Lourinhanosaurus antunesi, A New Upper Jurassic Allosauroid (Dinosauria: Theropoda) from Lourinhã, Portugal

Abstract

A new Allosauroid dinosaur (Saurischia: Theropoda) was found at Peralta near Lourinhã, Portugal (Upper Jurassic, Lower Tithonian). It is described under the name Lourinhanosaurus antunesi. It's diagnosable by the all vertebral centra longer than tall, neural spines of the anterior caudal vertebrae with a well-developed spike- like anterior process, the pubic blade is perforated by a large vertical ellipsoidal foramen and the lesser trochanter is well separate from the main body of the femur in lateral view. In the rib cage were found 32 gastroliths. This is the first non-avian theropod found with gastroliths.

Full text

Lourinhanosaurus antunesi,
A New Upper Jurassic Allosauroid
(Dinosauria: Theropoda)
from Lourinhã, Portugal
Octávio MATEUS1
Abstract
A new Allosauroid dinosaur (Saurischia: Theropoda) was found at Peralta
near Lourinhã, Portugal (Upper Jurassic, Lower Tithonian). It is described under the
name Lourinhanosaurus antunesi. It’s diagnosable by the all vertebral centra longer
than tall, neural spines of the anterior caudal vertebrae with a well-developed spike-
like anterior process, the pubic blade is perforated by a large vertical ellipsoidal
foramen and the lesser trochanter is well separate from the main body of the femur in
lateral view. In the rib cage were found 32 gastroliths. This is the first non-avian
theropod found with gastroliths.
Resumo
Descreve-se uma nova espécie de dinossauro alossauróide (Saurischia:
Theropoda) com o nome Lourinhanosaurus antunesi gen. et sp. nov. descoberto em
Peralta (Lourinhã, Portugal; Jurássico Superior, Titoniano inferior). É diagnosticável
por todos os corpos vertebrais mais compridos que altos, pelo processo anterior do
espinho neural das vértebras caudais anteriores bem desenvolvido e em forma de
espigão, pelo foramen na púbis elipsoidal e o trocanter anterior bem destacado do
resto do fémur em vista lateral. Na caixa torácica foram encontrados 32 gastrólitos.
Este é o primeiro terópode (excluindo as aves) conhecido com gastrólitos.
1. GEAL- Museu da Lourinhã, Departamento de Paleontologia, Rua João Luis de Moura, 2530
Lourinhã, Portugal; Email: museulourinha@mail.telepac.pt. Centro de Estudos Geológicos, Faculdade
de Ciências e Tecnologia da UNL, 2925-114 Caparica, PORTUGAL
Memórias da Academia de Ciências de Lisboa, 37: 111-124 1998

MEMÓRIAS DA ACADEMIA DE CIÊNCIAS DE LISBOA
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INTRODUCTION
Mesozoic reptile remnants are known in Portugal since long ago:
Sauvage (1897-98), Zbyszewski (1946), Lapparent and Zbyszewski (1957),
Antunes (1976) and Galton (1980)(1991) among others. Recently a theropod
nest with about 100 eggs was found in Paimogo, Lourinhã (Mateus et al.,
1997) about 7 km from the site where the specimen described here was found.
In the Lourinhã area (mainly Lower Tithonian and Upper
Kimmeridgian levels) were discovered several dinosaur remains ascribed to
Brachiosaurus atalaiensis (Sauropoda), Dacentrurus armatus (Stegosauria),
Dracopelta zbyszewskii (Ankylosauria), Hypsilophodon sp. (Ornithopoda),
Megalosaurus insignis and Megalosaurus pombali (Theropoda regarded as
nomina dubia by Molnar, 1990) as well as an unidentified Ceratosauria, other
remains from unidentified sauropods and unidentified Ornithopod (Galton,
1980; Galton, 1991; Helmdach, 1973-74; Lapparent & Zbyszewski, 1957; and
unpublished data).
In 1982 a farmer (Mr. Luis Mateus) discovered some fossil bones from
one individual in anatomical connection that he had removed and later
delivered to the GEAL- Museum of Lourinhã. Parts of the bones were already
without the hard sediment cover. Unfortunately the block containing the
dinosaur was broken in order to make the transportation easier. The resulting
fragments are kept at the GEAL- Museum of Lourinhã.
SYSTEMATIC PALAEONTOLOGY
Dinosauria Owen, 1842
Saurischia Seeley, 1888
Theropoda Marsh, 1881
Avetheropoda Paul, 1988
Allosauroidea Currie & Zhao, 1993
Lourinhanosaurus gen. nov.
Lourinhanosaurus antunesi gen. et sp. nov.
ABBREVIATIONS:
GEAL - Grupo de Etnologia e Arqueologia da Lourinhã;
ML - Museum of Lourinhã.

