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A REVIEW OF EUROPEAN TRIASSIC THEROPODS
Oliver W. M. RAUHUT
Axel HUNGERBÜHLER
A : 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 (H
), (?) Lilienster
-
nus airelensis C
&G , and Procompsognathus triassicus F . (?) L. airelensis
might represent a distinct genus, but more material is needed to confirm this. The genus
Syntarsus R
, 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 of their fossil remains, theropods were obviously rather rare elements of the Up-
per Triassic terrestrial vertebrate fauna of Europe.
R
: Le registre fossile des théropodes du Trias supérieur européen est révisé en terme
à la fois de validité des taxons proposés et de répartition stratigraphiques des restes fossi-
les. Seule trois espèces peuvent être actuellement considérées comme valides: Lilienster-
nus liliensterni (H
), (?) Liliensternus airelensis C &G et Procompsognathus
triassicus F
. (?) L. airelensis pourrait en fait appartenir à un nouveau genre, mais des
restes plus complets seraient nécessaires pour confirmer cette hypothèse. Le genre Syn-
tarsus R
, qui n'avait jusqu'à présent été décrit qu'en Afrique du Sud et en Amérique du
Nord, l'est également pour la première fois en Europe. Les théropodes apparaissent en Eu
-
rope à partir du Norien, et tous les fossiles identifiables appartiennent aux Coelophysoidea.
Si l'on en croit la pauvreté de leurs restes fossiles, il semble évident que les théropodes re
-
présentaient des éléments relativement rares au sein des faunes de vertébrés terrestres du
Trias supérieur d'Europe.
INTRODUCTION
In the light of new discoveries, especially in
South America, the early history of theropod dino
-
saurs has been of great interest recent years (e.g.
S
&N , 1992). Theropods are one of the
most diverse dinosaur groups in the Late Jurassic
and the Cretaceous, but little is still known about
their origin, radiation and early diversification. Espe
-
cially interesting in this respect is the first radiation of
theropod dinosaurs, or dinosaurs in general, in the
Upper Triassic (e.g. B
, 1982; B ,
1984, 1993). Our knowledge of Upper Triassic
theropods is mainly based on the American fossil
record, although theropod remains are known from
the Triassic all over the world (W
, 1990).
The aim of the present paper is to review Triassic
theropod records from Europe, in terms of the taxo
-
nomic validity of named species, and the stra
-
tigraphical distribution of theropod remains in the
European Triassic, to determine the first appear
-
ance and taxonomic diversity of theropod dinosaurs
in the Upper Triassic of Europe.
THEROPODS FROM THE TRIASSIC OF
EUROPE
In the following section, theropod taxa described
from Europe are reviewed. See Figure 1 for the geo
-
graphical occurrences, and Figure 2 for the stra
-
tigraphical distribution of theropodan remains.
75
artigos/papers
Avipes dillstedtianus HUENE, 1932
Age: Ladinian, Middle Triassic.
Occurrence: Grenzdolomit close to Dillstedt,
Thüringen, Germany.
Comments: A. dillstedtianus was described by
H
(1932) on the basis of the proximal ends of
three articulated metatarsals from the lowermost
Upper Triassic (Lettenkeuper) of Thuringia.
The proximal ends of the metatarsals are closely
appressed and deeper than wide (see H
, 1932:
taf. 1, f. 7). Distally, the shafts of the bones are sepa
-
rated from each other and appear to diverge.
As already mentioned by N
(1990), the
specimen probably represents a digitigrade animal,
but the metatarsals do not show any characters that
allow a referral to the Theropoda. Since there are a
variety of digitigrade animals in the Upper Triassic,
A. dillstedtianus can only be referred to as probable
Archosauria gen. et sp. indet. (see also N
,
1990).
Dolichosuchus cristatus HUENE, 1932
Age: Norian, Upper Triassic.
Occurrence: Lower or middle Stubensandstein,
Stuttgart-Kaltental, Baden-Württemberg, Germany.
Comments: Dolichosuchus cristatus is based
on an isolated tibia from the Stubensandstein of
southern Germany (H
, 1932). The specimen
(BMNH 38056) shows a large cnemial crest, and a
lateral ridge for the attachment of the fibula; these
characters indicate that it represents a theropod.
However, the poor preservation of the element
makes it generically and specifically indeterminate,
so that D. cristatus must be treated as a nomen du
-
bium. It should be noted, though, that, as already
76
O.W.M. RAUHUT & A. HUNGERBÜHLER
77
A REVIEW OF THE EUROPEAN TRIASSIC THEROPODS
mentioned by H (1934), the specimen shows
great similarities to the tibiae of the slightly younger
Liliensternus liliensterni (H
), and also of Dilo
-
phosaurus wetherilli (W
, 1984). This suggests
that it probably represents a member of the Coelo
-
physoidea ( = Dilophosaurus + Coelophysidae;
H
, 1994).
Halticosaurus longotarsus HUENE, 1907-8
Age: Norian, Upper Triassic.
Occurrence: Middle Stubensandstein, Weißer
Steinbruch, Pfaffenhofen, Baden-Württemberg,
Germany.
