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A Basal Sauropod Dinosaur from the Early Jurassic of Morocco

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A Basal Sauropod Dinosaur from the Early Jurassic of Morocco Published in Comptes Rendus Palevol 3(3): 199-208, 2004 which should be cited to reference this work A basal sauropod dinosaur from the Early Jurassic of Morocco Ronan Allain a,b,*, Najat Aquesbi c, Jean Dejax a, Christian Meyer d, Michel Monbaron e, Christian Montenat f, Philippe Richir a, Mohammed Rochdy c, Dale Russell g, Philippe Taquet a a Laboratoire de paléontologie, département « Histoire de la Terre », Muséum national d’histoire naturelle, UMR 8569 CNRS, 8, rue Buffon, 75005 Paris, France b Université Rennes 1, Geosciences, Campus de Beaulieu, av. du Général-Leclerc, 35042 Rennes cedex, France c Institut Agdal, ministère de l’Énergie et des Mines, BP 6 208, Rabat, Morocco d Naturhistorisches Museum Basel, Augustinergasse 2, CH-4001 Basel, Switzerland e Département de Géosciences, université de Fribourg, chemin du Musée 4 , CH-1700 Fribourg, Switzerland f Institut géologique Albert-de-Lapparent (IGAL), 13, bd de l’Hautil, 95092 Cergy-Pontoise, France g North Carolina Museum of Natural Sciences, and Department of Marine, Earth and Atmospheric Sciences, North Carolina State University, Box 8208, Raleigh, NC 27695, USA Received 12 February 2004; accepted 15 March 2004 Available online 30 April 2004 Abstract Continental strata of Early Jurassic age are seldom exposed, and little is known of the history of sauropod dinosaurs prior to the Middle Jurassic radiation of neosauropods. Well-preserved skeletons and skulls have not been recovered from strata older than the Middle Jurassic. Here we report, in the Early Jurassic of the Moroccan High Atlas, the discovery of the skeleton, including cranial material, of a new vulcanodontid sauropod. Tazoudasaurus naimi n.g., n.sp. represents with Vulcanodon the sister group of the eusauropods and the most complete basal sauropod material available to date. Résumé Parce que les dépôts continentaux du Jurassique inférieur affleurent rarement, l’histoire des dinosaures sauropodes est très mal connue avant la radiation des néosauropodes au Jurassique moyen. Les premiers crânes de sauropodes associés à des restes osseux bien préservés datent ainsi du Jurassique moyen. Nous rapportons ici la découverte, dans le Jurassique inférieur du Haut Atlas marocain, du squelette et du crâne d’un nouveau sauropode. Tazoudasaurus naimi n.g., n.sp. est le groupe frère de Vulcanodon et le sauropode basal le plus complet connu à l’heure actuelle. * Corresponding author. E-mail address: [email protected] (R. Allain). 1 Keywords: Dinosaurs; Sauropoda; Early Jurassic; Morocco; Phylogeny Mots clés : Dinosaures ; Sauropoda ; Jurassique inférieur ; Maroc ; Phylogénie Version française abrégée chevrons distaux bifurqués avec des processus anté- rieurs et postérieurs non fusionnés ; présence d’une Nous décrivons ici le squelette d’un nouveau sauro- crête sur la face latérale de l’extrémité proximale de la pode, incluant des restes crâniens, récolté dans des fibula. Tazoudasaurus présente aussi une combinaison pélites recouvrant des bancs carbonatés marins datés unique de synapomorphies de sauropode (préarticu- de la fin du Jurassique inférieur, de la nappe de Toun- laire étendu dorsoventralement ; couronne dentaire vue doute du Haut Atlas, au Maroc [9,17,24] (Fig. 1). Des en coupe, en forme de D ; émail dentaire plissé ; restes de dinosaures théropodes de grande et de centrum des vertèbres cervicales opisthocoele ; large moyenne taille, mais d’affinités pour l’instant incertai- épine neurale dorsale ; quadrupédie obligatoire ; crête nes, ainsi que des débris de végétaux, essentiellement deltopectorale de l’humérus réduite, rapport constitués de restes de fougères, de cycadales et de humérus/fémur supérieur à 0,60 ; diaphyse fémorale conifères, ont été découverts, associés aux restes de elliptique en coupe ; absence de foramens à la base du sauropode. processus ascendant de l’astragale ; phalange unguéale du premier doigt du pied en forme de faucille) et de 1. Paléontologie systématique symplésiomorphies de sauropodomorphe (20 dents dentaires, avec des denticules sur les marges antérieure Dinosauria Owen, 1842 et postérieure de la couronne dentaire ; extrémité anté- Saurischia Seeley, 1888 rieure du dentaire peu étendue dorsoventralement ; Sauropodomorpha Huene, 1932 rangées dentaires convergentes en forme de V ; ab- Sauropoda Marsh, 1878 sence de fosse quadratique postérieure ; présence d’un Vulcanodontidae Cooper, 1984 petit trochanter sur le fémur ; surface plantaire des Tazoudasaurus naimi gen. et sp. nov. phalanges unguéales des doigts II et III aplatie). Étymologie. Le nom de genre dérive de celui de la Tazoudasaurus se différencie de Barapasaurus localité type Tazouda et de sauros, lézard en grec. Le [13,14] et de