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113
Etymology- Derivatio nominis
Lourinhanosaurus refers the place (Lourinhã, Portugal) where the
specimen was found; antunesi after Prof. Miguel Telles Antunes, Portuguese
palaeontologist.
Holotype
ML370. Remnants from a single individual (see fig. 1): axial skeleton
(six cervical vertebrae with 6 ribs, five dorsal vertebrae with ribs, 5 sacral
vertebrae, 14 caudal vertebrae with 8 chevrons). Appendicular skeleton:
femora (left femur without tibial and fibular condyles; in right femur lacks the
proximal end); proximal part of the right tibia and fibula in anatomical
connection; two ilia, proximal parts of both pubes and ischia, and proximal end
of one metatarsus (?). Associated material: 32 gastroliths.
Fig. 1- Type of Lourinhanosaurus antunesi. Scale bar 0.5 m.
Generic diagnosis
As shown below, the Peralta (Lourinhã) dinosaur can be assigned as
Allosauroidea. This specimen differs from all the others in the group because
all vertebral centra are longer than tall, the neural spines of the anterior caudal
vertebrae with a well-developed spike-like anterior process, the pubic blade
perforated by a large vertical ellipsoidal foramen, and the lesser trochanter is
well separated from the main body axis of the femur in lateral view. These
characters point out to a new genus to which the name Lourinhanosaurus
antunesi is given.
Type species, Lourinhanosaurus antunesi sp. nov.

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Species diagnosis
As for the type genus (see above).
Locality and age
The dinosaur was discovered at Peralta, about 75 Km NW of Lisbon
(Portugal), near Lourinhã (UTM coordinates: MD707443) - Upper Jurassic
(Upper Kimmeridgian/Tithonian, Sobral Unit) Manuppella (this volume). The
remnants are preserved in a gray, micaceous, fine sandstone.
DESCRIPTION
The holotype of Lourinhanosaurus antunesi corresponds to a partial
skeleton that was found lying on its right side. It comprises part of the axial
skeleton, the pelvis girdle and hindlimb remained in anatomical connection,
not showing any diagenetic etching or deformation.
Vertebrae (Fig. 2; table 1)
The neural arch is fused to the centra. All vertebral centra are longer
than tall which is a diagnostic feature, no showed in any other similar
theropod.
All cervical vertebrae found were in articulation. The cervical centra
present one pair of pleurocoels. The centra are strongly opisthocoelous. The
ventral surface presents a median keel. In articulation the cervical series
produce a curve concave in dorsal view. There are paired ribs. Diapophyses are
reduced.
The eleventh to fourteenth vertebrae are present in Lourinhanosaurus
considering that bears 14 dorsal vertebrae as in Sinraptor (Currie & Zhao,
1993), Allosaurus (Madsen, 1976), and Acrocanthosaurus (Stovall &
Langston, 1950). The centra of the posterior dorsal vertebrae are amphicoelous
and relatively longer than in Allosaurus fragilis. The centra are not
pleurocoelous and are narrowed just ventrally to the neural canal. The dorsal
vertebrae exhibit prominent neural spines and diapophyses.
Five sacral vertebrae are present. Sacral centra are not pleurocoelous.

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All the caudal vertebra centra are longer than high. The caudal centra
are neither hollow nor cavernous. The anterior caudal vertebrae are moderately
Fig. 2- Lourinhanosaurus antunesi gen. et sp. nov. ,A: third caudal vertebra; B: twelfth caudal
vertebra. ap: anterior process of the neural spine; ns: neural spine; po:
postzygapophysis; pr- prezygapophysis; tp: transverse process. Scale bar 5 cm.
Table 1 - Measurements of the vertebrae of ML370 Lourinhanosaurus antunesi
D
12? D
13? D
14? S
4 S
5 C
1 C
2 C
3 C
4 C
5 C
6 C
9 C
10 C
11 C12
Length 7.3 7.5 7.6 6.7 7.1 6.2 6.4 7.0 6.4 7.0 7.0 7.0 7.0 6.4 6.2
Height anterior end 6.1 5.6 6.5 5.5 5.7 5.9 6.2 6.2 5.8 5.7 5.7 -- -- -- 3.1
Width anterior end 6.6 7.8 -- 5.7 5.2 6.3 -- -- -- 4.4 4.1 3.9 3.8 3.7 3.5
Measurements in centimetres.
amphicoelous. In the ventral surface of anterior caudal centra there is a
longitudinal groove. A decreasing transverse process is present, at least, to the

