A new titanosaur (Dinosauria: Sauropoda) from the Upper Cretaceous of Velaux-La-Bastide Neuve (southern France)

ABSTRACT

Remains of Garrigatitan meridionalis nov. gen. et sp. were found in two bonebeds of sequence 2 from the upper Campanian site of Velaux-La Bastide Neuve (Aix-en-Provence Basin, Bouches du Rhône department). The vertebrate assemblage is dominated by dinosaurs, including the titanosaur Atsinganosaurus velauciensis. Garrigatitan meridionalis presents three diagnostic characters: hourglass-shaped humeri (proximal and distal thirds of almost the same transversal width) in anterior and posterior views, ilium with a broad rounded hollow slightly posterior to the base of the pubic peduncle, proximolateral margin of the femur only slightly medially deflected. Garrigatitan was a small to medium-sized sauropod (sub/adult individuals between 4–6 metres and 2–2.5 tonnes), showing anatomical differences with Atsinganosaurus, and with the other Late Cretaceous Ibero-Armorican titanosaurs. Large titanosaurian specimens found at Velaux-La Bastide Neuve could belong to adult Garrigatitan individuals reaching a body length of at least 12 metres. Histological analysis of long bones shows features similar to other Late Cretaceous European titanosaurs, indicating that all individuals had reached skeletal maturity (presence of an EFS, heavy remodelling HOS 12 to 14). The new taxon is recovered within the clade Lirainosaurinae. Garrigatitan meridionalis increases the diversity of Late Cretaceous titanosaurs within the Ibero-Armorican Island.

KEYWORDS:

• France
• Garrigatitan meridionalis
• Late Cretaceous
• Lirainosaurinae
• Sauropoda
• Titanosauria

Acknowledgments

We would like to acknowledge A. Montaufier for the photographs (with financial support of the Lisea-Vinci group), and C. Zafra for the edition of the figures. We thank J. C. Corral and J. Alonso (Museo de Ciencias Naturales de Álava/Arabako Natur Zientzien Museoa, Vitoria-Gasteiz, Spain), B. Madarieta (Museo Vasco de Historia de la Medicina y de las Ciencias of Leioa, Spain), F. Ortega (Grupo de Biología Evolutiva, Universidad Nacional de Educación a Distancia, Spain), J. Le Loeuff (Musée des Dinosaures d’Espéraza, France), and P. Barrett and S. Maidment (Natural History Museum London, U.K.). F. Holwerda (Utrecht University, Utrecht, The Netherlands) and E. Tschopp (American Museum of Natural History, New York, USA) helped in the development of the phylogenetic analysis, and P. Mannion (UCL, U.K.) is thanked for the photographs of Normanniasaurus and his useful comments and help. The Willi Hennig Society sponsors the use of the TNT cladistics software. BJC thanks J. Laval for producing the thin sections for histologic purposes, V. Fischer for his useful critics on the long bone histology part, and Khimaerai (J. Jentgen) for having helped a lot to produce the histologic figures. We would like to thank the editor G. Dyke, and the useful comments and suggestions provided by S. Poropat and two anonymous reviewers, which greatly improved this manuscript.

This work has been developed thanks to the collaboration between Palaios (a research association presided by XV), the University of Poitiers, the Velaux Municipality (Y. Guérin, J.-P. Maggi, and C. Peru) with its heritage, culture and technical services (M. Calvier) and association “Les amis du patrimoine” (L. Melih), the environment department from CD 13 (M. Bourrelly, T. Tortosa, G. Michel, N. Mouly, and S. Amico), the ‘Service Départemental d’Incendie et de Secours’ (SDIS 13), and numerous volunteers during the field campaigns in 2009 and 2012 (particularly N. Bardet, T. Beauprez, A. Bernet, S. Berton, A. Cincotta, M. Bournoneville, D. Brossens, S. Goolaerts, L. Guiraud, T. Hublin, P. Lauters, B. Mertens, L. Villier and D. de Winter). This work was supported by the Ministry of Education and Communication (research grant VR1013 to Palaios association), the Bouches du Rhône department CD 13 proposals MAPADGAC23112010-1 and MAPADGAC16012014-1-AAPC. Research work of XPS was supported by the Spanish Ministerio de Ciencia e Innovación (project CGL2017-85038-P), the European Regional Development Fund, the Gobierno Vasco/Eusko Jaurlaritza (research group IT-1418-19) and the Universidad del País Vasco (UPV/EHU, research group PPG17/05), and that of BJC by a FRIA grant of the Fonds de la Recherche Scientifique. KS thanks the Fonds Wetenschappelijk Onderzoek for funding.

Disclosure statement

The authors declare no conflict of interest.

Additional information

Funding

This work was supported by the European Regional Development Fund; Fonds de la Recherche Scientifique - FNRS; Fonds Wetenschappelijk Onderzoek; Spanish Ministerio de Ciencia e Innovación [CGL2017-85038-P]; Gobierno Vasco/Eusko Jaurlaritza [IT-1418-19]; Bouches du Rhône department CD 13 [MAPADGAC16012014-1-AAPC,MAPADGAC23112010-1]; French Ministry of Education and Communication [VR1013]; Universidad del País Vasco [PPG17/05].