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An Early Spinosaurid Dinosaur from the Late Jurassic of Tendaguru (Tanzania) and the Evolution of the Spinosaurid Dentition

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An Early Spinosaurid Dinosaur from the Late Jurassic of Tendaguru (Tanzania) and the Evolution of the Spinosaurid Dentition An early spinosaurid dinosaur from the Late Jurassic of Tendaguru (Tanzania) and the evolution of the spinosaurid dentition Eric Buffetaut1 1 Centre National de la Recherche Scientifique, UMR 8538, Laboratoire de Géologie de l’Ecole Normale Supérieure, 24 rue Lhomond, 75231 Paris Cedex 05, France. E-mail : [email protected] Abstract - An isolated theropod tooth from the terminal Jurassic of the Tendaguru (Tanzania), originally referred to La- brosaurus (?) stechowi, is described as the type specimen of a new genus and species of spinosaurid theropod. It shows an enamel ornamentation reminiscent of Baryonyx and allied forms, but differs from all other known spinosaurids by the large size of the denticles borne by the carinae. The new taxon is the earliest currently known spinosaurid. It sheds some light on aspects of dental evolution in spinosaurids, which seems to have been characterised by reduction of the denticles. Keywords : Dinosauria, Theropoda, Spinosauridae, teeth, Late Jurassic, Tendaguru, Tanzania. Un dinosaure spinosauridé primitif du Jurassique supérieur du Tendaguru (Tanzanie) et l’évolution de la denture des spinosauridés - Une dent de théropode isolée du Jurassique terminal du Tendaguru (Tanzanie), rapportée à l’origine à Labrosaurus (?) stechowi, est décrite comme spécimen type d’un nouveau genre et d’une nouvelle espèce de thé- ropode spinosauridé. Elle montre une ornementation de l’émail rappelant Baryonyx et des formes apparentées, mais diffère de tous les autres spinosauridés connus par la grande taille des denticules portés par les carènes. Ce nouveau taxon est le plus ancien spinosauridé connu. Il éclaire certains aspects de l’évolution des dents chez les spinosauridés, qui semble avoir été caractérisée par la réduction des denticules. Mots-clés : Dinosauria, Theropoda, Spinosauridae, dents, Jurassique supérieur, Tendaguru, Tanzanie. INTRODUCTION examination of that material at the Museum für Naturkunde in Berlin, carried out in 2007. Among the many fossils collected by the German expeditions of 1909-1912 from the Late Jurassic dinosaur- bearing beds of what was then the south-eastern part of Institutional abbreviations Deutsch-Ostafrika (today Tanzania) were numerous iso- DMR : Department of Mineral Resources, Thailand. lated theropod teeth (230 according to Janensch, 1925), MB : Museum für Naturkunde der Humboldt-Universität, which were described by Janensch, first in a short prelimi- Berlin, Germany. NHMUK : Natural History Museum, Lon- nary paper (Janensch, 1920) and then in a large monograph don, UK. UT : Geology Department, University Al Fateh, (Janensch, 1925) on the “coelurosaurs and theropods” from Tripoli, Libya. the Tendaguru Beds. The precise identification of isolated theropod teeth is fraught with difficulties, and Janensch attempted both to provide identifications for as much as HISTORY OF PREVIOUS RESEARCH possible of the Tendaguru material and to describe distinct morphotypes in great detail. Besides teeth of various sizes The theropod teeth from Tendaguru bearing an or- showing the “usual” smooth, blade-like theropod pattern, namentation of ridges and grooves on the lingual face (and for which Janensch provided tentative identifications, the sometimes the labial face as well) were tentatively referred Berlin collection includes a few specimens which show by Janensch (1920, 1925) to the poorly known genus Labro- more unusual characters, including a more or less marked saurus Marsh, 1879, as Labrosaurus (?) stechowi. Janensch enamel ornamentation. Among these are teeth which show further distinguished several morphological “types” among considerable similarities with those of some spinosaurid this material. Janensch’s referral of these Tendaguru teeth to theropods (Fowler, 2007 ; Buffetaut, 2008). The present Labrosaurus was based on similarities, notably in enamel paper describes a new spinosaurid taxon on the basis of a re- ornamentation, with a tooth from the Morrison Formation ORYCTOS vol. 10, 2013, publié en ligne le 23 janvier 2012 1 described by Marsh (1896) as Labrosaurus sulcatus. The ge- “Obere Sauriermergel” by Janensch and other members of nus Labrosaurus, originally erected on postcranial elements, the German Tendaguru expeditions. In their recent revision has had an intricate taxonomic history, as Marsh applied it, of the local stratigraphy, Bussert et al. (2009) consider these under various specific names, to other specimens from North “Upper Saurian Marls” as a member of the Tendaguru Forma- America, including the above-mentioned tooth (see Madsen tion and designate them as the Upper Dinosaur Member. The & Welles, 2000, for a review). Madsen and Welles (2000) Upper Dinosaur Member is unconformably overlain by the considered that the North American material attributed to Rutitrigonia bornhardti-schwarzi Member, which contains Labrosaurus in fact belongs either to Allosaurus or to Cera- Early Cretaceous marine fossils, and it overlies the marine tosaurus, the “Labrosaurus sulcatus” tooth being in fact an Indotrigonia africana Member, which is probably Tithonian anterior tooth of Ceratosaurus. in age (Bussert et al., 2009). The Upper Dinosaur Member In their revision of the genus Ceratosaurus, Mad- has been variously considered as latest Jurassic or earliest sen and Welles (2000) discussed material from Tendaguru, Cretaceous in age (see review in Bussert et al., 2009). Bus- including the cranial and post-cranial elements described sert et al. (2009, p. 165) consider that “more work is needed by Janensch (1925) as Ceratosaurus (?) roechlingi and the to decide unambiguously whether the Upper Dinosaur Mem- teeth designated as Labrosaurus (?) stechowi. On the basis ber is wholly Late Jurassic or, in part, Early Cretaceous” ; in mainly of a quadrate, they concluded that a large species of their correlation chart, they consider it as Tithonian. Ceratosaurus was probably present at Tendaguru, and that Other ornamented theropod teeth from Tendag- the Labrosaurus (?) stechowi teeth, showing an ornamenta- uru referred by Janensch (1920, 1925) to Labrosaurus (?) tion similar to that of anterior teeth of Ceratosaurus, could stechowi come from what he called the “Mittlere Saurier- be referred to it. In their review of the Ceratosauria, Carrano mergel”, referred to as the Middle Dinosaur Member of the and Sampson (2008) discussed Ceratosaurus (?) roechlingi Tendaguru Formation by Bussert et al. (2009). This member and concluded that a basal ceratosaur was indeed present at underlies the Tithonian Indotrigonia africana Member, and Tendaguru, but they did not mention the teeth identified as overlies the Oxfordian to Kimmeridgian Nerinella Member. Labrosaurus (?) stechowi. Bussert et al. (2009) consider it as late Kimmeridgian in age. On the basis of Janensch’s work, Fowler (2007) suggested that the ornamented theropod teeth from Tenda- SYSTEMATIC DESCRIPTION guru should be reassessed by comparison with baryonych- ine teeth, and that they may support a ceratosaurian origin Dinosauria Owen, 1842 for baryonychines. Independently, after revising Janensch’s Theropoda Marsh, 1881 material at the Museum für Naturkunde in Berlin, Buffetaut Spinosauridae Stromer, 1915 (2008) concluded that while some of these teeth probably Ostafrikasaurus, n.g. belonged to ceratosaurs, others could be referred to an early Ostafrikasaurus crassiserratus, n. sp. spinosaurid. The present paper formalises this identifica- tion by erecting a new taxon for the spinosaurid teeth from Diagnosis : A spinosaurid dinosaur with teeth which Tenda guru. Although isolated theropod teeth have often are moderately compressed labiolingually. The enamel is been considered as having little systematic value, various finely wrinkled and bears ridges which are more numerous studies (e.g. Currie et al., 1990 ; Baszio, 1997 ; Sankey, 2001 ; and more pronounced on the lingual face than on the labial Sankey et al., 2002 ; Smith et al., 2005 ; Longrich, 2008 ; face. The anterior and posterior carinae are serrated. Differs Sankey, 2008 ; Cillari, 2010) have shown that they can be from all other spinosaurids in the relatively large size of the diagnostic when sufficient attention is paid to morphology denticles (2 to 4 denticles per mm). and enamel ornamentation. Spinosaurids have especially Type specimen : an isolated tooth, MB.R.1084, distinctive teeth that generally differ significantly from “Typus a” of Janensch (1925). those of other theropods, and among the group, differences Referred specimen : isolated tooth MB.R.1091, in morphology and ornamentation allow to distinguish taxa “Typus d” of Janensch (1925), from the Middle Dinosaur (Medeiros, 2006 ; Buffetaut et al., 2008). It therefore seems Member of the Tendaguru Formation. legitimate to erect a new taxon of early spinosaurid dinosaur Locus typicus : Tendaguru, southeastern Tanzania, on the basis of its distinctive teeth. quarry Om (Janensch, 1925). Stratum typicum : “Oberer Sauriermergel” (Jan- ensch, 1925), Upper Dinosaur Member of the Tendaguru GEOGRAPHICAL AND GEOLOGICAL SETTING Formation of Bussert et al. (2009). Derivatio nominis : Ostafrikasaurus, from the The specimens mentioned in the present paper were name used by the Germans ([Deutsch]-Ostafrika) for the collected on or in the vicinity of Tendaguru hill, in southeast- part of East Africa including Tendaguru at the time of the ex- ern Tanzania (see Bussert et al., 2009 for a recent detailed peditions of the Berlin Museum, and saurus, from σαυρóς, map of the area). Greek for a lizard ; crassiserratus, from the Latin
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