The "sauropod" from the Albian of Mesnil-Saint-Père (Aube, France):
a pliosaur, not a dinosaur.
Eric Buffetaut
Claude Colleté
Bruno Dubus2
Jean-Louis Petit2
Abstract: A vertebra from the Albian of Mesnil-Saint-Père (Aube, eastern Paris Basin), previously identified as the first caudal of a sauropod dinosaur, is shown to be a dorsal vertebra of a large pliosaur. The specimen resembles vertebrae from the Albian of England and eastern France that have been referred to the pliosaur Polyptychodon, a taxon in need of revision.
Key Words: Vertebra; Cretaceous; France; Plesiosauria; Sauropoda
Citation: Buffetaut E., Colleté C., Dubus B. & Petit J.-L. (2014).- The "sauropod" from the Albian of Mesnil-Saint-Père (Aube, France): a pliosaur, not a dinosaur.- Carnets Geol., Madrid, vol. W, no. X, p. yyy-zzz.
Résumé : Le "sauropode" de l'Albien de Mesnil-Saint-Père (Aube, France) est un pliosaure, non un dinosaure.- Il est montré qu'une vertèbre provenant de l'Albien de Mesnil-Saint-Père (Aube, E du Bassin de Paris), précédemment identifiée comme une première caudale d'un dinosaure sauropode, est en fait une vertèbre dorsale d'un grand pliosaure. Le spécimen évoque des vertèbres de l'Albien d'Angleterre et de l'Est de la France qui ont été attribuées au pliosaure Polyptychodon, taxon qui a besoin d'être révisé.
Mots-Clefs : Vertèbre ; Crétacé ; France ; Plesiosauria ; Sauropoda
Introduction 
In 2000, Knoll et al. described an isolated vertebra from the Albian of Aube (eastern Paris Basin) as an anterior caudal of a sauropod dinosaur (see also the French translation: Knoll et al., 2002). This identification was subsequently accepted by Allain and Pereda Suberbiola (2003), and on its basis Weishampel et al. (2004, p. 560) listed the occurrence of Sauropoda indet. in the Albian of Aube in their review of dinosaur distribution.
We have re-examined the specimen and found that its identification as a sauropod vertebra is erroneous. The vertebra from Mesnil-Saint-Père in fact does not belong to a dinosaur; as shown below, it is clearly a dorsal vertebra of a large pliosaur.
Redescription and reinterpretation of the specimen
As noted by Knoll et al. (2000), the vertebra found by one of us (J.-L. P.) at Mesnil-Saint-Père apparently comes from clayey layers overlying the "Sables verts" (Greensand), and is considered to be Early Albian in age. The specimen is in the J.-L. Petit collection in Troyes and can be examined through contact with the Association Géologique Auboise; a cast of it is kept at the Musée des Dinosaures, Espéraza, France.
The specimen (Fig. 1) is generally well-preserved, although some abrasion of the surface of the bone probably indicates transport. It consists of a fairly complete centrum: only the posterodorsal and right posterolateral regions are damaged and partly missing. Contrary to the description by Knoll et al. (2000), the neural arch is not completely missing, for its pedicels are clearly visible in the anterior half of the specimen, enclosing the floor of the neural canal (the posterior part of the pedicels is largely missing). The anterior articular surface is nearly circular in outline, and very shallowly concave. It shows a well-marked central elevation or papilla. Similarly, the posterior articular surface, which is slightly more concave, appears to have been circular in outline, and bears a central elevation. The lateral faces of the centrum are concave anteroposteriorly and show no sign of a rib articulation or transverse process. The ventral face is concave anteroposteriorly, with a prominent posterior margin (exaggerated by deformation). There are no chevron facets. No large foramina are visible on the ventral and lateral surfaces of the centrum, although small, irregularly distributed foramina, can be seen. Those visible on the left side of the centrum appear as oval pits with a maximum diameter of 4 mm.