Comments: Halticosaurus longotarsus, the type
species of the genus Halticosaurus, was described
by H
(1907-8) on the basis of some fragments
from the Stubensandstein of Pfaffenhofen (all bear
-
ing the collection number SMNS 12353). The mate
-
rial originally comprised a fragmentary dentary,
parts of cervical, dorsal, sacral, and caudal verte
-
brae, fragments of a humerus, an ilium and two
femora, and a complete metatarsal. Only one cervi-
cal vertebra (H
, 1907-8: taf. 97, f. 4), a dorsal
vertebral centrum (op. cit.: taf. 97, f. 7), another frag-
mentary vertebral centrum (op. cit.: taf. 97, f. 6), the
two femoral fragments (op. cit.: taf. 97, f. 1, 2) and
the metatarsus (op. cit.: taf. 97, f. 9) could be located
in the SMNS. All of the material is very badly pre-
served, and most of the fragments are not identifi
-
able even as theropods. Furthermore, the original
association of these specimens is very dubious. Ac
-
cording to H
(1907-8), the remains were found
"... together with Sellosaurus Fraasi and Teratosau
-
rus (?) minor [both synonyms of Sellosaurus gracilis;
note by the authors] ... in a marly layer intercalated in
the Stubensandstein ..." (H
, 1907-8: 231;
translated by O.R.), thus, it seems quite possible,
that some of the material might represent a pro
-
sauropod. Indeed, the collection number also in
-
cludes a skull of Sellosaurus H
(SMNS
12353a). The only remains that can be referred to
the Theropoda with some certainty are the femoral
fragments. One of them shows a spike-like lesser
trochanter and a significantly downturned femoral
head (Fig. 3), both characters also found in coelo
-
physoids. Although H. longotarsus must be treated
as a nomen dubium, the type series might therefore
include remains of a coelophysoid theropod.
H
(1921b) later referred some fragmentary
remains from the Norian of Halberstadt to the same
genus as cf. Halticosaurus longotarsus. This mate-
rial (now in the HMN) is extremely fragmentary as
well, and none of it can even be shown to be theropo-
dan with any certainty.
cf. Halticosaurus orbitoangulatus
H
UENE, 1932
Age: Norian, Upper Triassic.
Occurrence: Middle Stubensandstein, Weißer
Steinbruch, Pfaffenhofen, Baden-Württemberg,
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 end and most
of the skull roof is missing. The specimen has been
described in some detail by H
(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 H
, 1921a; S &
W
, 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
crocodile, rather than a theropod.
Liliensternus liliensterni (HUENE, 1934)
Age: Norian, Upper Triassic.
78
O.W.M. RAUHUT & A. HUNGERBÜHLER
Occurrences: Knollenmergel, Thüringen;
?Trossingen, Württemberg, Germany; ? Frick, Swit
-
zerland.
Comments: Liliensternus liliensterni is the best
represented Triassic theropod from Europe. The
taxon was originally described as Halticosaurus lili
-
ensterni by H
(1934), based on the associated,
but disarticulated remains of two individuals from
the Knollenmergel of Thüringen (HMN BM.R. 2175).
Later, W
(1984) placed the species in the new
genus Liliensternus and designated the larger indi
-
vidual as the lectotype. It must be noted, however,
that the material may represent more than two indi
-
viduals, and it seems almost impossible to separate
the remains belonging to the larger and smaller indi
-
viduals (W. D. Heinrich, pers. comm., 1996); there
-
fore it is, at present, best to retain the whole material
as the syntypes of the species.
The syntype of Liliensternus W
is one of the
largest known Triassic theropods, with an estimated
length of over5m(P
, 1988). Despite this large
size, the individuals represented by the type mate-
rial were probably juvenile to subadult, since the
neurocentral sutures are still visible in the vertebrae,
and only two fused sacrals are present (H
,
1934).
The anatomy of Liliensternus has briefly been
described by H
(1934), W (1984) and
R
&G (1990). The taxon shows several
derived characters shared with the Ceratosauria or
more restricted ingroups of that clade, including a
strong latero-ventral expansion of the dorsal rim of
the acetabulum, a strongly downturned femoral
head (H
, 1934; R &G , 1990), and
probably the presence of a subnarial gap (W
,
1984). By contrast with the illustration of the pelvis in
R
&G (1990: f. 5.7), which only shows
an obturator-notch in the pubis, a completely en
-
closed obturator foramen was present (H
,
1934: 159, taf. 15, f. 9b), opening dorso-medially as
in Syntarsus (R
, 1969).
Within Ceratosauria, R
&G (1990)
placed Liliensternus as a sister taxon to the clade
comprising Syntarsus R
and Coelophysis
C
, basing this relationship on the presence of a
well developed horizontal ridge on the maxilla. This
view is followed here.
Liliensternus airelensis CUNY &GALTON,
1993
Age: Rhaetian-Hettangian, Upper Triassic -
Lower Jurassic.
Occurrences: Couches d'Airel, Normandie,
France.
Comments: The holotype of this species com
-
prises a fragmentary associated vertebral column
and parts of the pelvis (Caen University, unnum
-
bered). The material was originally described as
Halticosaurus sp. by L
&L
(1966). C &G (1993) referred it to the ge
-
nus Liliensternus W
and made it the holotype
of a new species.
