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Home , Ceratosauria, Coelophysidae, Liliensternus, Paratypothorax, Procompsognathus

GAIA N° 15, LlSBOAlLISBON, DEZEMBRO/DECEMBER 1998, pp. 75-88 (ISSN: 0871-5424)

A REVIEW OF EUROPEAN 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 can presently be regarded as valid: liliensterni (HUENE), (?) Lilienster­ nus aire/ensis CUNY & GALTON, and triassicus FRAAS. (?) L. aire/ensis might represent a distinct , 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 , and all determinable fossils represent members of the , 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 (e.g. European Triassic, to determine the first appear­ SERENO & NOVAS, 1992). Theropods are one of the ance and taxonomic diversity of theropod most diverse groups in the Late in the Upper Triassic of Europe. and the , 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 -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 , 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 . Since there are a so that D. cristatus must be treated as a nomen du­ variety of digitigrade 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 ( = + ; 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 (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 and mos'. of the skull roof is missing. The specimen has been described in some detail by HUENE (1932), who re­ ferred this species to the family Podokesauridae within the Theropoda. However, the strongly anteri­ orly tapering antorbital fenestra closely resembles the condition seen in the sphenosuchian crocodile Saltoposuchus (see HUENE, 1921 a; SERENO & WILD, 1992). Moreover, the teeth show an almost circular cross section at the base of the crown and longitudinal striations, but they lack well defined cut­ ting edges and serrations. This is a condition often found in crocodylomorphs, but very rarely in thero­ pods. Thus, since the specimen also lacks any clear theropod synapomorphies, Halticosaurus orbitoan­ gulatus probably represents a sphenosuchian Fig. 3 - Halticosaurus longotarsus HUENE; SMNS crocodile, rather than a theropod. 12353; proximal end of righl femur, part of Ihe Iype series; Stubensandstein, Pfaffenhofen, Germany; cranial view. Liliensternus liliensterni (HUENE, 1934) Nole Ihe downlurned femoral head and Ihe spike-like lesser lrochanler (II). Scale bar indicales 1 cm. Age: Norian, Upper Triassic.

78 A REVIEW OF THE EUROPEAN TRIASSIC THEROPODS

Occurrences: Knollenmergel, Thuringen; Comments: The holotype of this species com­ ?Trossingen, Wurttemberg, Germany; ? Frick, Swit­ prises a fragmentary associated zerland. and parts of the pelvis (Caen University, unnum­ b~red) : The material was originally described as Comments: Liliensternus liIiensterni is the best Halticosaurus sp. by LARSONNEUR & LAPPARENT represented Triassic theropod from Europe. The (1966). CUNY & GALTON (1993) referred it to the ge­ taxon was originally described as Halticosaurus lili­ nus Liliensternus WELLES and made it the holotype ensterniby HUENE (1934), based on the associated, of a new species. but disarticulated remains of two individuals from the Knollenmergel ofThuringen (HMN BM.R. 2175). Although the material is rather fragmentary, itcan Later, WELLES (1984) placed the species in the new be said with some certainty that it does represent a genus Liliensternus and designated the larger indi­ distinct and diagnosable species. It must be noted, vidual as the lectotype. It must be noted, however, though, that its referral to the genus Liliensternus is that the material may represent more than two indi­ questionable, since there are several major differ­ viduals, and it seems almost impossible to separate ences in the cervical vertebrae (e.g. the presence of the remains belonging to the larger and smaller indi­ two pairs of pleurocoels in L. airelensis versus only viduals (w. D. Heinrich, pers. comm., 1996); there­ one pair in L. liIiensterni; WELLES, 1984; CUNY & fore it is, at present, best to retain the whole material GALTON, 1993). However, the significance of these as the syntypes of the species. differences for distinguishing genera may only be decided if new, more complete material of L. airelen­ The syntype of Liliensternus WELLES is one of the sis is found . In all other characters, L. airelensis is largest known Triassic theropods, with an estimated very similar to L. liliensterni; therefore, it can be re­ length of over 5 m (PAUL, 1988). Despite this large ferred to the Coelophysoidea with some certainty. size, the individuals represented by the type mate­ rial were probably juvenile to subadult, since the Procompsognathus triassicus FRAAS, 1913 neurocentral sutures are still visible in the vertebrae, and only two fused sacrals are present (HUENE, Age: Norian, Upper Triassic. 1934 ). Occurrence: Middle Stubensandstein, Weil1er The anatomy of Liliensternus has briefly been Steinbruch, Pfaffenhofen, Baden-Wurttemberg, described by HUENE (1934), WELLES (1984) and Germany. ROWE & GAUTHIER (1990). The taxon shows several Comments: Procompsognathus triassicus was derived characters shared with the or named by Fraas on the basis of " ... the major part of more restricted ingroups of that , including a an extremely delicate dinosaur skeleton, including strqng latero-ventral expansion of the dorsal rim of the skull, the middle part of the body with the legs the acetabulum, a strongly downturned femoral and the anterior part of the tail" (FRAAS, 1913: 1099; head (HUENE, 1934; ROWE & GAUTHIER, 1990), and translated by O.R.). HUENE (1921a) later referred probably the presence of a subnarial gap (WELLES, another partial skull and a left manus (both bearing 1984). By contrast with the illustration of the pelvis in the collection number SMNS 12352) from the same ROWE & GAUTHIER (1990: f. 5.7), which only shows locality to this species and gave a detailed descrip­ an obturator-notch in the pubis, a completely en­ tion ofthe type material (SMNS 12591). In his review closed obturator foramen was present (HUENE, of P triassicus, OSTROM (1981) noted thatthe partial 1934: 159, taf. 15, f. 9b), opening dorso-medially as skull and manus cannot belong to this species, but in Syntarsus (RAATH, 1969). he otherwise accepted the association of the skull Within Ceratosauria, ROWE & GAUTHIER (1990) and postcranium SMNS 12591. Hefurthermore con­ placed Liliensternus as a sister taxon to the clade cluded that P triassicus represents a primitive thero­ comprising Syntarsus RAATH and pod within its own family, the Procompsognathidae. COPE, basing this relationship on the presence of a SERENO & WILD (1992) reviewed the type mate­ well developed horizontal ridge on the maxilla. This rial again, and argued that the skull and the postcra­ view is followed here. nial skeleton represent different animals. According to this paper, the skull SMNS 12591 belongs to the Liliensternus airelensis CUNY & GALTON, crocodylomorph Sa/toposuchus connectens 1993 HUENE, which is known from the same locality, while Age: -Hettangian, Upper Triassic - the postcranial material represents an early thero­ Lower Jurassic. pod. Just one later, CHATTERJEE (1993) in a short abstract, argued against the crocodylomorph Occurrences: Couches d'Airel, Normandie, nature of the skull and referred it to the Theropoda France. again.

