, during the Violeta Riera3, Rodrigo Gaete4, Fabio M. Dalla Vecchia1, Late Cretaceous-Palaeocene interval in the southern Pyrenees. Josep Marmi1 and Albert G. Sellés1 They record a marine regression that began near the Campanian- 1Institut Català de Paleontologia Miquel Crusafont, Maastrichtian boundary. In the Tremp Formation four informal C/ Escola Industrial 23, 08201, Sabadell, Catalonia, Spain. lithostratigraphic units have been distinguished (Rosell et al., 2001) [email protected]; [email protected]; (Fig. 1). Dinosaurs bearing sites occur in the two lower units: the [email protected]; [email protected] “grey unit” and in the “lower red unit”. The first one consists in grey marls with abundant invertebrates that usually are interbedded with 2Grupo Aragosaurus-IUCA, Universidad de Zaragoza, layers of coal seams, limestones with charophytes and sandstones C/ Pedro Cerbuna 12, 50009, Zaragoza, Spain. beds. It deposited in lagoonal and coastal wetland settings with [email protected]; [email protected]; variable salinity that laterally evolve to a barrier-island. The age [email protected] ranges from the earliest to the latest Maastrichtian. 3Departament de Geologia, Universitat Autònoma de Barcelona, 08193, Cerdanyola del Vallès, Catalonia, Spain. [email protected]; [email protected]

4Museu de la Conca Dellà, C/ del Museu 4, 25650, Isona, Catalonia, Spain. [email protected]

The outcrops of the Aren and Tremp Formations in the southern Pyrenees, which are located in the Aragón and Catalunya communities, have yielded the most important collection of Late Cretaceous dinosaur remains of Europe. Bones, ichnites and eggs are found in a wide exposure that extends over one hundred kilometres. The importance of these outcrops was first noted by Talens (1955) and Lapparent (1958) in the middle 50s, and later corroborated by other scientists in the last decades (e.g., Casanovas Figure 1. Chronostratigraphy of the Tremp and Aren formations in the et al., 1993; Sanz et al., 1995). Recent fieldworks conducted by the southern Pyrenees (modified from Riera et al., 2009). From east to west: Universidad de Zaragoza and the Institut Català de Paleontologia Vallcebre Syncline, Coll de Nargó Syncline, Tremp Syncline, and Àger have uncovered new dinosaur sites. These new data, together with Synclines.

85 The “lower red unit” is mainly made of red lutites with teeth (Torices et al., 2004; Riera et al., 2009). Peculiar eggs sandstones, lacustrine limestones and palaeosols. These facies are (Sankofa pyrenaica) from a small theropod are known from the interpreted as floodplain alluvial and fluvial deposits with braided upper Arén Formation (López-Martínez and Vicens, 2012) and and meandering channels and palustrine-lacustrine limestones Prismatoolithidae-type eggshells are known from a few sites of the interbedded (Rosell et al., 2001). Its age is late Maastrichtian “lower red unit” (Galobart, 2006). (Riera et al., 2009; Oms et al., 2007; Oms and Canudo, 2004). Ornithopods

Dinosaurian fossil record Hadrosauroids are the most abundant dinosaurs in the Arén and Tremp Formations. About sixty localities with hadrosauroid We completed a continuous stratigraphic record for the bone remains and tracks have been identified, but no egg-sites Maastrichtian of the southern Pyrenees that provides a physical have been reported to date (Dalla Vecchia et al., 2011). Nearly all framework in which the dinosaur sites can be located. sites are located in the “lower red unit” whereas only a few are in the upper part of the underlying “grey unit”. The sampled record Sauropods indicates high hadrosaurid diversity in the upper Maastrichtian Sauropod evidence include tracks and trackways from two with at least three lambeosaurines - Pararhabdodon isonensis, basal levels (Fumanya and Orcau-2; Vila et al., 2005; Vila et al., Arenysaurus ardevoli and Blasisaurus canudoi – an euhadrosaurid 2011a) in the “grey unit” and a few sites with eggs and clutches and an indeterminate ‘hadrosaurine’ (Prieto-Márquez et al., 2006; (Font del Bullidor, Biscarri and Basturs-1, 2; Vila et al., 2010a; Pereda Suberbiola et al., 2009; Cruzado-Caballero et al., 2010a, López-Martínez et al., 2000; Sanz et al., 1995) at the top of the Arén Cruzado-Caballero et al., 2010b, Casanovas et al., 1999). Formation and the “grey unit” (Tremp Formation). Bone remains Ankylosaurs are scant and they occasionally occur in the upper part of the “grey unit” and in the “lower red unit”, probably reflecting differences To date, nodosaurid remains are probably the most infrequent in rock availability between the two units. In the “lower red unit”, dinosaur fossils in the south-Pyrenean basins. They include a few the eggs and clutches are abundant (e.g. Pinyes and Els Terrers teeth and some isolated postcranial material (López-Martínez et sites; Vila et al., 2010b; Vila et al., 2011b) being scarcer upwards. al., 2000; Riera et al., 2009; Escaso et al., 2010). Neither eggs Tracks are rare and they are only found near the K-Pg boundary. In nor tracks have been identified yet. They are always found in the terms of sauropod diversity, at least four distinct titanosaurs forms lower Maastrichtian deposits (lowermost part of the “grey unit” in can be distinguished in the upper part of the Tremp Formation, late the eastern Tremp Syncline), while they were never reported from Maastrichtian in age (Vila et al., 2009). the “lower red unit”.