FIG. 1 about here

Knoll et al. (2000) considered the specimen from Mesnil-Saint-Père as the first caudal vertebra of a sauropod dinosaur. However, this identification is untenable, for the simple reason that in sauropods the first caudal (like the other anterior caudals) bears well-developed transverse processes which extend over a large part of the lateral sides of the centrum (see, for instance, McIntosh, 1990, Fig. 16.7, p. 363, and Upchurch et al., 2004, Fig. 13.9, p. 286). Even if the neural arch was totally missing on the vertebra from Mesnil-Saint-Père, which is not the case, the inception of the transverse processes on the sides of the centrum would be visible. Instead, the lateral faces are smooth. According to Knoll et al. (2000), the vertebra from Mesnil-Saint-Père shows very strong similarities with the first caudal of Brachiosaurus brancai. However, a comparison with Janensch's description and figures Janensch (1950) shows that this is inaccurate: in Brachiosaurus brancai, as in other sauropods, there are well-developed transverse processes, inserting on the lateral faces of the centrum, down to nearly mid-height. Nothing of the sort can be seen on the specimen from Mesnil-Saint-Père; its well-preserved left side clearly shows the absence of any process. Consequently, there can be no doubt that the identification of the vertebra from Mesnil-Saint-Père as a sauropod caudal is erroneous. The characters of this vertebra do not fit in a sauropod vertebral column.
The specimen from Mesnil-Saint-Père can be identified as a dorsal vertebra of a large sauropterygian. The fact that there are no insertion facets for ribs on the lateral faces shows that it must have been located far posteriorly in the dorsal series, so that the rib facets were located on the now missing parts of the neural arch. The more or less cylindrical, although constricted, shape of the centrum is also in agreement with its identification as a sauropterygian vertebra. Well-marked central papillae on the articular surfaces, bearing or not a Notochordal pit, are a very frequent feature of sauropterygian vertebrae, whereas they are not common in sauropods. On the whole, the vertebra from Mesnil-Saint-Père closely resembles the dorsal vertebrae of large Cretaceous representatives of the Plesiosauria. In size and shape, for instance, it compares well with a large vertebra from the Upper Cretaceous of Elbing, Prussia (today Elblag, Poland) described by Schröder (1885) as Pliosaurus ? gigas (referred to as Plesiosauria indet. by Bardet and Godefroit, 1995).
One unusual feature of the vertebra from Mesnil-Saint-Père is the lack of large foramina on the ventral or lateral surfaces of the centrum. These "foramina subcentralia" (Storrs, 1991) are very common in the Plesiosauria, and their absence in the specimen from Aube may at first sight seem surprising. However, in some pliosaur taxa the foramina subcentralia are reduced or missing altogether. As noted by Williston (1903, p. 62), in the Late Cretaceous Brachauchenius lucasii, "the ventral vascular foramina, so characteristic of plesiosaur vertebrae, appear to be wanting throughout the whole series". Schumacher and Everhart (2005) report the same lack of ventral vascular foramina in a pliosaurid from the Turonian of Kansas. According to O’Keefe (2001), reduction of the foramina subcentralia is a derived character found, in the family Pliosauridae, in the genera Liopleurodon, Pliosaurus and Brachauchenius. The lack of such foramina in the specimen from Mesnil-Saint-Père is therefore not a sufficient reason to exclude it from the Plesiosauria, and may suggest that it belongs to a member of the family Pliosauridae (a conclusion also supported by the large size of the vertebra).