79 OW.M. RAUHUT & A. HUNGERBUHLER

The question whether the skull and postcranial synapomorphy of the Dinosauriformes, rather than skeleton were found in association originally and be­ a ceratosaurian one, the exact shape of this struc­ long to the same individual, is very difficult to an­ ture in P triassicus might be taken as an indication of swer. While HUENE (1921a: 360) and BERCKHEMER a ceratosaurian relationship for this taxon. A further (1938: 194) had no doubts about the association of argumemt for placing Procompsognathus in the the skull and postcranial skeleton , Fraas already Ceratosauria, and maybe even in the Coelophysoi­ noted that the holotype specimen " ... is the major dea, might be the presence of elongate dorsal verte­ part of a ... dinosaur skeleton in three pieces, ... " brae and more or less triangular dorsal transverse (FRAAS, 1914: 129; translation byO.R.). Since these processes. Forthe present, however, more material specimens were purchased from the local quarry is needed to confirm the systematic position of this manager, together with several other vertebrate fos­ species within Theropoda. sils (SERENO & WILD, 1992), no data on their original Although it is difficult to give a formal diagnosis association in the quarry exists, and they might well for P triassicus, it is provisionally regarded as a valid be from different parts of the quarry (R . Wild, pers. taxon here. Possible characters to distinguish itfrom comm., 1996). The skull has been reprepared, and a other Triassic theropods include an elongate detailed description, currently being carried out by hindlimb (ratios tibia/femur: c. 1.2; Mt III/femur: c. S. Chatterjee (pers. comm., 1996), will certainly re­ 0.74). veal new information on its anatomy and systematic position. Pending this new information, this review Pterospondy/us trie/bae JAEKEL, 1913 will only focus on the postcranial material. Age: ? Norian, Upper Triassic. Sereno & Wild (1992) listed six theropod synapo­ morphies in the postcranium of Procompsognathus Occurrence: Knollenmergel, Baerecke quarry, FRAAS, so that there can be little doubt as to the Halberstadt, Sachsen-Anhalt, Germany. theropod nature of this specimen. Its systematic po­ Comments: The species is based on an isolated sition within Theropoda, however, is much more dif­ dorsal vertebra, found within a shell of the Triassic ficult to establish. OSTROM (1981) and SERENO & turtle Proganoche/ys BAUR (JAEKEL 1913). The cen­ WILD (1992) noted that the pubis was similar to Coe­ trum is low and elongate (Fig. 4), resembling the /ophysis and especially CAMP in show­ condition seen in Procompsognathus. On this basis, ing a " ... slightly bowed, rectangular pubic apron ... " HUENE (1921b, 1932) referred this species to the (OSTROM, 1981: 193)and in being " ... anteroposteri­ Procompsognathidae and noted that it might be con­ orly compressed .. ." and lacking a pubic boot (SER­ generic with Procompsognathus. However, the ENO & WILD, 1992: 437). However, all of these transverse processes are both triangular and features are also present in prosauropods (e.g. strongly backturned, a condition also seen in Syntar­ _ HUENE, 1926) and do therefore probably represent sus (RAATH, 1969), and noted as a synapomorphy of the plesiomorphic character state for theropods. the Ceratosauria by ROWE & GAUTHIER (1990). Un­ Thus, the development of the pubis only implies that fortunately, not much can be said about the trans­ P triassicus is a primitive theropod. verse processes in the anterior dorsal vertebrae of SERENO & WILD also note the presence of a " ... Procompsognathus, except that they also seem to sigmoid trochanteric shelf ... identical to that in cera­ be more or less triangular (see above). tosaurs such as Syntarsus and Coe/ophysis." (SER­ The transverse processes in Di/ophosaurus ENO & WILD, 1992: 437). Although NOVAS (1996) WELLES and Lilienstemus are less strongly back­ noted that the presence of a trochanteric shelf is a turned and not as significantly triangular, as is the