Although the material is rather fragmentary, it can
be said with some certainty that it does represent a
distinct and diagnosable species. It must be noted,
though, that its referral to the genus Liliensternus is
questionable, since there are several major differ
-
ences in the cervical vertebrae (e.g. the presence of
two pairs of pleurocoels in L. airelensis versus only
one pair in L. liliensterni;W
, 1984; C &
G
, 1993). However, the significance of these
differences for distinguishing genera may only be
decided if new, more complete material of L. airelen
-
sis is found. In all other characters, L. airelensis is
very similar to L. liliensterni; therefore, it can be re
-
ferred to the Coelophysoidea with some certainty.
Procompsognathus triassicus FRAAS, 1913
Age: Norian, Upper Triassic.
Occurrence: Middle Stubensandstein, Weißer
Steinbruch, Pfaffenhofen, Baden-Württemberg,
Germany.
Comments: Procompsognathus triassicus was
named by Fraas on the basis of "... the major part of
an extremely delicate dinosaur skeleton, including
the skull, the middle part of the body with the legs
and the anterior part of the tail" (F
, 1913: 1099;
translated by O.R.). H
(1921a) later referred
another partial skull and a left manus (both bearing
the collection number SMNS 12352) from the same
locality to this species and gave a detailed descrip
-
tion of the type material (SMNS 12591). In his review
of P. triassicus,O
(1981) noted that the partial
skull and manus cannot belong to this species, but
he otherwise accepted the association of the skull
and postcranium SMNS 12591. He furthermore con
-
cluded that P. triassicus represents a primitive thero
-
pod within its own family, the Procompsognathidae.
S
&W (1992) reviewed the type mate
-
rial again, and argued that the skull and the postcra
-
nial skeleton represent different animals. According
to this paper, the skull SMNS 12591 belongs to the
crocodylomorph Saltoposuchus connectens
H
, which is known from the same locality, while
the postcranial material represents an early thero
-
pod. Just one year later, C
(1993) in a
short abstract, argued against the crocodylomorph
nature of the skull and referred it to the Theropoda
again.
79
A REVIEW OF THE EUROPEAN TRIASSIC THEROPODS
The question whether the skull and postcranial
skeleton were found in association originally and be
-
long to the same individual, is very difficult to an
-
swer. While H
(1921a: 360) and B
(1938: 194) had no doubts about the association of
the skull and postcranial skeleton, Fraas already
noted that the holotype specimen "... is the major
part of a ... dinosaur skeleton in three pieces, ..."
(F
, 1914: 129; translation by O.R.). Since these
specimens were purchased from the local quarry
manager, together with several other vertebrate fos
-
sils (S
&W , 1992), no data on their original
association in the quarry exists, and they might well
be from different parts of the quarry (R. Wild, pers.
comm., 1996). The skull has been reprepared, and a
detailed description, currently being carried out by
S. Chatterjee (pers. comm., 1996), will certainly re
-
veal new information on its anatomy and systematic
position. Pending this new information, this review
will only focus on the postcranial material.
Sereno & Wild (1992) listed six theropod synapo
-
morphies in the postcranium of Procompsognathus
F
, so that there can be little doubt as to the
theropod nature of this specimen. Its systematic po-
sition within Theropoda, however, is much more dif-
ficult to establish. O
(1981) and S &
W
(1992) noted that the pubis was similar to Coe-
lophysis and especially Segisaurus C
in show-
ing a "... slightly bowed, rectangular pubic apron ..."
(O
, 1981: 193) and in being "... anteroposteri-
orly compressed ..." and lacking a pubic boot (S
&W , 1992: 437). However, all of these
features are also present in prosauropods (e.g.
H
, 1926) and do therefore probably represent
the plesiomorphic character state for theropods.
Thus, the development of the pubis only implies that
P. triassicus is a primitive theropod.
S
&W also note the presence of a "...
sigmoid trochanteric shelf ... identical to that in cera
-
tosaurs such as Syntarsus and Coelophysis." (S
&W , 1992: 437). Although N (1996)
noted that the presence of a trochanteric shelf is a
synapomorphy of the Dinosauriformes, rather than
a ceratosaurian one, the exact shape of this struc
-
ture in P. triassicus might be taken as an indication of
a ceratosaurian relationship for this taxon. A further
argument for placing Procompsognathus in the
Ceratosauria, and maybe even in the Coelophysoi
-
dea, might be the presence of elongate dorsal verte
-
brae and more or less triangular dorsal transverse
processes. For the present, however, more material
is needed to confirm the systematic position of this
species within Theropoda.
Although it is difficult to give a formal diagnosis
for P. triassicus, it is provisionally regarded as a valid
taxon here. Possible characters to distinguish it from
other Triassic theropods include an elongate
hindlimb (ratios tibia/femur: c. 1.2; Mt III/femur: c.
0.74).
Pterospondylus trielbae JAEKEL, 1913
Age: ? Norian, Upper Triassic.
Occurrence: Knollenmergel, Baerecke quarry,
Halberstadt, Sachsen-Anhalt, Germany.