Measurements:
Height of anterior articular face of centrum: 137 mm	
Width of anterior articular face of centrum: 140 mm	
Ventral length of centrum: 108 mm
Estimating the total length of a pliosaur on the basis of an isolated vertebra is fraught with difficulties. On the basis of the proportions of the mounted skeleton of Kronosaurus queenslandicus, from the Lower Cretaceous of Australia, described by Romer and Lewis (1959) as being 12.80 m long, the pliosaur from Mesnil-Saint-Père may have been about 10 metres long. However, the total number of vertebrae in that specimen of  is uncertain and its total length may have been overestimated. Kear (2003, p. 291) describes Kronosaurus as a "large-bodied pliosaurid up to and probably in excess of 9 m". This would suggest an approximate length of 7 metres for the pliosaur from Mesnil-Saint-Père. Moreover, the proportions of the pliosaur from Aube may have been different from those of Kronosaurus, so the above estimates must be considered as very rough approximations. Be that as it may, the pliosaur vertebra from Mesnil-Saint-Père clearly belonged to a very large animal.
Remarks on the pliosaurs from the "Sables verts" of the eastern Paris Basin
The vertebra from Mesnil-Saint-Père is not the first pliosaur specimen to be reported from the Albian Greensand of the eastern Paris Basin. Barrois (1875) noted the occurrence of Polyptychodon teeth at Grandpré (Ardennes) and Louppy (Meuse). Sauvage (1882) described and illustrated teeth from Grandpré, and in 1903 described a tooth and eight vertebrae from Varennes-en-Argonne (Meuse). Corroy (1922) also mentioned Polyptychodon in the Albian of the Paris Basin and noted that the fossils from Varennes indicated an individual of an enormous size.
Polyptychodon was erected by Owen (1841) on the basis of isolated teeth, with two species, P. interruptus and P. continuus, which may in fact correspond to teeth from different positions in the tooth row (Sauvage, 1903). Teeth and other remains from many mid-Cretaceous formations, ranging in age from Albian to Turonian, in Europe and North America (Welles & Slaughter, 1963), have been referred to that genus. Although Bardet and Godefroit (1995, p. 182) considered it as "systematically well defined", Polyptychodon remains a relatively poorly known taxon. O’Keefe (2001) did not include it in his phylogenetic analysis and classification of the Plesiosauria. Nevertheless, and although they did not redefine the taxon, Bardet's and Godefroit's conclusion (Bardet & Godefroit, 1995, p. 183) that it is a "valid genus of Pliosauridae" is probably well-founded.
Although the type species of Polyptychodon was based on teeth, isolated vertebrae from deposits where such teeth have been found have been referred to this genus. Thus, Seeley (1876) described several vertebrae from the Cambridge Greensand as belonging to Polyptychodon. The Cambridge Greensand vertebrates are considered as Late Albian, reworked into the basal Cenomanian (Unwin, 2001). Photographs of a dorsal vertebra (Sedgwick Museum, CAMSM B57396) from that collection kindly provided by Dr Leslie Noè (Sedgwick Museum, Cambridge) show considerable similarities with the specimen from Mesnil-Saint-Père, both in the shape of the centrum and in the absence of large foramina subcentralia (Fig. 2).

FIG. 2 about here

The vertebrae from Varennes-en-Argonne referred by Sauvage (1903) to Polyptychodon are of particular interest because they appear to be reminiscent of the specimen from Mesnil-Saint-Père both in their morphology (most of them are described as having more or less concave articular faces with a strong central papilla) and in their size (vertebra "8" is described as being 140 mm high, 115 mm long and 135 mm wide), although most of them show costal facets on the sides of the centrum and must have come from more anterior parts of the vertebral column. However, whether the vertebrae described by Sauvage (1903) can safely be referred to Polyptychodon is uncertain; there is no clear evidence that the tooth and the vertebrae he mentioned were from a single individual (associated skeletal elements seem to be rare in the Sables verts, although Sauvage (1882) did describe a partly articulated theropod skeleton). This applies even more to the isolated vertebra from Mesnil-Saint-Père: pending a revision of Polyptychodon in general, and of the pliosaur remains from the Sables verts in particular, it seems safer to refer to it as Pliosauridae indet. The coexistence of large teeth referable to Polyptychodon and of large pliosaur vertebrae in the Albian Sables verts of the eastern Paris Basin does suggest, however, that they may belong to the same taxon.