Theropods Conclusions Theropod remains are rare and distributed in a few localities throughout the upper Arén Formation and Cretaceous part of The southern Pyrenees contain a rich paleontological record to Tremp Formation. They include teeth, eggshells, eggs and rarely understand the terrestrial environmental changes and biota successions bones. Five different theropod taxa have been identified, at least, of the last five million years of the Mesozoic. Over 300 sites with on the basis of dental morphology: Coelurosauria indet., cf. dinosaur remains have been identified to date, containing titanosaur Dromaeosauridae indet., cf. Richardoestesia, cf. Euronychodon sauropods, theropods, hadrosauroid ornithopods and nodosaurid and an indeterminate, large-sized taxon with plesiomorphic ankylosaurians, as well as many eggs sites and thousand of tracks.

86 On the basis of this fossil record, a well dated succession Latest Cretaceous of Arén (Huesca, Spain). Canadian Journal of dinosaurs is established. This is featured by the following of Earth Sciences, 47 (12): 1507-1517. statements: first, there is clear heterogeneity in the stratigraphic CRUZADO-CABALLERO, P., RUIZ-OMEÑACA, J.I. and distribution of the dinosaur taxa in the southern Pyrenees. Second, CANUDO, J.I. 2010b. Evidencias de la coexistencia de dinosaur sites are less frequent in the ‘grey unit’, so most of the hadrosaurinos y lambeosaurinos en el Maastrichtiano superior fossils are from the upper Maastrichtian “lower red unit” of the de la Península Ibérica (Arén, Huesca, España). Ameghiniana, Tremp Formation. Third, changes in dinosaur associations along 47(2): 153-164. the chronostratigraphic succession may reflect time-related events DALLA VECCHIA, F.M., GAETE, R., RIERA, V., OMS, O., (e.g., extinctions and colonizations) or biases (rock availability, PRIETO-MÁRQUEZ, A., VILA, A., GARCIA SELLÉS, A. sampling, environment, and ecology). and GALOBART, A. 2011. The hadrosaurid record in the Maastrichtian of the Eastern Tremp Syncline (Northern Spain). The abundance and diversity of dinosaur fossils in the upper In: Hadrosaur Symposium at the Royal Tyrrell Museum of Maastrichtian of the Pyrenees has a great potential to improve our Palaeontology. Abstract Volume: 38-44. knowledge of the extinction pattern of continental vertebrates at ESCASO, F., PÉREZ-GARCÍA, A., ORTEGA, F. and SANZ, J.L. the Cretaceous – Palaeogene boundary in Europe. 2010. Ankylosaurian evidence from the Upper Cretaceous of South Central Pyrennees (Lleida, Spain): a reappraisal. In: 8 Acknowledgements EAVP Meeting, Aix-en-Provence 2010. Abstract Volume: 33. GALOBART, A. 2006. Importancia del registro español de huevos We would like to thank the Projects of the Subdirección de dinosaurio. In: Los dinosaurios en el siglo XXI, Nuevas general de Proyectos de Investigació, Ministerio de Economía respuestas al inagotable enigma de los dinosaurios, (J.L. Sanz, y Competitividad: CGL2011-30069-C02-01/02 and CGL2010- Ed). Colección Metatemas, Tusquets Editores, Barcelona, 99, 16447.BV acknowledges support from the Ministerio de 98-119 p. Ciencia e Innovación (Subprograma Juan de la Cierva LAPPARENT, A.F. 1958. Découverte d’un gisement d’oeufs de (MICINN-JDC) 2011). Dinosauriens dans le Crétacé supérieur du bassin de Tremp (Province de Lérida, Espagne). Comptes Rendus de l’ Académie References des Sciences de Paris: 1879-1880. LÓPEZ-MARTÍNEZ, N. and VICENS, E. 2012. A new peculiar CASANOVAS, M.L., SANTAFÉ, J.V. and ISIDRO, A. 1993. dinosaur egg, Sankofa pyrenaica oogen. nov. oosp. nov. from the Pararhabdodon isonense n. gen. n. sp. (Dinosauria). Estudio Upper Cretaceous coastal deposits of the Aren Formation, South- morfológico, radiotomográfico y consideraciones biomecánicas. Central Pyrenees, Lleida, Catalonia, Spain. Palaeontology, Paleontologia i Evolució, 26-27: 121-131. 55(2): 325-339. CASANOVAS, M.L., PEREDA-SUBERBIOLA, X, SANTAFÉ, J.V. LÓPEZ-MARTÍNEZ, N., MORATALLA, J.J. and SANZ, J.L. and WEISHAMPEL, D.B. 1999. A primitive euhadrosaurian dinosaur 2000. Dinosaurs nesting on tidal flats. Palaeogeography, from the uppermost Cretaceous of the Ager syncline (southern Palaeoclimatology, Palaeoecology, 160(1-2): 153-163. Pyrenees, Catalonia). Geologie en Mijnbouw, 78: 345-356. OMS, O. and CANUDO, J.I. 2004. Datación magnetoestratigráfica CRUZADO-CABALLERO, P., Pereda-Suberbiola, X. and de los dinosaurios del Cretácico terminal (Maastrichtiense Ruiz-Omeñaca, J.I. 2010a. Blasisaurus canudoi gen. et sp. superior) de Arén (Huesca, Unidad Surpirenaica Central). nov., a new lambeosaurine dinosaur (Hadrosauridae) from the Geotemas, 6(5): 51-54.

