501-Özi-5.8. 20.11.

2007 12:13 Uhr Seite 173 (Schwarz Bogen)

N. Jb. Geol. Paläont. Abh.

2007, vol. 246/2, p. 173 – 181, Stuttgart, November 2007, published online 2007

A maniraptoran (Theropoda, Dinosauria) sacrum from the

Upper Cretaceous of the Haţeg Basin (Romania) – in search
of the lost pterosaurs of Baron FRANZ NOPCSA
Attila Ősi and Istv án F őzy, Budapest
With 2 figures

ŐSI, A. & FŐZY, I. (2007): A maniraptoran (Theropoda, Dinosauria) sacrum from the Upper
Cretaceous of the Haţeg Basin (Romania) – in search of the lost pterosaurs of Baron FRANZ NOPCSA.
– N. Jb. Geol. Paläont. Abh., 246: 173–181; Stuttgart.

Abstract: A fragmentary sacrum from the Upper Cretaceous of the Haţeg Basin is described here.
On the basis of a combination of traits, such as large pleurocoels on the first sacral, a prominent
neural platform and completely fused neural spines dorsally, it is referred here to Paraves. Com-
parison with other non-avian theropod sacra indicates that the sacrum of Ornithodesmus cluniculus
from Barremian of the Isle of Wight is the most similar to the Haţeg specimen. The specimen is
also compared with the sacrum of birds and pterodactyloid pterosaurs. The size of the bone suggests
a total body-length of about 1.2-1.5 meters for the animal.

Key words: sacrum, Paraves, Dromaeosauridae, Late Cretaceous, Haţeg Basin.

1. Introduction After NOPCSA’s last mention on Transylvanian

Pterosaur remains from the Upper Cretaceous of the
Haţeg Basin have been known since 1899 (NOPCSA
1899). Later, NOPCSA repeatedly mentioned pterosaur
remains from the area (e.g. NOPCSA 1905, 1923),
but never gave a list, a detailed description, or any
illustration of the available specimens (for a review of
the story of NOPCSA’s Transylvanian pterosaurs, see
BUFFETAUT et al. [2003]). The first detailed report of
pterosaur remains including figures from the area was
provided by BUFFETAUT et al. (2002), 103 years after
the first mention of pterosaurs. This material, how-
ever, represents newly discovered fossils which belong
to a large-sized pterosaur, Hatzegopteryx thambema
(BUFFETAUT et al. 2003).
pterosaurs (NOPCSA 1926) no information was
available about these fossils till 1995, when CORALIA
JIANU and DAVID WEISHAMPEL rediscovered one
specimen (Ob. 1948) in the Vertebrate Collection of
the Geological Institute of Hungary (MÁFI) (JIANU
et al. 1997). The authors considered this specimen as
being part of NOPCSA’s pterosaur collection, but it was
neither figured nor described.
With the help of LÁSZLÓ KORDOS we also checked
the Vertebrate Collection of the MÁFI, where we
relocated specimen Ob. 1948, but no other pieces of
NOPCSA’s mysterious pterosaur remains. Specimen
Ob. 1948 is catalogued as “Pterosauria sacrum from
Szentpéterfalva” with a remark that it was in the
possession of BARON NOPCSA.

DOI: 10.1127/0077-7749/2007/0246-0173 0077-7749/07/0246-0173 $ 2.25

© 2007 E. Schweizerbart’sche Verlagsbuchhandlung, D-70176 Stuttgart
501-Özi-5.8. 20.11.2007 12:13 Uhr Seite 174 (Schwarz Bogen)

174 A. Ősi and I. Főzy

Fig. 1 (Legend see p. 175)

501-Özi-5.8. 20.11.2007 12:13 Uhr Seite 175 (Schwarz Bogen)

A maniraptoran sacrum from the Upper Cretaceous of the Haţeg Basin (Romania) 175

This bone (Figs. 1-2) is a fragmentary sacrum Paraves indet.