Conclusions
Although sauropod remains do occur in the Albian of the eastern Paris Basin (see Buffetaut, 1995, for a review), and have recently been reported from the Sables verts (Buffetaut, 2002), the vertebra from Mesnil-Saint-Père clearly does not belong to a dinosaur, and it obviously cannot be used to support the conclusions of Knoll et al. (2000) on the possible occurrence of brachiosaurids in the Albian of Europe. However, its identification as a dorsal vertebra of a large pliosaurid raises the question of the plesiosaurs and pliosaurs from the Sables verts of the eastern Paris Basin. A certain amount of material is present in the collections of various museums (notably in Lille, Nancy and Charleville-Mézières), but has received little attention since the beginning of the 20th century. A revision of these marine reptiles is clearly needed.
Acknowledgements
We thank Dr Leslie Noè (Sedgwick Museum, Cambridge) for information (including photographs) about the vertebrae referred to Polyptychodon by Seeley (1876). Thanks also to the referees (Marie-Céline Buchy, Mike Everhart, Jean Le Loeuff, Jean-Michel Mazin and Leslie Noè) for their useful comments.
Bibliographic references
Allain R. & Pereda Suberbiola X. (2003).- Dinosaurs of France.- Comptes Rendus Palevol, Paris, vol. 2, no. 1, p. 27-44.
Bardet N. & Godefroit P. (1995).- Plesiosaurus houzeaui Dollo, 1909 from the Upper Campanian of Ciply (Belgium) and a review of the Upper Cretaceous plesiosaurs from Europe.- Bulletin de l'Institut royal des Sciences naturelles de Belgique (Sciences de la Terre), Bruxelles, vol. 65, p. 179-186.
Barrois C. (1875).- Les reptiles du terrain crétacé du N.E. du Bassin de Paris.- Bulletin scientifique, historique et littéraire du Département du Nord et des Pays voisins, Lille, vol. 7, p. 73-83.
Buffetaut E. (1995).- Dinosaures de France.- BRGM, Orléans, 144 p.
Buffetaut E. (2002).- New data from old finds: the dinosaurs from the Early Cretaceous Greensand ("Sables verts") of the eastern Paris Basin. In: 7th European Workshop on Vertebrate Palaeontology, Sibiu (Romania).- Abstracts Volume, Ars Docendi Publishing House, Bucharest, p. 9.
Corroy G. (1922).- Les reptiles néocomiens et albiens du Bassin de Paris.- Comptes Rendus de l'Académie des Sciences, Paris, t. 174, p. 1192-1194.
Janensch W. (1950).- Die Wirbelsäule von Brachiosaurus brancai.- Palaeontographica Supplementbände, Stuttgart, vol. VII, p. 27-93.
Kear B.P. (2003).- Cretaceous marine reptiles of Australia: a review of taxonomy and distribution.- Cretaceous Research, vol. 24, no. 3, p. 277-303.
Knoll F., Colleté C., Dubus B. & Petit J.-L. (2000).- On the presence of a sauropod dinosaur (Saurischia) in the Albian of Aube (France).- Geodiversitas, Paris, vol. 22, no. 3, p. 389-394.
Knoll F., Colleté C., Dubus B. & Petit J.-L. (2002).- De la présence d'un dinosaure sauropode (Saurischia) dans l'Albien de l'Aube.- Bulletin Annuel de l'Association Géologique Auboise, Troyes, vol. 23, p. 27-33.
McIntosh J.S. (1990).- Sauropoda. In: Weishampel D.B., Dodson P. & Osmolska H. (eds.), The Dinosauria.- University of California Press, Berkeley - CA, p. 345-401.
O’Keefe F.R. (2001).- A cladistic analysis and taxonomic revision of the Plesiosauria (Reptilia: Sauropterygia).- Acta Zoologica Fennica, Helsinki, vol. 213, p. 1-63.