A 1 em c

Fig . 4 - pterospondy/us trie/bae JAEKEL; isolated dorsal vertebra, holotype; Knollenmergel, Halberstadt, Germany. A - Left lateral view. B - Dorsal view. C - Posterior view. Modified from HUENE (1921b). Scale bar indicates 1 em.

80 A REVIEW OF THE EUROPEAN TRIASSIC THEROPODS

case in Syntarsus or P. trielbae, indicating that this 1993), but the sacral ribs of this species are much character may rather represent a synapomorphy of more massive than seems to be the case in the type a more restricted ingroup ofthe Ceratosauria. Thus, specimen of Sa/topus HUENE. Characters like elon­ the vertebra from Halberstadt probably represents a gated hind limbs, bipedality, mainly three digits used member ofthe Coelophysidae. P. trielbae cannot be in locomotion, and habits are also present formally diagnosed and must be treated as a nomen in primitive dinosauriformes like Marasuchus SER­ dubium . ENO & ARCUCCI, 1994. Therefore, S. e/ginensis can only be treated as a probable dinosauriform nomen elginensis HUENE, 1910 dubium. Age: Upper , Upper Triassic. ? Syntarsus sp. Occurrence: Lossiemouth Sandstone Forma­ Age: ? Norian, Upper Triassic. tion, Grampian, Scotland. Occurrence: Pant-y-ffynnon fissure filling, Comments: This taxon is based on a partial South Glamorgan, Wales. postcranial skeleton of a small, long-limbed tetrapod from the Late Carnian of Elgin, Scotland (HUENE, Comments: A few theropod remains were found 1910; BENTON & WALKER, 1985). The specimen in the fissure fillings of southern Wales (BMNH R 3915) comprises the major parts of a dor­ and south-east England. The most significant of sal, sacral and caudal vertebral column, fragments these is an articulated left pelvic girdle (Fig. 5A), in­ of the forelimb and the hindlimbs. Skull remains are cluding parts of the sacrum, the posteriormost dor­ not present (contra NORMAN, 1990). The material is sal vertebrae and an associated left femur (Fig. 5B­ extremely poorly preserved. The bones are either in­ C), lacking the distal end (BMNH PV RU P 77/1 and dicated as imprints (counterslab) or at least superfi­ RU P 76/1) from the locality Pant-y-ffynnon in south­ cially remineralized as goethite (see BENTON & ern Wales (WARRENER, 1983). WALKER, 1985); original bone material is preserved The dorsal vertebrae are long and rather low, only occasionally (pers. obs.). Not much can there­ with low, but long neural spines. The sacral verte­ fore be said about the anatomy of the animal in de­ brae are fused, with the sutures between single ver­ tail. tebrae being only visible as slight swellings. The The skeleton, as preserved on the main slab, lies pubis and ischium are fused to the ilium, but not to on its belly, as judged by the impressions of the neu­ each other. The completely preserved ilium of the ral spines in the counterslab and the orientation of specimen is only 52 mm long. It is dolichoiliacic, the the hindlimbs. From both, the specimen on the main preacetabular part being significantly shorter than slab and the impressions on the counterslab, the the postacetabularone, and shows a well developed number of sacral vertebrae is probably two, rather brevis shelf and a strong lateral expansion of the than four, as argued by HUENE (1910), or three, as dorsal rim of the acetabulum. The distal end of the noted by NORMAN (1990). In contrast with the illus­ pubis is missing. The preserved part shows a slightly trations by HUENE (1910: taf. 1), the ilium is rather bowed shaft, a small, dorso-medially opening obtu­ short, especially its preacetabular part. Again, the rator foramen and a much larger pubic foramen un­ impressions ofthis bone in the counterslab show the derneath it. In anterior view, the fused pubes form a shortness of the ilium much better than the reminer­ rather broad apron. The ischium also lacks its distal alized bone remains on the main slab. The limbs are end. The obturator process is not offset from the pu­ very long and slender, the lower elements being es­ bic peduncle and distally there is a small notch be­ pecially elongated (ratio tibia/femur: c. 1.4). The fe­ tween the process and the ischial shaft. The femur mur is slightly curved. In the lower limbs, the tibia shows a downturned femoral head and a well devel­ and fibula seem to be of subequal width, and the oped trochanteric shelf. The fourth trochanter is rep­ metatarsals are long and slender. As judged by the resented by a long, low, proximally placed flange. impressions of the toes of the left hindlimb in the In all of their characters, the specimens are ex­ main slab, the number of digits used for locomotion tremely similar to the coelophysid Syntarsus was probably three, but there is an impression of a (RAATH, 1969; ROWE & GAUTHIER, 1990); thus, they further, significantly shorter digit medial to these. are tentatively referred to this genus, despite their In conclusion, the material represents a small, significantly older age (see also WARRENER, 1983). cursorial bipedal animal with elongated hindlimbs. A very similar theropod appears also to be present in However, the shortness of the ilium, the low number the Upper Triassic (Norian) of the Ghost Ranch of sacral vertebrae and the subequal width of the quarry of North America (PAUL, 1993). However, lower limb bones make it seem rather unlikely that it there is some confusion about the theropod speci­ represents a theropod. A short ilium and on ly two mens from this locality, and the theropod described sacrals are found in REIG (NOVAS, as Coelophysis by COLBERT (1989) is clearly distinct