Comments: The species is based on an isolated
dorsal vertebra, found within a shell of the Triassic
turtle Proganochelys B
(J 1913). The cen-
trum is low and elongate (Fig. 4), resembling the
condition seen in Procompsognathus. On this basis,
H
(1921b, 1932) referred this species to the
Procompsognathidae and noted that it might be con-
generic with Procompsognathus. However, the
transverse processes are both triangular and
strongly backturned, a condition also seen in Syntar
-
sus (R
, 1969), and noted as a synapomorphy of
the Ceratosauria by R
&G (1990). Un
-
fortunately, not much can be said about the trans
-
verse processes in the anterior dorsal vertebrae of
Procompsognathus, except that they also seem to
be more or less triangular (see above).
The transverse processes in Dilophosaurus
W
and Liliensternus are less strongly back
-
turned and not as significantly triangular, as is the
80
O.W.M. RAUHUT & A. HUNGERBÜHLER
A B C
case in Syntarsus or P. trielbae, indicating that this
character may rather represent a synapomorphy of
a more restricted ingroup of the Ceratosauria. Thus,
the vertebra from Halberstadt probably represents a
member of the Coelophysidae. P. trielbae cannot be
formally diagnosed and must be treated as a nomen
dubium.
Saltopus elginensis HUENE, 1910
Age: Upper Carnian, Upper Triassic.
Occurrence: Lossiemouth Sandstone Forma
-
tion, Grampian, Scotland.
Comments: This taxon is based on a partial
postcranial skeleton of a small, long-limbed tetrapod
from the Late Carnian of Elgin, Scotland (H
,
1910; B
&W , 1985). The specimen
(BMNH R 3915) comprises the major parts of a dor
-
sal, sacral and caudal vertebral column, fragments
of the forelimb and the hindlimbs. Skull remains are
not present (contra N
, 1990). The material is
extremely poorly preserved. The bones are either in
-
dicated as imprints (counterslab) or at least superfi-
cially remineralized as goethite (see B
&
W
, 1985); original bone material is preserved
only occasionally (pers. obs.). Not much can there-
fore be said about the anatomy of the animal in de-
tail.
The skeleton, as preserved on the main slab, lies
on its belly, as judged by the impressions of the neu-
ral spines in the counterslab and the orientation of
the hindlimbs. From both, the specimen on the main
slab and the impressions on the counterslab, the
number of sacral vertebrae is probably two, rather
than four, as argued by H
(1910), or three, as
noted by N
(1990). In contrast with the illus
-
trations by H
(1910: taf. 1), the ilium is rather
short, especially its preacetabular part. Again, the
impressions of this bone in the counterslab show the
shortness of the ilium much better than the reminer
-
alized bone remains on the main slab. The limbs are
very long and slender, the lower elements being es
-
pecially elongated (ratio tibia/femur: c. 1.4). The fe
-
mur is slightly curved. In the lower limbs, the tibia
and fibula seem to be of subequal width, and the
metatarsals are long and slender. As judged by the
impressions of the toes of the left hindlimb in the
main slab, the number of digits used for locomotion
was probably three, but there is an impression of a
further, significantly shorter digit medial to these.
In conclusion, the material represents a small,
cursorial bipedal animal with elongated hindlimbs.
However, the shortness of the ilium, the low number
of sacral vertebrae and the subequal width of the
lower limb bones make it seem rather unlikely that it
represents a theropod. A short ilium and only two
sacrals are found in Herrerasaurus R
(N ,
1993), but the sacral ribs of this species are much
more massive than seems to be the case in the type
specimen of Saltopus H
. Characters like elon
-
gated hindlimbs, bipedality, mainly three digits used
in locomotion, and cursorial habits are also present
in primitive dinosauriformes like Marasuchus S
&A , 1994. Therefore, S. elginensis can
only be treated as a probable dinosauriform nomen
dubium.
? Syntarsus sp.
Age: ? Norian, Upper Triassic.
Occurrence: Pant-y-ffynnon fissure filling,
South Glamorgan, Wales.
Comments: A few theropod remains were found
in the Late Triassic fissure fillings of southern Wales
and south-east England. The most significant of
these is an articulated left pelvic girdle (Fig. 5A), in
-
cluding parts of the sacrum, the posteriormost dor
-
sal vertebrae and an associated left femur (Fig. 5B-
C), lacking the distal end (BMNH PV RU P 77/1 and
RU P 76/1) from the locality Pant-y-ffynnon in south-
ern Wales (W
, 1983).
The dorsal vertebrae are long and rather low,
with low, but long neural spines. The sacral verte-
brae are fused, with the sutures between single ver-
tebrae being only visible as slight swellings. The
pubis and ischium are fused to the ilium, but not to
each other. The completely preserved ilium of the
specimen is only 52 mm long. It is dolichoiliacic, the
preacetabular part being significantly shorter than
the postacetabular one, and shows a well developed
brevis shelf and a strong lateral expansion of the
dorsal rim of the acetabulum. The distal end of the
pubis is missing. The preserved part shows a slightly
bowed shaft, a small, dorso-medially opening obtu
-
rator foramen and a much larger pubic foramen un
-
derneath it. In anterior view, the fused pubes form a
rather broad apron. The ischium also lacks its distal
end. The obturator process is not offset from the pu
-
bic peduncle and distally there is a small notch be
-
tween the process and the ischial shaft. The femur
shows a downturned femoral head and a well devel
-
oped trochanteric shelf. The fourth trochanter is rep
-
resented by a long, low, proximally placed flange.