Kotasaurus [33,34] du Jurassique infé- nom spécifique vient de naïmi, élancé en arabe, et fait rieur d’Inde, et de Gongxianosaurus [11] du Jurassique référence à la petite taille de l’holotype. inférieur de Chine, sur la base de nombreux caractères anatomiques. Il présente, en revanche, beaucoup plus Holotype. To 2000–1, musée des Sciences de la d’affinités avec Vulcanodon, du Jurassique inférieur du Terre de Rabat, Maroc ; squelette partiellement articulé Zimbabwe [5,23], duquel il diffère cependant par l’ab- comprenant du matériel crânien, qui inclut une mandi- sence de sillon médian sur la face ventrale des vertè- bule gauche complète, carré, jugal, postorbitaire, pa- bres dorsales et par des épines neurales caudales plus riétal, frontal et exoccipital. allongées antéropostérieurement. Une analyse phylo- Autre spécimen. To 2000–2, restes associés d’un génétique incluant 29 taxons, dont les genres maro- spécimen juvénile. cains Tazoudasaurus et Atlasaurus, a été réalisée. Les Localité et Horizon. Douar de Tazouda, près du 235 caractères utilisés dans cette analyse sont tirés de village de Toundoute, dans la province de Ouarzazate, la récente phylogénie de Wilson [30], auxquels nous Haut Atlas, Maroc. Nappe de Toundoute, série conti- avons rajouté le caractère « surface articulaire proxi- nentale détritique du Toarcien, qui recouvre en concor- male des doigts II et III du pied significativement plus dance des carbonates marins datés du début et du large que haute », précédemment considéré comme milieu du Jurassique inférieur. une autapomorphie de Vulcanodon [31]. Ce caractère Diagnose. Sauropode primitif, défini par les autapo- soutient la position de Tazoudasaurus comme groupe morphies suivantes (Fig. 2a–n) : lame osseuse s’éten- frère de Vulcanodon à l’extérieur des Eusauropoda, dant sur la marge postérodorsale du postorbitaire ; dans les six arbres les plus parcimonieux obtenus, 2 après utilisation de la commande mhennig*, bb* du 1. Introduction logiciel Hennig86 [8] (Fig. 3). Les récentes datations effectuées sur les basaltes du plateau du Karoo mon- Sauropod remains were documented from the trent que Vulcanodon serait contemporain de Tazouda- Middle Jurassic strata of the High Atlas Mountains in saurus [7,15], même si de nombreux auteurs conti- Morocco through the descriptions of ‘Cetiosaurus’ nuent à le placer dans le Jurassique basal (Hettangien) mogrebiensis [16] and Atlasaurus imelakei [20,21]. [3–5,10,13,23,28,30,32]. Les deux taxons sont regrou- Recently, we collected a partly articulated sauropod pés ici au sein de la famille monophylétique des Vul- skeleton, including cranial material, from the Early canodontidae, qui exclut les autres sauropodes connus Jurassic of the High Atlas near Toundout, in the Prov- au Jurassique inférieur [5,25,26,32]. ince of Ouarzazate (Fig. 1). In this area, the entire Atlasaurus imlakei du Jurassique moyen (Batho- Jurassic series, decoupled from its base, has been over- nien) du Haut Atlas central [20,21] est le groupe frère thrusted to the south where it overlies the Cretaceous– des Neosauropoda (Fig. 3) [30] et est donc beaucoup Eocene [17]. The two principal fossiliferous sites are plus dérivé que Tazoudasaurus. La succession, dans le near the summit of a detrital series about 300 m thick Haut Atlas, de ces deux taxons montre que des change- that conformably overlies early Lower Jurassic marine ments majeurs interviennent dans la morphologie des carbonates containing algal spheres, and benthic fora- sauropodes durant les 20 millions d’années qui sépa- miniferal and mollusc shells [9,24]. The two sites, rent le Toarcien du Bathonien. Certains de ces change- separated from each other stratigraphically by about ments semblent liés à l’alimentation (perte de la fenê- 30 m, are sedimentologically and palaeobotanically tre mandibulaire externe, robustesse mandibulaire similar. Associated with the sauropod material were accrue au niveau de la symphyse, forme en U de la isolated elements of medium-sized and large theropods symphyse, réduction du nombre de dents), alors que of uncertain affinities. Greenish, channel-filling silts d’autres peuvent être reliés à la locomotion et à la taille and sands are preserved within a red, relatively unfos- du corps (configuration en arche des métacarpiens, siliferous clastic sequence. Submacroscopic woody extension transversale de l’extrémité proximale du ti- debris include homoxylous particles devoid of paren- bia et changement d’orientation de la crête cnémiale, chyma, evenly distributed uniseriate rays, tracheids forme en faucille des phalanges unguéales du pied). bordered by uniseriate pits of abietinian form, and Tazoudasaurus est le sauropode primitif le plus cross-fields with piceoid oculipores bearing narrow, complet connu à ce jour. En tant que tel, il partage de perpendicular slits. The anatomy suggests affinities nombreux
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