described here in detail. Due to the lack of earlier
illustrations or descriptions, however, it is impossible
to identify which of NOPCSA’s publications include
reference to this bone. Nevertheless, we suggest that
this sacrum could be the specimen that NOPCSA first
referred as a Coeluride (?) sacrum (NOPCSA 1905).
Probably it is the same bone that was later reinter-
preted by NOPCSA (1914) as a pterosaur notarium
(BUFFETAUT et al. 2003).
NOPCSA in 1915 and later in 1923 mentioned simi-
larity between the Haţeg pterosaur material and the
remains of Ornithodesmus. One can only speculate
whether this similarity is based on a comparison of
the Ob. 1948 sacrum with that of Ornithodesmus
cluniculus SEELEY, 1887 (BMNH R187) from the
Barremian of the Isle of Wight, as discussed below,
or NOPCSA studied other specimens that were sub-
sequently lost, and compared them with “Ornitho-
desmus” latidens SEELEY, 1901 (redescribed as
Istiodactylus latidens by HOWSE et al. [2001]).
The specimen Ob. 1948 does not resemble the
sacrum of pterodactyloid pterosaurs, but rather it is
more similar to that of maniraptoran dinosaurs,
especially to that of dromaeosaurids and birds.
Institutional abbreviations: AMNH, American Mu-
seum of Natural History, New York, USA; BMNH, Natural
History Museum, London United Kingdom; IGM, Mon-
golian Institute of Geology, Ulan Bataar, Mongolia; MÁFI,
Geological Institute of Hungary, Budapest, Hungary;
MCNA, Museo de Ciencias Naturales de Alava; MDE,
Musée des Dinosaures, Espéraza, France; SMC, Sedgwick
Museum, Cambridge, England; TMP, Royal Tyrrell
Museum of Paleontology, Drumheller, Canada.
2. Systematic paleontology
Dinosauria OWEN, 1842
Theropoda MARSH, 1881
Maniraptora GAUTHIER, 1986
Paraves SERENO, 1997 sensu HOLTZ & OSMÓLSKA
(2004)
Figs. 1-2
Refer red specimen: MÁFI Ob. 1948 fragmentary
sacrum, collected by GYULA HALAVÁTS.
Locality and horizon: Sinpetru (Szentpéterfalva),
Haţeg Basin, western Romania.
Age: Maastrichtian (GRIGORESCU & CSIKI 2002).
3. Description and comparison with
non-avian theropods
The specimen Ob. 1948 is a 44 mm long, anterior part
of a fragmentary sacrum that is slightly arched along
its length (Fig. 1E-H). It consists of two completely
ankylosed vertebral centra. The partial neural arches
form a co-osssified, rigid structure extending from the
anterior end of the second centrum presumably to the
level of the end of the third vertebral centrum. This
massive, composite neural arch is crushed both
anteriorly and posteriorly suggesting at least four
fused sacral vertebra. The first preserved centrum is
relatively deep, almost as high as wide. The anterior
articular surface, although slightly crushed dorsally,
appears to be slightly concave. The ventral surface of
this centrum is slightly concave anteroposteriorly
and convex transversely. This centrum is similar to
the first centrum of the sacrum of the possible
dromaeosaurid Ornithodesmus (HOWSE & MILNER
1993; NORELL & MAKOVICKY 1997, 2004; BARSBOLD
& OSMÓLSKA 1999; NAISH et al. 2001) but the
anterior and posterior ends of Ornithodesmus are
more expanded transversely.
The sacra of alvarezsaurids (e. g. Patagonykus,
Mononykus) differs from the Haţeg specimen in
having transversely compressed centra (CHIAPPE et al.
2002). On the studied specimen (MÁFI Ob. 1948) the
junction between the first and the second centra is not
visible due to the complete ankylosis of the bones,
in addition that area is slightly crushed ventrally
(Fig. 1C, D). Furthermore, no characteristic transverse
widening of the junction is observed, in contrast with

Fig. 1. Fragmentary sacrum from the Upper Cretaceous of the Haţeg Basin (MÁFI Ob. 1948). A-B – dorsal view;
C-D – ventral view; E-F – left lateral view; G-H – right lateral view. Abbreviations: ass1, anterior surface of the first sacral
vertebra; cav, cavity; cns, co-ossified neural spines; ctrp, crushed transverse process; dsnc, dorsal surface of the
neural canal; fzy, fused zygapophyses; inf, intervertebral fenestra; lp, lateral pillar; nc, neural canal; np, neural plat-
form; plc, pleurocoel; pre, prezygapophysis; s1, first sacral vertebra; s 2, second sacral vertebra; trp, transverse process;
vt, ventral trough.
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176 A. Ősi and I. Főzy