81 O. w.M. RAUHUT & A. HUNGERBOHLER

It ts

;..

A B c

Fig. 5 - ?Syntarsus sp.; fissure fillings , Pant-y-ffynnon, Wales. A - BMNH PV RU P 77/1; left side of the pel vis; lateral view; pf - pubic foramen. B - BMNH RU P 76/1; left femur; medial view; ft - fourth trochanter, It -lesser trochanter. C - BMNH RU P 76/1 ; left femur; cranial view; ts - trochanteric shelf. Scale bars indicate 1 cm. from the Welsh specimens. In all the comparable gated prezygapophyses. It shows a rather broad characters (shape of the dorsal vertebral centra, ventral groove, similar to that found in the caudal width of pubes in anterior view, development of tro­ vertebrae of Liliensternus. However, the specimen chanteric shelf), the specimens are furthermore is generically and specifically indeterminable, so very similar to Proeompsognathus, and new discov­ that T posthumus must be regarded as a nomen du­ eries might prove that they are referable to this ge­ bium. nus. Velocipes guerichi HUENE, 1932 Tanystrophaeus posthumus HUENE, 1907-8 Age: Norian, Upper Triassic. Age: Norian, Upper Triassic. Occurrence: Lissauer Breccia, Gorny Slask, Occurrences: ? Middle Stubensandstein, Poland. Stuttgart-Heslach, Baden-WOrttemberg , Germany. Comments: The holotype of V. gueriehi is a Comments: T posthumus is based on a single proximal end of a long bone. HUENE (1932) de­ caudal vertebra (SMNS 4385) collected by S.F.J . scribed it as the upper half of a left theropod fibula, von Kapff in the sixties of the 19th century (A.H., un­ but the specimen is extremely poorly preserved and publ. data). The specimen has first been described even its identification as a fibula may be doubted. and figured by MEYER (1865: 114, pI. 27, fig . 4-6), but Therefore, V gueriehi can only be regarded as a no­ he did not assign it to a particular taxon. HUENE men dubium (see also NORMAN , 1990). (1907-8) first realized the theropod nature ofthe ver­ tebra and made it the type of a new species. The "Zanclodon" cambrensis NEWTON, 1899 specimen is now kept in the collections ofthe Staatli­ Age: Rhaetian, Upper Triassic. ches Museum fOr Naturkunde Stuttgart, where it is labelled as "Nierosaurus sp ... . Occurrence: Rhaetic beds, Mid-Glamorgan, Wales. The vertebra represents a relatively stout poste­ rior caudal with strongly elongated prezygapophy­ Comments: This species is based on a natural ses (Fig. 6). No transverse processes are present, mould in a sandstone slab from the Uppermost Tri­ and the neural spine is very low. assic of southern Wales. It was first described by NEWTON (1899) as Zane/odon cambrensis and later This element is readily identified as a theropod referred to the genus Mega/osaurus BUCKLAND. caudal vertebra, because of the presence of elon- MOLNAR, KURZANOV & DONG (1990) noted that this