87 OMS, O., DINARÈS-TURELL, J., VICENS, E., ESTRADA, R., TORICES, A., RUIZ-OMEÑACA, J.I., CANUDO, J.I. and LÓPEZ- VILA, B., GALOBART, À. and BRAVO, A. M. 2007. Integrated MARTÍNEZ, N. 2004. Nuevos datos sobre los dinosaurios terópodos stratigraphy from the Vallcebre Basin (southeastern Pyrenees, Spain): (Saurischia: Theropoda) del Cretácico superior de los Pirineos Sur- New insights on the continental Cretaceous-Tertiary transition Centrales (Huesca y Lleida). Geo-Temas, 6(5): 71-74. in southwest Europe. Palaeogeography, Palaeoclimatology, VILA, B., OMS, O. and GALOBART, À. 2005. Manus-only Palaeoecology, 255: 35-47. titanosaurid trackway from Fumanya (Maastrichtian, Pyrenees): PEREDA SUBERBIOLA, X., CANUDO, J.I., CRUZADO- further evidence for an underprint origin. Lethaia, 38: CABALLERO, P., BARCO, J.L., LÓPEZ-MARTÍNEZ, N., OMS, 211-218. O. and RUIZ-OMEÑACA, J.I. 2009. The last hadrosaurid dinosaurs VILA, B,. GALOBART, A., CANUDO, J.I., LE LOEUFF, J. and of Europe: A new lambeosaurine from the Uppermost Cretaceous of OMS, O. 2009. Late Cretaceous sauropod diversity in Southern Aren (Huesca, Spain). Comptes Rendus Palevol, 8: 559-572. Europe. Journal of Vertebrate Paleontology 29(3):196A. PRIETO-MÁRQUEZ, A., GAETE, R., RIVAS, G., GALOBART, VILA, B., GALOBART, À, OMS, O., POZA, B. and BRAVO, À. and BOADA, M. 2006. Hadrosauroid dinosaurs from the A.M. 2010a. Assessing the nesting strategies of Late Cretaceous Late Cretaceous of Spain: Pararhabdodon isonensis revisited titanosaurs: 3-D clutch geometry from a new Megaloolithid and Koutalisaurus kohlerorum, gen. et sp. nov. Journal of eggsite. Lethaia, 43: 197-208. Vertebrate Paleontology, 26(4): 929-943. VILA, B., JACKSON, F.D., FORTUNY, J., SELLÉS, A.G. and RIERA, V., OMS, O., GAETE, R. and GALOBART, À. 2009. GALOBART, À. 2010b. 3-D modelling of megaloolithid The end-Cretaceous dinosaur succession in Europe: the Tremp clutches: insights about nest construction and dinosaur basin record (Spain). Palaeogeography, Palaeoclimatology, behaviour. PLoS ONE 5 (5): e10362. Palaeoecology, 283: 160-171. VILA, B., MORATALLA, J.J., GAETE, R., SANTOS, V. and ROSELL, J., LINARES, R. and LLOMPART, C. 2001. El GALOBART, À. 2011a. New titanosaur trackways from southern “Garumniense” Pirenaico. Revista de la Sociedad Geológica de Pyrenees: Orcau-2 locality (Late Cretaceous) revisited. In: España, 14 (1-2): 47-57. Dinosaur Track Symposium. Abstracts Volume: 34-35. SANZ, J.L., MORATALLA, J.J., DÍAZ-MOLINA, M., LÓPEZ- VILA, B., RIERA, V., BRAVO, A.M., OMS, O., VICENS, E., MARTÍNEZ, N., KÄLIN, O. and VIANEY-LIAUD, M. 1995. ESTRADA, R. and GALOBART, À. 2011b. The chronology of Dinosaurs nests at the sea shore. Nature, 376: 731-732 dinosaur oospecies in south-western Europe: Refinements from TALENS, J. 1955. Descubrimiento de dinosaurios en Tremp the Maastrichtian succession of the eastern Pyrenees. Cretaceous (Lérida). Estudios geológicos, 25: 86. Research, 32: 378-386.