In all of their characters, the specimens are ex
-
tremely similar to the coelophysid Syntarsus
(R
, 1969; R &G , 1990); thus, they
are tentatively referred to this genus, despite their
significantly older age (see also W
, 1983).
A very similar theropod appears also to be present in
the Upper Triassic (Norian) of the Ghost Ranch
quarry of North America (P
, 1993). However,
there is some confusion about the theropod speci
-
mens from this locality, and the theropod described
as Coelophysis by C
(1989) is clearly distinct
81
A REVIEW OF THE EUROPEAN TRIASSIC THEROPODS
from the Welsh specimens. In all the comparable
characters (shape of the dorsal vertebral centra,
width of pubes in anterior view, development of tro-
chanteric shelf), the specimens are furthermore
very similar to Procompsognathus, and new discov-
eries might prove that they are referable to this ge-
nus.
Tanystrophaeus posthumus HUENE, 1907-8
Age: Norian, Upper Triassic.
Occurrences: ? Middle Stubensandstein,
Stuttgart-Heslach, Baden-Württemberg, Germany.
Comments: T. posthumus is based on a single
caudal vertebra (SMNS 4385) collected by S.F.J.
von Kapff in the sixties of the 19th century (A.H., un
-
publ. data). The specimen has first been described
and figured by M
(1865: 114, pl. 27, fig. 4-6), but
he did not assign it to a particular taxon. H
(1907-8) first realized the theropod nature of the ver
-
tebra and made it the type of a new species. The
specimen is now kept in the collections of the Staatli
-
ches Museum für Naturkunde Stuttgart, where it is
labelled as "Nicrosaurus sp.".
The vertebra represents a relatively stout poste
-
rior caudal with strongly elongated prezygapophy
-
ses (Fig. 6). No transverse processes are present,
and the neural spine is very low.
This element is readily identified as a theropod
caudal vertebra, because of the presence of elon
-
gated prezygapophyses. It shows a rather broad
ventral groove, similar to that found in the caudal
vertebrae of Liliensternus. However, the specimen
is generically and specifically indeterminable, so
that T. posthumus must be regarded as a nomen du-
bium.
Velocipes guerichi HUENE, 1932
Age: Norian, Upper Triassic.
Occurrence: Lissauer Breccia, Gorny Slask,
Poland.
Comments: The holotype of V. guerichi is a
proximal end of a long bone. H
(1932) de
-
scribed it as the upper half of a left theropod fibula,
but the specimen is extremely poorly preserved and
even its identification as a fibula may be doubted.
Therefore, V. guerichi can only be regarded as a no
-
men dubium (see also N
, 1990).
"Zanclodon" cambrensis NEWTON, 1899
Age: Rhaetian, Upper Triassic.
Occurrence: Rhaetic beds, Mid-Glamorgan,
Wales.
Comments: This species is based on a natural
mould in a sandstone slab from the Uppermost Tri
-
assic of southern Wales. It was first described by
N
(1899) as Zanclodon cambrensis and later
referred to the genus Megalosaurus B
.
M
,K &D (1990) noted that this
82
O.W.M. RAUHUT & A. HUNGERBÜHLER
A
B
C
dentary agrees with Megalosaurus in six of the nine
characters, listed by M
(1976) to distinguish
between Allosaurus and Megalosaurus; the other
three cannot be determined. Three of the characters
were regarded as shared derived features: the an
-
gular rostral margin, separate interdental plates and
exposed replacement teeth (M
,K &
D
, 1990). However, an angular rostral margin is
also found in Liliensternus liliensterni (H
,
1934), Syntarsus kayentakate R
, 1989, and
even the prosauropod Sellosaurus gracilis H
(GPIT PV 18318a; pers. obs.). The interdental
plates are separate in Plateosaurus engelhardti
M
(HMN MB.R. 1937; pers. obs.), Dilophosau-
rus wetherilli (W
)(W , 1984), Sinraptor
dongi C
&Z , 1993, and Compsognathus
longipes W
(O , 1978), amongst many
others. The third character was described "... re-
placement teeth exposed at base between interden
-
tal plates ..." by M
(1976: 10). This character is
clearly correlated with the separate interdental
plates. Thus, all of these characters probably repre
-
sent the plesiomorphic conditions within theropods
and cannot therefore be used to link "Z." cambrensis
with Megalosaurus. Although all determinable char
-
acters of the specimen agree quite well with Lilien
-
sternus liliensterni or Dilophosaurus wetherilli (e.g.
the presence of very low and broad interdental
plates), its systematic position must remain un
-
solved.
"Z." cambrensis does not show any derived char
-
acters that would allow a formal diagnosis of the spe
-
cies. It might represent a distinct taxon, but at
present it can only be regarded as a nomen dubium.
DISCUSSION
S
G K
D T R
A synthesis of the Mid-European Keuper succes
-
sion is compiled in Figure 2, largely based on the de
-
posits in South Germany. The lithologic description
below follows G
&G (1991) and
A
&B (1992), and we refer to these
publications for further readings.