82 A REVIEW OF THE EUROPEAN TRIASSIC THEROPODS

A B

Fig. 6 - Tanys/rophaeus pas/humus HUENE; SMNS 4385; isolated caudal vertebra, holotype; Stubensandstein, Stuttgart-Heslach, Germany. A - Right lateral view. B - Dorsal view. Scale bars indicate 1 cm.

dentary agrees with in six of the nine below follows GEYER & GW INNER (1991) and characters, listed by MADSEN (1976) to distinguish AIGNER & BACHMANN (1992), and we refer to these between and Megalosaurus; the other publications for further readings. three cannot be determined. Three of the characters The Lower Keuper or Lettenkeuper starts with were regarded as shared derived features: the an­ the Grenz-bonebed, which is unconformably overly­ gular rostral margin, separate interdental plates and ing the marine Muschelkalk. The Lower Keuper is exposed replacement teeth (MOLNAR, KURZANOV & mainly a succession of variegated shales interca­ DONG, 1990). However, an angular rostral margin is lated with thin dolomitic beds, probably laid down in also found in Lilienstemus lifienstemi (HUENE, a brackish-water environment. Several fluvial sand­ 1934), Syntarsus kayentakate ROWE, 1989, and stone beds occur regionally in different stratigraphic even the prosauropod Sellosaurus gracilis HUENE positions within the succession. A gradual shift to­ (GPIT PV 18318a; pers. obs.). The interdental wards a marine depositional environment results in plates are separate in engelhardti the Gipskeuperdeposits (stacked dolomite-gypsum MEYER (HMN MB.R. 1937; pers. obs.), Dilophosau­ sequences, topped by deposits of a playa-like envi­ rus wetherilli (WELLES) (WELLES, 1984), ronment). The Schilfsandstein is again probably a dongi CURRIE & ZHAO, 1993, and fluvial rather than a deltaic sandstone as presumed 10ngipeSWAGNER (OSTROM, 1978), amongst many previously, which has yielded a variety of temno­ others. The third character was described " ... re­ spondyl amphibians (BENTON, 1994a). Coloured placementteeth exposed at base between interden­ mudstones deposited in an arid desert-like environ­ tal plates ... " by MADSEN (1976: 10). This character is ment make up the Rote Wand and Kieselsandstein. clearly correlated with the separate interdental The latter unit includes a series of dolomitic beds, plates. Thus, all ofthese characters probably repre­ the Lehrbergschichten, and, interdigitating with the sent the plesiomorphic conditions within theropods shales, a fluvial sandstone (Kieselsandstein). and cannot therefore be used to link "Z." cambrensis with Mega/osaurus. Although all determinable char­ Similarly, the following Stubensandstein can be acters of the specimen agree quite well with Lilien­ subdivided into fluvially dominated deposits (Sand­ stern us liliensterni or Dilophosaurus wetherilli (e.g. steinkeuper) protruding from the margins into the the presence of very low and broad interdental German Basin towards the north and northwest, plates), its systematic position must remain un­ where they interdigitate with variegated playa pe­ solved. lites and lacustrine dolomites (Steinmergelkeuper) in the depositional centre of the German Basin. The "Z." cambrensis does not show any derived char­ marginal deposits comprise four heterochronous acters that would allow a formal diagnosis of the spe­ large-scaled alluvial fans termed first to fourth Stu­ cies. It might represent a distinct taxon, but at bensandstein. Note that the next following unit, the present it can only be regarded as a nomen dubium. Knollenmergel, in parts evidently represent a het­ erochronous facies deposited in a playa environ­ DISCUSSION ment: in southern WOrttemberg, the lower beds of the Knollenmergel replace the third and fourth Stu­ STRATIGRAPHY OF THE GERMAN KEUPER AND bensandstein laid down contemporaneously in cen­ DISTRIBUTION OF THEROPOD REMAINS tral and eastern WOrttemberg (BRENNER & Asynthesis olthe Mid-European Keuper succes- VILLINGER, 1981). In northern and eastern Ger­ sion is compiled in Figure 2, largely based on the de­ many, the Knollenmergel is topped by thick fluvial posits in South Germany. The lithologic description sandstones ("Hauptsandstein") grading into flood-