Fig. 2 (Legend see p. 177)

501-Özi-5.8. 20.11.2007 12:13 Uhr Seite 177 (Schwarz Bogen)
A maniraptoran sacrum from the Upper Cretaceous of the Haţeg Basin (Romania)
that of dromaeosaurids and oviraptorosaurids. The
complete fusion of the two centra, however, resembles
the synsacrum of birds rather than the sacrum of
Velociraptor mongoliensis (IGM 100/986, NORELL
& MAKOVICKY 1999) or that of Sinornithosaurus
millenii (XU et al. 1999) where the junctions are
well-exposed. In ventral view, the sacral centra of
Sinornithosaurus strongly differ from that of the
Haţeg sacrum in having deeply concave ventral sur-
faces both anteroposteriorly and transversely (XU et
al. 1999). On the basis of the ventral shape of Ob.
1948, a nearly equal length of the two centra or
perhaps a slightly greater length of the second
centrum is suggested. Similar to Ornithodesmus,
(HOWSE & MILNER 1993) the second centrum has
flattened ventral faces and it shows neither transverse
convexity, nor anteroposterior concavity. The second
centrum is the last preserved among the centra of Ob.
1948, and the posterior half of its ventral surface bears
a shallow and wide medial trough ventrally (Fig. 1D).
This is not similar to those mentioned in Ornitho-
desmus (HOWSE & MILNER 1993) but rather (although
not completely) similar to the groove seen on the
fourth centrum of the possible Saurornitholestes
sacrum of BMNH 4463 (HOWSE & MILNER 1993, fig.
6 originally interpreted as being from a troodontid but
see NORELL & MAKOVICKY 1997: 22, and LE LOEUFF
& BUFFETAUT 1998: 109). This structure is also
similar to the ventral groove of Variraptor mechi-
norum (MDE –D169, LE LOEUFF & BUFFETAUT 1998)
between the third and fourth sacrals. Important to note
that the development of this ventral groove is variable
even among different individuals of one genus (e. g.
Saurornitholestes), and it appears more frequently in
older individuals (P. MAKOVICKY, pers. comm.).
The posterior end of the second centrum is strongly
divergent, indicating that the transverse processes of
the third sacral overlapped the junction between the
third and fourth sacrals (Fig. 1C, D). The first sacral
vertebra bears large pleurocoels, which open laterally
below the transverse processes (Fig. 1G, H). These
pleurocoels have a longitudinally elongate, oval shape
similarly to those of the first sacral of Ornithodesmus
(HOWSE & MILNER 1993). In contrast to Variraptor
Fig. 2. Fragmentary sacrum from the Upper Cretaceous
of the Ha¸teg Basin (MÁFI Ob. 1948). A-B – anterior view;
C-D – posterior view. For abbreviations see Fig. 1.
177
(LE LOEUFF & BUFFETAUT 1998), Ornithodesmus
(HOWSE & MILNER 1993), Saurornitholestes langstoni
(TMP 67.20.36, NORELL & MAKOVICKY 1997), and
Velociraptor (IGM 100/986; NORELL & MAKOVICKY
1999) no pleurocoels are present on the second sacral
vertebra. Presence of pleurocoels on the first sacral,
however, is not a unique feature of dromaeosaurids,
because other theropod groups e.g. allosaurids and
some coelurosaurs (HOLTZ et al. 2004), or caenag-
nathids (OSMÓLSKA et al. 2004; P. MAKOVICKY pers.
comm.) and may be the therizinosauroid Alxasaurus
elesitaiensis (CLARK et al. 2002) possess these
openings.
The transverse processes originate at progressively
lower levels from the first to the third sacral (Fig.
1E-H). The transverse processes of the first sacral
arise close to the posterior end of the vertebra postero-
dorsal to the pleurocoel. They are directed laterally
and upward with an angle of 45°. The transverse
processes of the second vertebra are slightly damaged
at their distal ends and are directed laterally and
upward with an angle of approximately 35°. Laterally
these broken processes have a vertically elongate
triangular cross-section. Furthermore, their ventral
sides are concave and together with lateral excavations
on the centrum they form a concave triangular area.
This is present in the case of first sacral too, but also