MEMÓRIAS DA ACADEMIA DE CIÊNCIAS DE LISBOA
116
fifteenth caudal vertebra. Neural spines decrease in size caudalwards. The
transverse processes are long and

MEMÓRIAS DA ACADEMIA DE CIÊNCIAS DE LISBOA
117
placed near the middle of centrum. The neural spines of the anterior caudals
are thin and blade-like, with the anterior process spike-like present to, at least,
the fifteenth caudal vertebrae (see fig. 2). The anterior processes of the caudal
neural spines are more or less developed amongst allosauroids but not so much
as in Lourinhanosaurus. Except for the first caudal vertebrae the centra have
an anterior and posterior chevron facet and corresponding chevron. The
chevrons are deeper than long and not expanded distally. The chevron shafts
are slightly curve posteriorly. The chevron bases have paired anterior and
posterior processes. The haemal canal is close in the proximal end.
Pelvic Girdle (Fig. 3)
The three pelvic elements are not fused.
In lateral view the posterodorsal margin of ilia describes a slightly
ventral curve. The posterior end of ilium is pointed, and less truncated than in
Allosaurus. Almost whole of the blade border bears some roughness
corresponding to the pelvic muscle's insertions. The preacetabular portion is
expanded dorsoventrally; the postacetabular portion is significantly longer than
the preacetabular one. Pubic peduncle forms an angle of nearly 45º with the
main axis. The anterior end is expanded, and projects slightly beyond the pubic
peduncle. The posterior end is much more expanded beyond the ischial
peduncle. The medial blade is fused to the fifth sacral vertebra. The pubis
peduncle is longer than ischial peduncle but both are at the same level. The
pubic peduncle is twice as long anteroposteriorly as broad transversely.
The two pubes are in articulation. The pubis is strong and straight, with
postero-proximal expanded ischiac articulation. The pubic plate presents a
large, vertical ellipsoid pubic foramen or fenestra. The border of the
acetabulum is longer than in Allosaurus fragilis. Unfortunately the distal ends
of both pubes are not complete and the presence of the pubic boot, which is
very diagnostic of Allosauridae (Holtz, 1994), cannot be ascertained.
The ischium projects caudoventrally. The shaft is straight with a
minimum diameter of 2.2 centimetres. There is a slender “neck” at the pubic
peduncle. The obturator process is placed proximally with trapezoidal shape (it
is narrower near the ischial shaft than in the anterior edge). There is a notch
between the pubic and obturator processes.

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Fig. 3- Lourinhanosaurus antunesi gen. et sp. nov., reconstruction of the pelvic girdle. act:
acetabulum; il: ilium; ila: iliac articulation; ilp: iliac peduncle; is: ischium; isc: ischiac
articulation; isp: ischial peduncle; meb: medial blade; obf: obturator foramen or
fenestra; op: obturator process; pb: pubic blade; pu: pubis; pup: pubic peduncle of
ilium; Scale bar 5 cm. The ilium was reconstructed with the features of both ilia.
Femora, right tibia and fibula (Fig. 4 and 5)
The femoral head is medially directed. The fourth trochanter is long
and it is situated on the posterior surface of the femoral shaft where there is an
obvious muscle scar. Length of the fourth trochanter is, at least twice the femur
shaft diameter just below. The greater trochanter does not bear a cleft. It
presents an aliform lesser (or anterior) trochanter extended by a distally placed
lamella of bone, well separated from the main body of femur in lateral view
(see fig.4).

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The distal condyles are separated by a deep intercondylar groove. This
groove is more developed posteriorly than anteriorly. There is no crista
tibiofibularis (which is characteristic of Ceratosauria).
Fig. 4- Lourinhanosaurus antunesi gen. et sp.
nov., A- left femur in lateral view; B-
right femur in posterior view. h-
femoral head, lt- lesser trochanter, tr4-
fourth trochanter. Scale bar 5 cm.
Fig. 5- Lourinhanosaurus antunesi gen. et sp.
nov., right tibia and fibula in lateral
(A), and proximal (B) view. cf- crista
fibularis of tibia; cn- cnemial crest; fi-
fibula; Scale bar 5 cm.
The proximal articular surface of the tibia is flattened. The cnemial
crest is much less developed than in Allosaurus fragilis and does not arise out
of the lateral surface as in tyrannosaurids. The medial surface is curved
outwards and shows the muscular insertion. The lateral and posterior borders
are grooved to form a buttress for the proximal attachment of the fibula. The
tibial shaft has a sharp ridge (crista fibularis) placed laterally, for clasping the
fibula. The shaft is slightly curved anteriorly. The tibia seems shorter than the
femur.
The shaft of the fibula is slender and straight. It is expanded
proximally, being convex in lateral view and concave in medial view. The
distal end is lacking.