83 o. w.M. RAUHUT & A. HUNGERBOHLER

plain deposits, which both are by convenience re­ lower Stubensandstein because of the co­ ferred to the Lower Rhaetian. These again are occurrence of Aetosaurustherein and in the datable overlain unconformably by marine sandstones and Calcare di Zorzino in North Italy. This would point to­ shales, which can be safely dated as Rhaetian by a wards a Middle to Late Norian Age forthe whole Stu­ fauna including the bivalve Rhaetavicula contorta bensan(Jstein. However, as outlined before, the (PORTLOCK) . In South Germany, the Rhaetian is rep­ referral o( to the lower Stubensandstein resented by much condensed and patchy deltaic de­ is debatable, and it could well occur in the middle posits ("Rhat-sandstone" and "Rhat-shales") Stubensandstein (e.g. BRENNER, 1973). Further­ following the marine Rhaetic transgression. more, LUCAS & HUNT (1993) proposed the Revuel­ tian biochron (Lower Norian) for the Southwestern About half of the taxa treated here are derived United States, characterised, among other verte­ from the South German Stubensandstein, and for brates, by the Paratypothorax LONG & BAL­ this reason it is necessary to review the stratigraphic LE W (see also HUNT & LUCAS , 1992). Para­ position of this unit in more detail. The specimens occurs in the lower Stubensandstein from Pfaffenhofen (Procompsognathus triassicus, (WILD, 1991), but also in the middle (A.H., unpubl. the theropod nomen dubium Halticosaurus longo­ data), arguing for an Early Norian age of both tarsus, and the archosaurian "Halticosaurus" orbito­ subunits. However, the utility of vertebrate bio­ angulatus) can be safely placed into the middle chrons for dating strata on widely separate conti­ Stubensandstein (STOLL, 1929; BRENNER, 1978). nents has yet to be tested. We consider the Much more difficult to establish is the lithostra­ Stubensandstein spanning mostofthe Norian as the tigraphic position of Tanystrophaeus posthumus more likely interpretation at the moment. Both hy­ (HESLACH) and Oolichosuchus cristatus (KAL TEN­ potheses may be tested when dates are established TAL), since there are numerous abandoned Stuben­ for specimens referable to the Mystrio­ sand stein quarries around Stuttgart and the exact suchus FRAAS from the marine Dachsteinkalk of the provenance of the specimens was never recorded. Austrian Alps (reported preliminarily by BUFFETAUT, The Heslach quarry, also the source of a number of 1994). In the German Basin, Mystriosuchus is en­ important phytosaur remains and the rauisuchian tirely restricted to the middle Stubensandstein. MEYER (KAPFF, 1859; MEYER, 1861; GALTON, 1985b), corresponds probably to the site Lilienstemus lilienstemi comes from the Knollen­ "Heslacher Wand", which is most likely middle Stu­ mergel of Thuringia and can therefore be dated as bensandstein (BRENNER, 1978). The quarry near Late Norian with some certainty (see above). This Kaltental which produced the famous association of conclusion is furthermore substantiated by the fact 22 specimens of the stagonolepidid Aetosaurus thatthe remains of L.lilienstemiwere found in asso­ FRAAS was placed in the lower Stubensandstein ciation with remains of Plateosaurus engelhardti (WILD, 1989), but it is by no means sure whether 00- (HUENE, 1934), which is well known from the Late lichosuchus comes from the same site. Additionally, Norian of southern Germany and Switzerland all the lithostratigraphic referrals are debated, and (SANDER, 1992). the only safe conclusion is that both specimens must Syntarsus sp. comes from Pant-y-ffynnon have been derived from lower or middle Stuben­ quarry, one of the numerous fissure fillings in Car· sandstein. boniferous limestone strata of Wales and Southwest Two alternative hypotheses have been advo­ England. These fillings have a complex depositional cated to calibrate the lithostratigraphic units of the history and dating is difficult (see FRASER, 1994), German Keuper Succession (discussed in BENTON, being mainly achieved by their contents of verte­ 1994a, 1994b). Both unanimously agree that the brates. Pant-y-ffynnon has yielded, among other Stubensandstein must be considered Norian in age, taxa, the crocodylomorph CRUSH but conflicting palynologic, magnetostratigraphic (which has been suggested as a synonym of the and palaeoclimatic evidence has led to different as­ middle Stubensandstein Sa/toposuchus) and the sessments of the time span of this unit within the gliding diapsid Kuehneosaurus ROBINSON . Verte­ stage. In the first interpretation, the Stubensand­ brates seem to exclude a post-Triassic age, but the stein comprises almost the whole Norian, except for locality can be considered Late Triassic at best, the Knollenmergel, which is commonly referred to ranging anywhere from Late Carnian to Rhaetian the uppermost part of the Norian. According to the (BENTON & SPENCER, 1995). alternative view, the Stubensandstein is restricted to The age of Pterospondylus trielbae is usually the upper part of the Norian only. Currently, there is given as Rhaetian (e.g. HUENE, 1932; NORMAN, not sufficient data available to correlate one of the 1990). The section in the Baerecke quarry, as sum­ subunits of the Stubensandstein with the alpine Tri­ marized by SANDER (1992), consisted of two suc­ assic strata convincingly and to clinch this question. cessions: a lower claystone layer, which is WILD (1989) assigned a Middle Norian age to the considered to represent the equivalent of the Knol-