including the pleurocoels. Transverse processes of the
third sacral are only stumps preserved in a low level
on the posterior end of the second centrum.
It is not clear whether or not the prezygapophyses
of the first sacral were fused together with the
postzygapophyses of the last dorsal (Fig. 1E, F).
Posteriorly, however, the postzygapophyses of the first
sacral are fused with the prezygapophyses of the
second sacral. This feature is present between the
zygapophyses of the second and third vertabrae, too.
Thus the zygapophyses form a sinuous ridge on
both sides of the sacrum that appears to be a dromaeo-
saurid feature (NORELL & MAKOVICKY 1997, 2004).
Dorsally, these fused segments extend into trans-
versely flattened lateral pillars of the neural arch (Fig.
1F). Deep, almost rectangular cavities are situated
between these pillars. This structure is clearly ob-
served on the sacrum of Ornithodesmus in right lateral
view (HOWSE & MILNER 1993, text-fig. 1B). The
neural canal has an almost circular cross-section
anteriorly (Fig. 2A, B), while posteriorly it is 1.5
times wider than high and possesses much greater
linear dimensions (Fig. 2C, D).
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178 A. Ősi and I. Főzy
The completely ankylosed neural arch is a massive
structure and almost rectangular in cross-section. In
lateral view, it descends ventrally at about 15° to the
axis of the sacrum (Fig. 1E-H). The damaged neural
spines are represented only by their bases but they
are completely co-ossified to form a continuous
blade-like structure dorsally, similar to that of
Ornithodesmus, the sacrum of BMNH 4463, Velo-
ciraptor, Varirpator and Deinonychus (OSTROM 1976;
HOWSE & MILNER 1993; NORELL & MAKOVICKY
1997; LE LOEUFF & BUFFETAUT 1998). The fusion of
neural spines, however, is not an exclusive dromaeo-
saurid feature. Apart from this group, it appears also
in ceratosaurs (TYKOSKI & ROWE 2004), tyrannosaurs
(HOLTZ 2004), and at least on the cranial and central
portions of the sacrum of alvarezsaurids (CHIAPPE
et al. 2002). In oviraptorosaurs neural spines contact
but do not fuse to each other (OSMÓLSKA et al. 2004).
This continuous neural blade is thickened ventrally
by fusing continuously with the lateral laminae of the
neural platform (Fig. 1A, B). This neural platform is
supported ventrally by the above mentioned lateral
pillars that extend dorsally from the fused zygapo-
pyseal constructions.
4. Comparison with birds and ptero-
dactyloids
Comparison of the Haţeg sacrum with that of birds
indicated several similarities. For example, the fusion
of the first two centra without any suture, the great
expansion of the posterior end of the second centra
involving the third transverse process, as well as the
form and position of the anterior transverse processes
and the fused neural spines forming a massive neural
blade (e. g. in Rahonavis [FORSTER et al. 1998] and
even in modern birds), suggest a close relationship.
The transverse processes on the third sacral are only
stumps preserved in a low level on the posterior end of
the second centrum. These processes widen greatly
at the posterior end of the second vertebra; a similar
structure appears also on the anterior sacrals of
several different groups of recent birds (e.g. Struthio
camelus, Otis tarda, Bubo bubo) and shows the area
where the ilia fit the closest to the sacrum. Gobipteryx
minuta (KUROCHKIN 1996) and some El Brete spe-
cimens (CHIAPPE & WALKER 2002) bear a shallow
ventral groove on its centra that is similar to that
metioned in Ob. 1948.
Some features presented in Ob. 1948, however, are
not found in birds. A single great pleurocoel below the
transverse process of the first sacral and the well-
developed neural platform do not appear so markedly
in birds. In addition, no median ridge on the ventral
surface of the anterior sacrals is present, which is
prominent anteriorly on the synsacrum of several
