GAIA N° 15, LlSBOAlLISBON, DEZEMBRO/DECEMBER 1998, pp. 75-88 (ISSN: 0871-5424) A REVIEW OF EUROPEAN TRIASSIC THEROPODS Oliver W. M. RAUHUT Institut fOr Palaontologie, Freie Universitat Berlin. MalteserstraBe 74-100, Hs D, 0-12249, BERLIN. GERMANY E-mail: [email protected] Axel HUNGERSOHLER Royal Ontario Museum, Departement of Paleobiology. 100 Queen's Park, Toronto, ONTARIO M5S 2C6. CANADA ABSTRACT: The Upper Triassic theropod fossil record from Europe is reviewed in terms of va­ lidity of proposed taxa and the stratigraphical distribution of theropod remains, Only three species can presently be regarded as valid: Liliensternus liliensterni (HUENE), (?) Lilienster­ nus aire/ensis CUNY & GALTON, and Procompsognathus triassicus FRAAS. (?) L. aire/ensis might represent a distinct genus, but more material is needed to confirm this. The genus Syntarsus RAATH, formerly known from southern Africa and North America is described from Europe for the first time. Theropods are first known from the fossil record in Europe in the Norian, and all determinable fossils represent members of the Coelophysoidea, Judged by the rarity oftheirfossil remains, theropods were obviously rather rare elements ofthe Up­ per Triassic terrestrial vertebrate fauna of Europe. RESUME: Le registre fossile des theropodes du Trias superieur europeen est revise en terme a la fois de validite des taxons proposes et de repartition stratigraphiques des restes fossi­ les, Seule trois especes peuvent etre actuellement considerees comme valides: Lilienster­ nus liliensterni (HUENE), (?) Liliensternus aire/ensis CUNY & GALTON et Procompsognathus triassicus FRAAS. (?) L. aire/ensis pourrait en fait appartenir a un nouveau genre, mais des , restes plus complets seraient necessaires pour confirmer cette hypothese, Le genre Syn­ tarsus RAATH, qui n'avaitjusqu'a present ete decrit qu'en Afrique du Sud et en Amerique du Nord, I'est egalement pour la premiere fois en Europe, Les theropodes apparaissent en Eu­ rope a partir du Norien, et tous les fossiles identifiables appartiennent aux Coelophysoidea, Si I'on en croit la pauvrete de leurs restes fossiles, il semble evident que les theropodes re­ presentaient des elements relativement rares au sein des faunes de vertebres terrestres du Trias superieur d'Europe. INTRODUCTION The aim ofthe present paper is to review Triassic theropod records from Europe, in terms of the taxo­ In the light of new discoveries, especially in nomic validity of named species, and the stra­ South America, the early history of theropod dino­ tigraphical distribution of theropod remains in the saurs has been of great interest recent years (e.g. European Triassic, to determine the first appear­ SERENO & NOVAS, 1992). Theropods are one of the ance and taxonomic diversity of theropod dinosaurs most diverse dinosaur groups in the Late Jurassic in the Upper Triassic of Europe. and the Cretaceous, but little is still known about their origin, radiation and early diversification. Espe­ THEROPODS FROM THE TRIASSIC OF cially interesting in this respect is the first radiation of EUROPE theropod dinosaurs, or dinosaurs in general, in the Upper Triassic (e.g. BONAPARTE, 1982; BENTON, In the following section, theropod taxa described 1984, 1993). Our knowledge of Upper Triassic from Europe are reviewed. See Figure 1 forthe geo­ theropods is mainly based on the American fossil graphical occurrences, and Figure 2 for the stra­ record, although theropod remains are known from tigraphical distribution of theropodan remains. the Triassic all overthe world (WEISHAMPEL, 1990). 75 artigos/papers O. WM. RAUHUT & A. HUNGERBOHLER N + Heilbronn I • Pfaffen­ hofeo Halberstadt Heslach~ * Kaltental Dillstedt G ro Ber -# Gleichberg Ttibingen Trossingen I Sigmaringen • I ! , , 1 I 25km , Upper Stubensandstein Middle Stubensandsteio Fig. 1 - Map olthe vertebrate localities mentioned in the text. The map of South Germany is based on BENTON (1994a). Avipes dillstedtianus HUENE, 1932 A. dillstedtianus can only be referred to as probable Archosauria gen. et sp. indet. (see also NORMAN , Age: Ladinian, Middle Triassic. 