The Lower Keuper or Lettenkeuper starts with
the Grenz-bonebed, which is unconformably overly
-
ing the marine Muschelkalk. The Lower Keuper is
mainly a succession of variegated shales interca
-
lated with thin dolomitic beds, probably laid down in
a brackish-water environment. Several fluvial sand-
stone beds occur regionally in different stratigraphic
positions within the succession. A gradual shift to-
wards a marine depositional environment results in
the Gipskeuper deposits (stacked dolomite-gypsum
sequences, topped by deposits of a playa-like envi-
ronment). The Schilfsandstein is again probably a
fluvial rather than a deltaic sandstone as presumed
previously, which has yielded a variety of temno-
spondyl amphibians (B
, 1994a). Coloured
mudstones deposited in an arid desert-like environ
-
ment make up the Rote Wand and Kieselsandstein.
The latter unit includes a series of dolomitic beds,
the Lehrbergschichten, and, interdigitating with the
shales, a fluvial sandstone (Kieselsandstein).
Similarly, the following Stubensandstein can be
subdivided into fluvially dominated deposits (Sand
-
steinkeuper) protruding from the margins into the
German Basin towards the north and northwest,
where they interdigitate with variegated playa pe
-
lites and lacustrine dolomites (Steinmergelkeuper)
in the depositional centre of the German Basin. The
marginal deposits comprise four heterochronous
large-scaled alluvial fans termed first to fourth Stu
-
bensandstein. Note that the next following unit, the
Knollenmergel, in parts evidently represent a het
-
erochronous facies deposited in a playa environ
-
ment: in southern Württemberg, the lower beds of
the Knollenmergel replace the third and fourth Stu
-
bensandstein laid down contemporaneously in cen
-
tral and eastern Württemberg (B
&
V
, 1981). In northern and eastern Ger
-
many, the Knollenmergel is topped by thick fluvial
sandstones ("Hauptsandstein") grading into flood
-
83
A REVIEW OF THE EUROPEAN TRIASSIC THEROPODS
A B
plain deposits, which both are by convenience re
-
ferred to the Lower Rhaetian. These again are
overlain unconformably by marine sandstones and
shales, which can be safely dated as Rhaetian by a
fauna including the bivalve Rhaetavicula contorta
(P
). In South Germany, the Rhaetian is rep
-
resented by much condensed and patchy deltaic de
-
posits ("Rhät-sandstone" and "Rhät-shales")
following the marine Rhaetic transgression.
About half of the taxa treated here are derived
from the South German Stubensandstein, and for
this reason it is necessary to review the stratigraphic
position of this unit in more detail. The specimens
from Pfaffenhofen (Procompsognathus triassicus,
the theropod nomen dubium Halticosaurus longo
-
tarsus, and the archosaurian "Halticosaurus" orbito
-
angulatus) can be safely placed into the middle
Stubensandstein (S
, 1929; B , 1978).
Much more difficult to establish is the lithostra
-
tigraphic position of Tanystrophaeus posthumus
(H
) and Dolichosuchus cristatus (K
), since there are numerous abandoned Stuben
-
sandstein quarries around Stuttgart and the exact
provenance of the specimens was never recorded.
The Heslach quarry, also the source of a number of
important phytosaur remains and the rauisuchian
Teratosaurus M
(K , 1859; M , 1861;
G
, 1985b), corresponds probably to the site
"Heslacher Wand", which is most likely middle Stu-
bensandstein (B
, 1978). The quarry near
Kaltental which produced the famous association of
22 specimens of the stagonolepidid Aetosaurus
F
was placed in the lower Stubensandstein
(W
, 1989), but it is by no means sure whether Do
-
lichosuchus comes from the same site. Additionally,
all the lithostratigraphic referrals are debated, and
the only safe conclusion is that both specimens must
have been derived from lower or middle Stuben
-
sandstein.
Two alternative hypotheses have been advo
-
cated to calibrate the lithostratigraphic units of the
German Keuper Succession (discussed in B
,
1994a, 1994b). Both unanimously agree that the
Stubensandstein must be considered Norian in age,
but conflicting palynologic, magnetostratigraphic
and palaeoclimatic evidence has led to different as
-
sessments of the time span of this unit within the
stage. In the first interpretation, the Stubensand
-
stein comprises almost the whole Norian, except for
the Knollenmergel, which is commonly referred to
the uppermost part of the Norian. According to the
alternative view, the Stubensandstein is restricted to
the upper part of the Norian only. Currently, there is
not sufficient data available to correlate one of the
subunits of the Stubensandstein with the alpine Tri
-
assic strata convincingly and to clinch this question.
W
(1989) assigned a Middle Norian age to the
lower Stubensandstein because of the co-
occurrence of Aetosaurus therein and in the datable
Calcare di Zorzino in North Italy. This would point to
-
wards a Middle to Late Norian Age for the whole Stu
-
bensandstein. However, as outlined before, the
referral of Aetosaurus to the lower Stubensandstein
is debatable, and it could well occur in the middle
Stubensandstein (e.g. B
, 1973). Further
-
more, L
&H (1993) proposed the Revuel
-
tian biochron (Lower Norian) for the Southwestern
United States, characterised, among other verte
-
brates, by the aetosaur Paratypothorax L
&B
(see also H &L , 1992). Para-
typothorax occurs in the lower Stubensandstein
(W
, 1991), but also in the middle (A.H., unpubl.
data), arguing for an Early Norian age of both
subunits. However, the utility of vertebrate bio
-
chrons for dating strata on widely separate conti
-
nents has yet to be tested. We consider the
Stubensandstein spanning most of the Norian as the
more likely interpretation at the moment. Both hy
-
potheses may be tested when dates are established
for specimens referable to the phytosaur Mystrio-
suchus F
from the marine Dachsteinkalk of the
Austrian Alps (reported preliminarily by B
,
1994). In the German Basin, Mystriosuchus is en-
tirely restricted to the middle Stubensandstein.