fossil and recent birds (e. g. Apsaravis ukhaana [IGM
100/1017] CLARKE & NORELL 2002; Gargantuavis
philoinos [MDE C3-525] BUFFETAUT & LE LOEUFF
1998; Enaliornis barretti [SMC B55282] GALTON &
MARTIN 2002; Corvus corax). The synsacrum com-
posed by at least 8 centra in Enantiornithes differs
from Ob. 1948 in having laterally compressed centra
and in the lack of pleurocoels (CHIAPPE 1996). Un-
fortunately, the number of sacrals, as a characteristic
feature of the bird synsacrum, is not recognizable on
Ob. 1948.
During our study of the Haţeg sacrum we also
made comparisons with the sacrum of pterodactyloid
pterosaurs. These examinations indicated several
differences between the sacrum of pterodactyloids and
the Haţeg specimen. It is mportant to note, however,
that the sacrum is known only in a few pterodactyloid
taxa. Among azhdarchids, the only group of con-
firmed pterosaurs represented in the area (BUFFETAUT
et al. 2002), Zhejiangopterus is the only genus
with preserved sacrum but it “is obscured by the
pelvic girdle prohibiting its description” (CAI & WEI
1994).
The sacral series of a partially preserved ptero-
dactyloid pelvis (AMNH 22569) from the Aptian of
Brazil (BENNETT 1990), similarly to other ptero-
dactyloid sacra, differs from the Haţeg sacrum in the
lack of a neural platform. In addition, in AMNH
22569 the neural spines are separated elements and
probably fused only at their top as a supraneural plate,
similar to Pteranodon ingens (EATON 1910) and to
Coloborhynchus spielbergi (VELDMEIJER 2003). The
unfused neural spines are also characteristic in
Anhanguera piscator (KELLNER & TOMIDA 2000) and
in the anterior sacrals of Anhanguera santanae
(WELLNHOFER 1991). The low and anteroposteriorly
wide transverse processes of the anterior sacrals
present in Pteranodon, Anhanguera, and in Colobor-
hynchus, in addition to the lack of pleurocoels, also
distinguish them from the Haţeg sacrum. The sacrum
of Istioactylus latidens (HOWSE et al. 2001) is charac-
terized by progressive narrowing of centra posteriorly,
obliquely backwards-oriented transverse processes
and the absence of a neural platform and a ventral
trough which clearly distinguish it from the Haţeg
specimen.
501-Özi-5.8. 20.11.2007 12:13 Uhr Seite 179 (Schwarz Bogen)
A maniraptoran sacrum from the Upper Cretaceous of the Haţeg Basin (Romania)
A sacrum (MCNA 2583) composed of at least five
vertebrae from the Late Cretaceous of Laño (Spanish
Basque Country, BUFFETAUT 1999) is rather more
similar to the sacrum from the Haţeg Basin in the
lack of intervertebral sutures. BUFFETAUT (1999)
shortly described MCNA 2583 as cf. Azhdarcho, but
mentioned the possibility that it may represent a bird.
The above comparisons clearly indicate the mani-
raptoran affinity of the Haţeg sacrum and appear to
exclude its pterodactyloidean relationship. Several of
the above mentioned features (e.g. general form of the
vertebral centra, the completely fused neural arch,
shape of transverse processes) are shared also with
dromaeosaurids and birds. Nevertheless, we em-
phasize that due to the fragmentary nature of Ob.
1948, several of important characters (e.g. the total
number of sacrals, the presence or absence of pleuro-
coels and the ventral groove posteriorly, as well as the
shape and orientation of the transverse processes)
remain unknown.
Thus, here we tentatively refer the Haţeg sacrum to
the clade Paraves, with the comment that future dis-
coveries of more complete specimens may help to
identify it more precisely.
5. Discussion
Although the fragmentary nature of the sacrum of
Ob. 1948 prevents us from identifying the total
number of co-ossified vertebrae, the above mentioned
features may suggest the dromaeosaurid affinities
of the specimen, and thus five or six sacrals could
be supposed (NORELL & MAKOVICKY 1997, 2004).
OSTROM (1976) reported that in old individuals the