Owen R. (1840-1845 [1841]).- Odontography.- Hippolyte Baillière, London, 655 p.
Romer A.S. & Lewis A.D. (1959).- A mounted skeleton of the giant plesiosaur Kronosaurus.- Breviora, Cambridge - MA, vol. 112, p. 1-15. URL: https://www.biodiversitylibrary.org/part/45
Sauvage H.E. (1882).- Les reptiles trouvés dans le Gault de l'Est du Bassin de Paris.- Mémoires de la Société géologique de France (3éme série), Paris, t. II, no. 4, p. 1-43.
Sauvage H.E. (1903).- Sur la présence du genre Polyptychodon dans les Sables verts de la Meuse.- Bulletin de la Société d'Histoire naturelle d'Autun, vol. 16, p. 321-323.
Seeley H.G. (1876).- On an associated series of cervical and dorsal vertebrae of Polyptychodon, from the Cambridge Upper Greensand, in the Woodwardian Museum of the University of Cambridge.- Quarterly Journal of the Geological Society of London, vol. 32, p. 433-436.
Schröder H. (1885).- Saurierreste aus der baltischen oberen Kreide.- Jahrbuch der königlich Preussischen geologischen Landesanstalt und Bergakademie zu Berlin, Jahr 1884, p. 293-333.
Schumacher B.A. & Everhart M.J. (2005).- A stratigraphic and taxonomic review of plesiosaurs from the old "Fort Benton Group" of Central Kansas: a new assessment of old records.- Paludicola, Rochester - NY, vol. 5, p. 33-54.
Storrs G.W. (1991).- Anatomy and relationships of Corosaurus alcovensis<i/> (Diapsida: Sauropterygia) and the Triassic Alcova Limestone of Wyoming.- Bulletin of the Peabody Museum of Natural History, New Haven - CT, vol. 44, p. 1-151.
Upchurch P., Barrett P.M. & Dodson P. (2004).- Sauropoda. In: Weishampel D.B., Dodson P. & Osmolska H. (eds.), The Dinosauria.- Second Edition. University of California Press, Berkeley - CA, p. 259-322.
Unwin D.M. (2001).- An overview of the pterosaur assemblage from the Cambridge Greensand (Cretaceous) of Eastern England.- Mitteilungen des Museums für Naturkunde Berlin, Geowissenschaftliche Reihe, vol. 4, p. 189-221.
Weishampel D.B., Barrett P.M., Coria R.A., Le Loeuff J., Xu X., Zhao X., Sahni A., Gomani E.P. & Noto C.R. (2004).- Dinosaur distribution. In: Weishampel D.B., Dodson P. & Osmolska H. (eds.), The Dinosauria.- Second Edition. University of California Press, Berkeley - CA, p. 517-606.
Welles S.P. & Slaughter B.H. (1963).- The first record of the plesiosaurian genus Polyptychodon (Pliosauridae) from the New World.- Journal of Paleontology, Lawrence - KS, vol. 37, no. 1, p. 131-133.
Williston S.W. (1903).- North American plesiosaurs. Part I.- Field Columbian Museum (Geological Series), Chicago - IL, Publ. 73, , vol. 2, no. 1, p. 1-79.
Figures
 
PLEASE DO NOT EMBED A FIGURE IN THE TEXT
	
Figure 1: Dorsal vertebra of a large pliosaur from the Albian of Mesnil-Saint-Père, Aube (J.-L. Petit collection, Troyes), in anterior (A), left lateral (B), posterior (C) and ventral (D, anterior end to the left) views.
PLEASE DO NOT EMBED A FIGURE IN THE TEXT
	
Figure 2: Dorsal vertebra referred by Seeley (1876) to Polyptychodon, from the Cambridge Greensand (Sedgwick Museum, Cambridge, CAMSM B57396), in right lateral (A), anterior (B) and ventral (C, anterior end to the left) views. Photos courtesy of Dr Leslie Noè (Sedgwick Museum).