Liliensternus liliensterni comes from the Knollen-
mergel of Thuringia and can therefore be dated as
Late Norian with some certainty (see above). This
conclusion is furthermore substantiated by the fact
that the remains of L. liliensterni were found in asso-
ciation with remains of Plateosaurus engelhardti
(H
, 1934), which is well known from the Late
Norian of southern Germany and Switzerland
(S
, 1992).
Syntarsus sp. comes from Pant-y-ffynnon
quarry, one of the numerous fissure fillings in Car
-
boniferous limestone strata of Wales and Southwest
England. These fillings have a complex depositional
history and dating is difficult (see F
, 1994),
being mainly achieved by their contents of verte
-
brates. Pant-y-ffynnon has yielded, among other
taxa, the crocodylomorph Terrestrisuchus C
(which has been suggested as a synonym of the
middle Stubensandstein Saltoposuchus) and the
gliding diapsid Kuehneosaurus R
. Verte
-
brates seem to exclude a post-Triassic age, but the
locality can be considered Late Triassic at best,
ranging anywhere from Late Carnian to Rhaetian
(B
&S , 1995).
The age of Pterospondylus trielbae is usually
given as Rhaetian (e.g. H
, 1932; N ,
1990). The section in the Baerecke quarry, as sum
-
marized by S
(1992), consisted of two suc
-
cessions: a lower claystone layer, which is
considered to represent the equivalent of the Knol
-
84
O.W.M. RAUHUT & A. HUNGERBÜHLER
lenmergel, overlain by sand- and siltstones of proba
-
bly Rhaetian age. According to J
(1913), at
least a part of the turtle remains were derived from
the lower part of the section of the Baerecke quarry.
Since the holotype of P. trielbae was found within a
turtle carapace, it might therefore be of Late Norian
age, rather than Rhaetian.
"Zanclodon" cambrensis comes from a sand
-
stone bed in southern Wales. The slab was derived
from a stack of building material, and its exact stra
-
tigraphic position (above or below the Rhaetavicula
contorta beds) is uncertain (N
, 1899). How
-
ever, the sandstone unit, where the specimen came
from is considered to be Rhaetian (W
et
al., 1980).
The stratigraphic position of ?Liliensternus
airelensis has been discussed in some detail by
C
&G (1993). The specimen might be ei
-
ther late Rhaetian or early Hettangian in age.
T
T T F E
Only three theropod species from the Upper Tri
-
assic of Europe can presently be regarded as being
valid (see also T
I): Liliensternus liliensterni,
?Liliensternus airelensis and Procompsognathus
triassicus. Furthermore, a species of Syntarsus was
most probably present in the Upper Triassic of
Europe as well. Both L. liliensterni and P. triassicus
come from the Norian of southern Germany, the
specimen of Syntarsus derives from probably No
-
rian fissure fillings in Wales, while ? L. airelensis was
found in sediments spanning the Triassic-Jurassic
boundary in northern France, and might even be of
lowermost Jurassic age.
Apart from the remains discussed above, only a
few other fragments can be assigned to the Thero
-
poda with any certainty. G
(1985a) noted a
probable theropod femur fragment from the Stuben
-
sandstein of Pfaffenhofen, B
&W
(1986) described some probable theropod teeth
85
A REVIEW OF THE EUROPEAN TRIASSIC THEROPODS
N
AME
A
GE
O
CCURENCE
T
AXONOMIC
S
TATUS
Avipes dillstedtianus
H
UENE
, 1932
Ladinian
Lettenkeuper, Thuringia,
Germany
? Archosauria, nomen
dubium
Dolichosuchus cristatus
H
UENE
, 1932
Norian Stubensandstein, Baden-
Württemberg, Germany
Theropoda, nomen
dubium
Halticosaurus longotarsus
(H
UENE
, 1907-8)
Norian Stubensandstein, Baden
Württemberg, Germany
Theropoda, nomen
dubium (at least partim)
"Halticosaurus" orbitoangulatus
H
UENE, 1932
Norian Stubensandstein, Baden-
Württemberg, Germany
? Crocodylomorpha,
nomen dubium
? Liliensternus airelensis
C
UNY &GALTON 1993
Rhaetian-
Hettangian
Airel quarry, Normandy,
France
Theropoda, valid species
Liliensternus liliensterni
(H
UENE
, 1934)
Norian Knollenmergel, Thuringia,
Germany
Theropoda, valid species
Procompsognathus triassicus
F
RAAS
, 1913
Norian Stubensandstein, Baden
Württemberg, Germany
Theropoda, provisionally
regarded as valid species
Pterospondylus trielbae
J
AEKEL, 1913
? Norian Knollenmergel, Thuringia,
Germany
Theropoda, nomen
dubium
Saltopus elginensis
H
UENE
, 1910
Carnian Lossimouth Sandstone
Formation, Scotland
Dinosauriformes,
nomen dubium
Tanystrophaeus posthumus
H
UENE, 1907-8
Norian Stubensandstein, Baden-
Württemberg, Germany
Theropoda, nomen
dubium
Velocipes guerichi
H
UENE
, 1932
Norian Lissauer Breccia, Gorny
Slask, Poland
Vertebrata, nomen dubium
"Zanclodon" cambrensis
N
EWTON, 1899
Rhaetian Rhaetic sandstone, Wales Theropoda, nomen
dubium
from the Norian of France, and S (1992) re
-
ferred several teeth from the Knollenmergel of Frick
(Switzerland) and Trossingen (southern Germany)
to cf. Liliensternus. However, since teeth of thero
-
pod dinosaurs do not differ significantly from the ser
-
rated teeth of other Triassic archosaurs, the
identification of isolated serrated teeth from Triassic
beds as theropod is always doubtful. For the same
reason, taxa based on isolated teeth (e.g. "Megalo
-
saurus" cloacinus Q
,"Megalosaurus" ob
-
tusus H
,"Plateosaurus" ornatus H ; see
H
, 1907-8, 1932; G , 1985b) are not in
-
cluded in this review.