last dorsal might have been co-ossified with the first
sacral to form a sacrum composed of six vertebrae.
Thus the first vertebra of the Haţe g sacrum may re-
present the last dorsal. However, on the basis of the
complete ankylosis (no suture is observed) of the first
two preserved vertebral centra, it is suggested here
that they more likely represent the first two sacrals.
Furthermore the strong ankylosis between the first
two vertebrae (Fig. 1C, D) indicates maturity. Com-
parison of the size of the Haţeg sacrum with that of
dromaeosaurids suggests a small-bodied animal. In
Ob. 1948 the length of the first two sacrals is appro-
ximately 30 mm (note the fragmentary condition of
the posterior part of the second centrum). This length
is 33 mm in Ornithodesmus (HOWSE & MILNER 1993),
34.8 mm (IGM 100/985), 39.6 mm (IGM 100/985)
in Velociraptor (NORELL & MAKOVICKY 1999), and
28.2 mm in Sinornithosaurus millenii (XU et al.
179
1999). This indicates a total body length of about
1.2-1.5 meters for the Haţeg maniraptoran that is
almost similar to that of Ornithodesmus.
Although remains of theropod dinosaurs from the
Haţeg Basin have been known since the time of
NOPCSA (ANDREWS 1913; NOPCSA 1915), they are few
in number, and except for some associated material
(CSIKI & GRIGORESCU 2005), most of the specimens
represent fragmentary bones and teeth. For a review of
theropod remains of the area see CSIKI & GRIGORESCU
(1998) and KESSLER et al. (2005). Thus the sacrum,
presented here, is very important for a better re-
cognition of the theropods of the Late Cretaceous
ecosystem of the Haţeg Basin.
The first mention of dromaeosaurid theropods from
the area is by LE LOEUFF et al. (1992), where they
referred the “European elopterygids” to the Dromaeo-
sauridae. Recently, KESSLER et al. (2005) reported a
newly discovered distal femur and reinterpreted the
Elopteryx material as possibly representing alvarez-
saurids.
The first indisputable remains of dromaeosaurid
dinosaurs from the area were described by WEIS-
HAMPEL & JIANU (1996) on the basis of an articulated
frontal and parietal that is apparently closely related to
Saurornitholestes. The size of these bones, however,
suggests a significantly larger animal compared with
the above discussed Haţeg maniraptoran. Later, as a
result of extensive excavations and screen-washing,
several authors (e.g. CSIKI & GRIGORESCU 1998;
GRIGORESCU & CSIKI 2002; CODREA et al. 2002)
mentioned dromaeosaurid teeth from the Haţeg Basin.
The size of these teeth appears to be more similar to
that of the maniraptoran, discussed here.
6. Conclusions
Restudy of the Vertebrate Collection of the Geological
Institute of Hungary provided only a fragmentary
sacrum (Ob 1948; rediscovered by C. JIANU and D.
WEISHAMPEL in 1995) that probably represents a piece
of the assemblage that NOPCSA referred to pterosaurs.
Detailed study and comparison of this specimen with
the sacra of non-avian theropods, birds and pterosaurs,
indicated that the Haţeg specimen is not from a
pterosaur but it is more similar to the sacrum of
dromaeosaurids and birds, especially to that of
Ornithodesmus, thus is referred here tentatively to
Paraves. The size of the Haţeg sacrum indicates a total
body length of about 1.2-1.5 metres for the Haţeg
maniraptoran.
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180 A. Ősi and I. Főzy

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Manuscript received: May 30th, 2007.
Revised version accepted by the Stuttgart editor: July 19th,
2007.
Addresses of the authors:
ATTILA ŐSI 1, 2, ISTVÁN FŐZY 3, 1 Hungarian Academy of
Sciences – Hungarian Natural History Museum, Research
Group for Palaeontology, Ludovika tér 2, 1083 Budapest,
Hungary; 2 Eötvös Loránd University, Department of
Paleontology, Pázmány Péter sétány 1/c, 1117 Budapest,
Hungary; 3 Hungarian Natural History Museum, Ludovika
tér 2, 1083 Budapest, Hungary;
e-mails: hungaros@freemail.hu; fozy@nhmus.hu