1990). Occurrence: Grenzdolomit close to Dillstedt, ThOringen, Germany. Dolichosuchus cristatus HUENE, 1932 Comments: A. dillstedtianus was described by Age: Norian, Upper Triassic. HUENE (1932) on the basis of the proximal ends of Occurrence: Lower or middle Stubensandstein, three articulated metatarsals from the lowermost Stuttgart-Kaltental, Baden-WOrttemberg, Germany. Upper Triassic (Lettenkeuper) of Thuringia. Comments: Dolichosuchus cristatus is based The proximal ends of the metatarsals are closely on an isolated tibia from the Stubensandstein of appressed and deeper than wide (see HUENE, 1932: southern Germany (HUENE, 1932). The specimen taf. 1, f. 7). Distally, the shafts of the bones are sepa­ (BMNH 38056) shows a large cnemial crest, and a rated from each other and appear to diverge. lateral ridge for the attachment of the fibula; these As already mentioned by NORMAN (1990), the characters indicate that it represents a theropod. specimen probably represents a digitigrade animal, However, the poor preservation of the element but the metatarsals do not show any characters that makes it generically and specifically indeterminate, allow a referral to the Theropoda. Since there are a so that D. cristatus must be treated as a nomen du­ variety of digitigrade animals in the Upper Triassic, bium. It should be noted, though, that, as already 76 A REVIEW OF THE EUROPEAN TRIASSIC THEROPODS Chronostrat. LithostratIgraphy System Stage SW Germany Thuringia Jurassic Hettangian Lias Lias (X Lias (X ;::: o:s ."",..... ~ u. RMt RMt o:s ,.;::: ~ .... Knolleomergel ~ o:s Dolomit· ."", mergelkeuperl .... Stein· 0 mergelkeuper Z Kieselsandstein U. Gipskeuper <1) ."", Rote Wand I .... Schilfsandstein ~. Schilfsandstein Gipskeuper u L. Gipskeuper L. Lettenkeuper renzbone Muschel· U. Muschelkalk U. Muschelkalk kalk 1 Dolichosuchus cristatus HUENE 3 Syntarsus sp. [JJ] sandstone Tanystrophaeus posthumus HUENE 4 Liliensternus liliensterni (HUENE) pelite 2 Halticosaurus longotarsus HUENE 5 Pterospondylus trielbae JAEKEL I I Procomposagnathus triassicus FRAAS 6 "Zanclodon" cambrensis NEWTON dolomitic 7 (?)Liliensternus airelensis CUNY & GALTON B limestone Fig. 2· Synthesis olthe lithostratigraphy 01 the German Keuper succession and its tentative correlation with the Trias· sic stages. modified Irom AIGNER & BACHMANN (1992). The approximate stratigraphie positions olthe taxa are plotted in the diagrammatic lithostratigraphic section, taxa found outside of Germany are inserted in frames. 77 O. w.M. RAUHUT & A. HUNGERBOHLER mentioned by HUENE (1934), the specimen shows served, and most of the fragments are not identifi­ great similarities to the tibiae of the slightly younger able even as theropods. Furthermore, the original Lilienstemus lilienstemi (HUENE), and also of Dilo­ association of these specimens is very dubious. Ac­ phosaurus wetherilli (WELLES, 1984). This suggests cording to HUENE (1907-8), the remains were found that it probably represents a member of the Coelo­ " ... together with Sellosaurus Fraasi and Teratosau­ physoidea ( = Dilophosaurus + Coelophysidae; rus (?) minor [both synonyms of Sellosaurus gracilis; HOLTZ, 1994). note by the authors] ... in a marly layer intercalated in the Stubensandstein ... " (HUENE, 1907-8: 231; Halticosaurus longotarsus HUENE, 1907-8 translated by O.R.), thus, it seems quite possible, Age: Norian, Upper Triassic. that some of the material might represent a pro­ sauropod. Indeed, the collection number also in­ Occurrence: Middle Stubensandstein, Weil1er cludes a skull of Sellosaurus HUENE (SMNS Steinbruch, Pfaffenhofen, Baden-WOrtlemberg, 12353a). The only remains that can be referred to Germany. the Theropoda with some certainty are the femoral Comments: Halticosaurus longotarsus, the type fragments. One of them shows a spike-like lesser species of the genus Halticosaurus, was described trochanter and a significantly downturned femoral by HUENE (1907-8) on the basis of some fragments head (Fig. 3), both characters also found in coelo­ from the Stubensandstein of Pfaffenhofen (all bear­ physoids. Although H. longotarsus must be treated ing the collection number SMNS 12353). The mate­ as a nomen dubium, the type series might therefore rial originally comprised a fragmentary dentary, include remains of a coelophysoid theropod. parts of cervical, dorsal, sacral, and caudal verte­ HUENE (1921b) later referred some fragmentary brae, fragments of a humerus, an ilium and two remains from the Norian of Halberstadt to the same femora, and a complete metatarsal. Only one cervi­ genus as cf. Halticosaurus longotarsus. This mate­ cal vertebra (HUENE, 1907-8: taf. 97, f. 4), a dorsal rial (now in the HMN) is extremely fragmentary as vertebral centrum (op. cit.: taf. 97, f. 7), anotherfrag­ well, and none of it can even be shown to be theropo­ mentary vertebral centrum (op. cit.: taf. 97, f. 6), the dan with any certainty. two femoral fragments (op. cit.: taf. 97, f. 1, 2) and the metatarsus (op. cit.: taf. 97. f. 9) could be located cf. Halticosaurus orbitoangulatus in the SMNS. All of the material is very badly pre- HUENE, 1932 Age: Norian, Upper Triassic. Occurrence: Middle Stubensandstein, Weil1er Stein bruch, Pfaffenhofen, Baden-WOrtlemberg, Germany. Comments: This taxon is based on a partial skull (SMNS 12353b) from the Pfaffenhofen quarry in the middle Stubensandstein of southern Germany. The skull is badly crushed, and the anterior elld