In addition to the skeletal remains, supposed
theropod footprints have been described from differ
-
ent localities (e. g. H
, 1990). However, there
are many sources of error in identifying prints (K
&B , 1996), and, apart from theropods, other
animals including protodinosaurs or early ornithis
-
chians might have produced tridactyl prints, hence
footprint evidence is not taken into consideration
here.
Thus, the earliest certain records of theropod di-
nosaurs in Europe come from the upper parts of the
Lower Norian. Since there are vertebrate localities
in the pre-Norian Upper Triassic in Europe, and es-
pecially in Germany, the Upper Triassic is well repre-
sented by terrestrial sediments (see above), the
appearance of theropod dinosaurs at that time is in-
terpreted here as representing the first radiation of
theropods in Europe.
Furthermore, all identifiable theropod remains
appear to belong to the Coelophysoidea, within the
Ceratosauria. The presence of the genus Syntarsus
in particular is interesting; this taxon is well known
from the lowermost Jurassic of southern Africa and
North America (R
&G , 1990), and
might have been present in the Norian of North
America as well (P
, 1993). Since the oldest
theropods are known from the Carnian of South
America (S
&N , 1992), and coelophy
-
soids first appear in the fossil record in the early No
-
rian (R
&G , 1990), there would appear
to have been a rapid radiation of ceratosaurian
theropods in the late Carnian / early Norian. These
conclusions are in general accordance with ideas
proposed by B
(1984, 1993a), that dinosaur
radiation was rapid in the Upper Triassic, possibly
following an extinction event.
An interesting aspect of the Upper Triassic thero
-
pod fossil record, not only of Europe, but in general,
is the complete absence of theropods other than
herrerasaurids and ceratosaurs. Given the sister-
group relationship of ceratosaurs and tetanurans
(G
, 1986; H , 1994), the latter group
must have been present in the Late Triassic. The
lack of tetanuran fossils from the Triassic either re
-
flects geographic isolation of this group during that
time, or their rareness in Upper Triassic vertebrate
faunas.
In this context, another aspect is worth mention
-
ing: there are many fossil vertebrate localities, and
dinosaurs are quite abundant in the European Up
-
per Triassic (see e.g. W
, 1990), but thero
-
pod remains are rather rare. Although this may
partly be due to the taphonomy of the localities
(S
, 1992), it might reflect genuine rarity of
theropods in the Upper Triassic vertebrate fauna of
Europe.
ACKNOWLEDGEMENTS
We thank Sandra Chapman, London, W.-D.
Heinrich, Berlin, and R. Wild, Stuttgart, for access to
specimens under their care. G. Cuny, Bristol, is
thanked for providing casts of L. airelensis and for
translating the abstract into French. Special thanks
are due to Dave Unwin and Mike Benton, both Bris
-
tol, for comments on the manuscript. Thanks are
also due to Ben Edwards, London, for bringing the
fissure-filling specimens to our attention. Simone
Klutzny, Bristol, is thanked for discussions while this
paper was written. We thank the referees, Thomas
Holtz, College Park, and Bernardino Pérez-Moreno,
Madrid, for useful comments. O.R. is supported by
the EC under TMR grant ERBFMBICT 961013 and
A.H. under ERBCHBICT 930521, which is gratefully
acknowledged.
ABBREVIATIONS
BMNH - Natural History Museum, London, UK;
GPIT - Institut und Museum für Geologie und
Paläontologie Tübingen, Germany; HMN - Museum
für Naturkunde, Humboldt Universität, Berlin, Ger
-
many; SMNS - Staatliches Museum für Naturkunde
Stuttgart, Germany.
REFERENCES
86
O.W.M. RAUHUT & A. HUNGERBÜHLER
87
A REVIEW OF THE EUROPEAN TRIASSIC THEROPODS
88
O.W.M. RAUHUT & A. HUNGERBÜHLER