84 A REV/EW OF THE EUROPEAN TR/ASS/C THEROPODS

TABLE I Summary of the age, occurence, and taxonomic status of theropod taxa from the Triassic of Europe.

NAME AGE ' OCCURENCE TAXONOMIC STATUS

Avipes dillstedtianus Ladinian Lettenkeuper, Thuringia, ? Archosauria, nomen HUENE,1932 Germany dubium Dolichosuchus cristatus Norian Stubensandstein, Baden- Theropoda, nomen HUENE, 1932 Wurttemberg, Germany dubium Ha/ticosaurus /ongotarsus Norian Stubensandstein, Baden Theropoda, nomen (HUENE, 1907-8) Wurttemberg, Germany dubium (at least partim) "Halticosaurus" orbitoangu/atus Norian Stubensandstein, Baden- ? , HUENE, 1932 Wurttemberg, Germany nomen dubium ? Liliensternus aire/ensis Rhaetian- Airel quarry, Normandy, Theropoda, valid species CUNY & GALTON 1993 Hettangian France Liliensternus /iliensterni Norian Knollenmergel, Thuringia, Theropoda, valid species (HUENE, 1934) Germany Proeompsognathus triassicus Norian Stubensandstein, Baden Theropoda, provisionally FRAAS, 1913 Wurttemberg, Germany regarded as valid species Plerospondy/us trie/bae ? Norian Knollenmergel, Thuringia, Theropoda, nomen JAEKEL, 1913 Germany dubium Sa/topus e/ginensis Carnian Lossimouth Sandstone Dinosauriformes, HUENE, 1910 Formation, Scotland nomen dubium Tanystrophaeus posthumus Norian Stubensandstein, Baden- Theropoda, nomen HUENE, 1907-8 Wurttemberg, Germany dubium Ve/oeipes gueriehi Norian Lissauer Breccia, Gorny Vertebrata, nomen dubium HUENE , 1932 Slask, Poland "Zane/odon" cambrensis Rhaetian Rhaetic sandstone, Wales Theropoda, nomen NEWTON, 1899 dubium

lenmergel, overlain by sand-and siltstonesofproba­ THE TRIASSIC THEROPOD FAUNA OF EUROPE bly Rhaetian age. According to JAEKEL (1913), at Only three theropod species from the Upper Tri­ least a part of the turtle remains were derived from assic of Europe can presently be regarded as being the lower part of the section of the Baerecke quarry. valid (see also TABLE I): Liliensternus liIiensterni, Since the holotype of P trie/bae was found within a ?Liliensternus airelensis and Procompsognathus turtle carapace, it might therefore be of Late Norian triassicus. Furthermore, a species of Syntarsus was age, rather than Rhaetian. most probably present in the Upper Triassic of "Zane/odon" eambrensis comes from a sand­ Europe as well. Both L. /iliensterni and P triassicus stone bed in southern Wales. The slab was derived come from the Norian of southern Germany, the from a stack of building material, and its exact stra­ specimen of Syntarsus derives from probably No­ tigraphic position (above or below the Rhaetavieu/a rian fissure fillings in Wales, while? L. aire/ensiswas eontorta beds) is uncertain (NEWTON , 1899). How­ found in sediments spanning the Triassic-Jurassic ever, the sandstone unit, where the specimen came boundary in northern France, and might even be of from is considered to be Rhaetian (WARRINGTON et lowermost Jurassic age. a/., 1980). Apart from the remains discussed above, only a The stratigraphic position of ?Liliensternus few other fragments can be assigned to the Thero­ aire/ensis has been discussed in some detail by poda with any certainty. GALTON (1985a) noted a CUNY & GALTON (1993). The specimen might be ei­ probable theropod femur fragment from the Stuben­ ther late Rhaetian or early Hettangian in age. sandstein of Pfaffenhofen, BUFFETAUT & WOUTERS (1986) described some probable theropod teeth

