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A New Polycotylid Plesiosaur from the Late Cretaceous (Turonian) of Morocco

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A New Polycotylid Plesiosaur from the Late Cretaceous (Turonian) of Morocco C. R. Palevol 2 (2003) 307–315 Systematic Palaeontology (Vertebrate Palaeontology) A new polycotylid plesiosaur from the Late Cretaceous (Turonian) of Morocco Nathalie Bardet a,*, Xabier Pereda Suberbiola a,b, Nour-Eddine Jalil c a Département « Histoire de la Terre », UMR 8569 du CNRS, Muséum national d’histoire naturelle, 8, rue Buffon, 75005 Paris, France b Departamento de Estratigrafía y Palaeontología, Facultad de Ciencias, Universidad del País Vasco/EHU, Apartado 644, 48080 Bilbao, Spain c Laboratoire de paléontologie, département des sciences de la Terre, faculté des sciences Semlalia, université Cadi-Ayyad, bd Moulay-Abdellah, PO Box 2390, 40000 Marrakech, Morocco Received 28 April 2003; accepted 1 July 2003 Presented by Philippe Taquet Abstract A new Polycotylidae (Plesiosauria) from the Late Cretaceous (Turonian) of southern Morocco is described. Thililua longicollis gen. et sp. nov. is based on a complete cranium and 37 associated vertebrae. It differs from other polycotylids in several cranial and especially vertebral characters, such as the occurrence of 30 cervical vertebrae, whose centra are nearly as long as high and bear laterally longitudinal ridges. Thililua is the first polycotylid hitherto found in Africa and under subtropical palaeolatitudes. To cite this article: N. Bardet et al., C. R. Palevol 2 (2003). © 2003 Académie des sciences. Published by Éditions scientifiques et médicales Elsevier SAS. All rights reserved. Résumé Un nouveau plésiosaure polycotylidé du Crétacé supérieur (Turonien) du Maroc. Un nouveau Polycotylidae (Plesiosauria) provenant du Crétacé supérieur (Turonien) du Sud du Maroc est décrit. Thililua longicollis gen. et sp. nov. est basé sur un crâne complet et 37 vertèbres associées. Il diffère des autres polycotylidés par des caractères crâniens et surtout vertébraux, tels que la présence de 30 vertèbres cervicales dont les centres sont presque aussi longs que hauts et portent latéralement des crêtes longitudinales. Thililua est le premier polycotylidé trouvé jusqu’à présent en Afrique et sous des paléolatitudes subtropicales. Pour citer cet article : N. Bardet et al., C. R. Palevol 2 (2003). © 2003 Académie des sciences. Published by Éditions scientifiques et médicales Elsevier SAS. All rights reserved. Keywords: Plesiosauria; Polycotylidae; Late Cretaceous; Turonian; Morocco; New taxon Mots clés : Plesiosauria ; Polycotylidae ; Crétacé supérieur ; Turonien ; Maroc ; Nouveau taxon * Corresponding author. E-mail address: [email protected] (N. Bardet). © 2003 Académie des sciences. Published by Éditions scientifiques et médicales Elsevier SAS. All rights reserved. doi:10.1016/S1631-0683(03)00063-0 308 N. Bardet et al. / C. R. Palevol 2 (2003) 307–315 Version française abrégée calcaire du Cénomanien–Turonien, Crétacé supérieur, Turonien inférieur [10]. Introduction Diagnose. Polycotylidé caractérisé par un prémax- illaire portant des processus étendus latéralement entre Les Polycotylidae sont des plésiosaures à cou court, les narines et les foramens frontaux ; des orbites connus durant le Crétacé et trouvés essentiellement en régulièrement ovales sans processus supraorbitaires ; Amérique du Nord [4,9,19,20]. La monophylie des cinq dents prémaxillaires, au moins 22 maxillaires et Polycotylidae, bien que longtemps mise en doute, est 29 dentaires ; une symphyse mandibulaire portant actuellement appuyée par plusieurs travaux [1,4,13]. 15 paires de dents ; 30 vertèbres cervicales dont les Classiquement inclus au sein des pliosaures [3,18], les centres sont presque aussi longs que hauts et qui por- polycotylidés sont de nos jours considérés comme tent latéralement des crêtes longitudinales qui sont étant plus proches des plésiosaures élasmosauridés à dédoublées de la 19e à la 22e vertèbre. long cou [1,4,5,13], une hypothèse déjàémise il y a prèsd’un siècle [20]. Discussion Contexte géologique Thililua longicollis est rapporté aux Polycotylidae Le spécimen (MNHGr.PA.11710, Muséum (sensu [4,5,13]) de par la combinaison des caractères d’Histoire naturelle de Grenoble) provient de la région suivants : prémaxillaire dont le processus dorsal est de Goulmima, Province d’Er-Rachidia, Sud du Maroc, étendu postérieurement et sépare les frontaux (par- bien que sa provenance exacte reste inconnue (Fig. 1). allélisme avec les pliosauridés) ; contact étendu entre Les gisements localisés lors de travaux de terrain près le maxillaire et le squamosal ; jugal formant une barre des villages de Tadirhourst et Asfla ont livré une riche subrectangulaire horizontale (comme chez les élasmo- faune de vertébrés marins [2,6], ainsi que des ammo- sauridés) ; splénial inclu dans la symphyse mandibu- nites. Les fossiles sont préservés dans des nodules laire. ovoïdes situés au sommet (unité 4) de la barre calcaire Thililua longicollis partage également avec tous les du Cénomanien–Turonien [7,8].L’unité 4 est d’âge polycotylidés(Dolichorhynchops, Georgiasaurus, Turonien inférieur (présence de l’ammonite Mammites Polycotylus, Sulcusuchus, Trinacromerum) notamment) et correspond à un environnement de [4,5,11,14,16,17,19,20], sauf Edgarosaurus muddi plate-forme ouverte, lié au maximum de la transgres- (rapportée provisoirement aux polycotylidés, voir [9]), sion cénomano-turonienne [10]. un crâne gracile à long museau, dont la partie préorbi- Systématique taire représente environ 60% de la longueur totale et une dentition uniforme sans dents caniniformes. De Plesiosauria de Blainville, 1835 plus, le prémaxillaire contacte le pariétal et il n’y a pas Polycotylidae Cope, 1869 de foramen pinéal, comme chez Dolichorhynchops Thililua gen. nov. osborni, Trinacromerum bentonianum et Sulcusuchus Étymologie. De Thililua, dieu aquatique dans la erraini (à la différence d’Edgarosaurus) [4,9,11]. Thi- mythologie berbère [12]. lilua possède enfin un foramen frontal au-dessus de Espèce type. Thililua longicollis. l’orbite et une symphyse mandibulaire longue (environ Diagnose. Comme pour l’espèce type. la moitié de la longueur totale), comme chez Dolicho- Thililua longicollis sp. nov. rhynchops et Trinacromerum [5]. Ce dernier caractère Étymologie. En référence à son long cou. est aussi présent chez Polycotylus latipinnis [16].En Holotype. MHNGr.PA.11710, crâne et mandibule attendant une révision complète du groupe et en subcomplets et 37 vertèbres associées articulées, com- l’absence de diagnose consensuelle (comparer [4] et prenant les séries cervicale et pectorale complètes ainsi [9]), l’ensemble de ces caractères est ici considéré que les quatre premières vertèbres dorsales (Figs. 2 et provisoirement comme caractéristique des Polycotyl- 3; Tableaux 1 et 2). idae ou de clades moins inclusifs. Localité et horizon type. Près de Goulmima, prov- Thililua a des dents longues et finement striées, ince d’Er-Rachidia, Sud du Maroc ; unité 4, Barre comme chez Dolichorhynchops et Sulcusuchus, à la N. Bardet et al. / C. R. Palevol 2 (2003) 307–315 309 différence de celles de Polycotylus, Trinacromerum et Atlas of Morocco (Fig. 1). Though its exact location Edgarosaurus, qui sont robustes et fortement striées remains unknown, fieldwork has allowed us to locate [4,9,11]. several fossiliferous localities near the villages of Ta- Thililua diffère de Dolichorhynchops et de Trinac- dirhourst and Asfla, north of Goulmima, Er-Rachidia romerum par l’absence d’un processus supraorbitaire, Province. These sites have yielded a rich fauna of la présence d’un contact prémaxillaire–pariétal situé marine vertebrates, including actinopterygians [6], postérieurement aux orbites et une facette costale marine reptiles [2], as well as ammonites. The fossils portée par le centrum de l’axis seulement [4,5,19].De are preserved in ovoid calcareous nodules that occur plus, son suspensorium est vertical, contrairement à near the top of a Cenomanian–Turonian limestone bar, Trinacromerum [4]. a reference level in North Africa [7,8]. The fossilifer- Finalement, Thililua longicollis diffère des autres ous nodules are concentrated in Unit 4 of Ferrandini et polycotylidés par les autapomorphies mentionnées al. [10], which is Early Turonian in age according to dans la diagnose. the ammonite association (mainly Mammites). This D’un point de vue paléobiogéographique, Thililua unit corresponds to an open platform environment re- est le premier polycotylidé trouvé jusqu’à présent en lated to the maximum of the Cenomanian–Turonian Afrique et sous des paléolatitudes subtropicales. transgressive phase [10]. 1. Introduction 3. Systematic palaeontology Polycotylidae is a group of short-necked plesiosaurs Plesiosauria de Blainville, 1835 exclusive of the Cretaceous. Their remains are essen- Polycotylidae Cope, 1869 tially known from North America [4,9,19,20],but Thililua gen. nov. more fragmentary material has also been described in Etymology. From Thililua, aquatic god in the Ber- Asia [15,17], South America [11], Australia and New- ber mythology [12]. Zealand [15]. They stratigraphically range from the Type species. Thililua longicollis. Aptian–Albian to the Maastrichtian [15]. The validity Diagnosis. As for type species. of the Polycotylidae is still under debate, but its mono- Thililua longicollis sp. nov. phyly has been supported by recent works [1,4,13]. The Polycotylidae currently includes Polycotylus Cope, 1869 [4,16,20], Trinacromerum Cragin, 1888 [4], Dolichorhynchops Williston, 1902 [4,5,19], Geor- giasaurus Otschev, 1977 [14,17], Sulcusuchus Gas- parini & Spalletti, 1990 [11] and, tentatively, Edgaro- saurus Druckenmiller, 2002 [9]. Concerning their phylogenetic relationships within the
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