85 o. w.M. RAUHUT & A. HUNGERBOHLER

from the Norian of France, and SANDER (1992) re­ lack of tetanuran fossils from the Triassic either re­ ferred several teeth from the Knollenmergel of Frick flects geographic isolation of this group during that (Switzerland) and Trossingen (southern Germany) time, or their rareness in Upper Triassic vertebrate to cf. Lifiensternus. However, since teeth of thero­ faunas. pod dinosaurs do not differ significantly from the ser­ In this context, another aspect is worth mention­ rated teeth of other Triassic , the ing: there are many fossil vertebrate localities, and identification of isolated serrated teeth from Triassic dinosaurs are quite abundant in the European Up­ beds as theropod is always doubtful. For the same per Triassic (see e.g. WEISHAMPEL, 1990), butthero­ reason, taxa based on isolated teeth (e.g . "Mega/o­ pod remains are rather rare. Although this may saurus" c/oacinus QUENSTEDT, "Mega/osaurus" ob­ partly be due to the taphonomy of the localities tusus HENRY, "P/ateosaurus" ornatus HUENE; see (SANDER, 1992), it might reflect genuine rarity of HUENE, 1907-8, 1932; GALTON, 1985b) are not in­ theropods in the Upper Triassic vertebrate fauna of cluded in this review. Europe. In addition to the skeletal remains, supposed theropod footprints have been described from differ­ ACKNOWLEDGEMENTS ent localities (e. g. HADERER, 1990). However, there We thank Sandra Chapman, London, W.-D. are many sources of error in identifying prints (KING Heinrich, Berlin, and R. Wild, Stuttgart, for access to & BENTON, 1996), and, apart from theropods, other specimens under their care. G. Cuny, Bristol, is animals including protodinosaurs or early ornithis­ thanked for providing casts of L. airelensis and for chians might have produced tridactyl prints, hence translating the abstract into French. Special thanks footprint evidence is not taken into consideration are due to Dave Unwin and Mike Benton, both Bris­ here. tol, for comments on the manuscript. Thanks are Thus, the earliest certain records of theropod di­ also due to Ben Edwards, London, for bringing the nosaurs in Europe come from the upper parts of the fissure-filling specimens to our attention. Simone Lower Norian. Since there are vertebrate localities Klutzny, Bristol , is thanked for discussions while this in the pre-Norian Upper Triassic in Europe, and es­ paper was written. We thank the referees, Thomas pecially in Germany, the Upper Triassic is well repre­ Holtz, College Park, and Bernardino Perez-Moreno, sented by terrestrial sediments (see above), the Madrid, for useful comments. O.R. is supported by appearance of theropod dinosaurs at that time is in­ the EC under TMR grant ERBFMBICT 961013 and terpreted here as representing the first radiation of A.H. under ERBCHBICT 930521, which is gratefully theropods in Europe. acknowledged. Furthermore, all identifiable theropod remains ABBREVIATIONS appear to belong to the Coelophysoidea, within the Ceratosauria. The presence ofthe genus Syntarsus BMNH - Natural History Museum, London , UK; in particular is interesting; this taxon is well known GPIT - Institut und Museum fOr Geologie und from the lowermost Jurassic of southern Africa and Palaontologie TObingen, Germany; HMN - Museum North America (ROWE & GAUTHIER, 1990), and fOr Naturkunde, Humboldt Universitat, '3erlin, Ger·· might have been present in the Norian of North many; SMNS - Staatliches Museum fOr Naturkunde America as well (PAUL, 1993). Since the oldest Stuttgart, Germany. theropods are known from the Carnian of South America (SERENO & NOVAS, 1992), and coelophy­ REFERENCES soids first appear in the fossil record in the early No­ AIGNER, T. & BACHMANN, G.H. (1992) - Sequence-stratigraphic rian (ROWE & GAUTHIER, 1990), there would appear framework of the Germanic Triassic. Sediment. Geol., 80: to have been a rapid radiation of ceratosaurian 115-135. theropods in the late Carnian / early Norian. These BENTON , M.J. (1984) - Fossil of the German Late Triassic and the origin of the dinosaurs, in REIF, W.-E. & WESTPHAL, F. conclusions are in general accordance with ideas (Eds.), Third Symposium on Mesozoic Terrestrial Ecosys­ proposed by BENTON (1984, 1993a), that dinosaur te ms, TObingen, Short Papers, TObingen, pp. 13-18. radiation was rapid in the Upper Triassic, possibly BENTON, M.J. & WALKER, A.D. 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