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Table 2 - Measurements of ML370 Lourinhanosaurus antunesi
Greatest length of Ilia* 46.0
Mid-diameter of femoral shaft 5.2
Length of femur * 54.0
Width of proximal end of fibula 7.9
Width of proximal end of tibia 12.7
Measurements in centimetres.
*- Estimated measurement.
Associated material
Gastroliths
Gastroliths have been described in Ornithopoda, Ceratopsia,
Prosauropoda, Sauropoda, Lacertilia, Crocodylia, extant birds and seals
(Stokes, 1987; Christiansen, 1996) but not in non-avian theropods until now.
The specimen had 32 gastroliths and the enveloping sediment preserved the
negative imprint of 3 additional gastroliths. The maximum observed gastrolith
length is 22 millimetres. Near the pebbles were three small bone fragments that
seemed to be food remains. The gastroliths have been found in the rib cage
below the eleventh dorsal vertebra. The high number, concentration and
relative size of the gastroliths suggest that they belong to this specimen, and
that they had not been swallowed when eating other dinosaur’s stomach.
DISCUSSION
This specimen shares the following theropod synapomorphies:
presacral vertebrae with pleurocoels; five sacral vertebrae; long preacetabular
process on the ilium; pronounced brevis fossa on caudal part of the ilium;
femur convex cranially; fibula closely appressed to the tibia and attached to a
tibial crest; and thin walled, hollow, long bones (see Gauthier, 1986). The
species is clearly a tetanuran and avetheropod (neotetanuran sensu Sereno et al.
1994, Allosauroidea + Coelurosauria) because the chevrons bases have paired
anterior and posterior processes, the iliac-ischial articulation is smaller than the
iliac-pubic articulation, ischial obturator notch is present, the femoral anterior
trochanter is blade-shaped, the iliac preacetabular fossa is present, and iliac

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pubic peduncle is twice as long anteroposteriorly as broad transversely (Sereno
et al., 1996).
No features cited in Sereno et al. (1994; 1996) are available to
diagnostic Lourinhanosaurus as Allosauroidea because only cranial characters
are used. However because of the mid cervical centra least than 20% broader
than tall, the elevation of the anterior face present in mid cervical centra, the
developed lesser trochanter, the presence of more than 15 caudal vertebrae
with transverse processes, the ischial obturator flange trapezoidal and the
reduced fibular fossa it is possible to classify this species as a non
Carcharodontosaurid nor Coelurosaurian Avetheropoda.
Holtz (1994) produced a single, most parsimonious cladogram of the
Theropoda. The data-matrix was criticised by Clark et al. (1994) and Charig &
Milner (1997). After that, Holtz (1995, 1996) and Sereno et al. (1996)
published new theropod relationship data. Using the data-matrix of Holtz
(1994) with the modifications introduced by Charig & Milner (1997) the
specimen from Lourinhã was placed as a basal Avetheropoda Paul, 1988. For
this study the Hennig86 programme was run. The cladogram has a consistency
index (C.I.) of 49%, a retention index (R.I.) of 70% and 236 steps length (see
fig. 6 A ).
The data-matrix of Sereno et al. (1996) was also run in Hennig86 with
Lourinhanosaurus characters showing a C.I. of 81%, R.I. of 84% and 80 steps
Fig. 6- Cladogram of the Theropoda including Lourinhanosaurus, A- after HOLTZ (1994)
with the modifications of CHARIG & MILNER (1997); B- SERENO et al. (1996).

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length. The Lourinhanosaurus was placed within Neotetanurae in a polytomy
among Sinraptoridae, Cryolophosaurus, Monolophosaurus, Allosaurus and
Carcharodontosauridae (see fig. 6B).
The following features justify including Lourinhanosaurus in the
Allosauroidea: transverse processes of middle caudal series placed near the
middle of the centrum, rather than posteriorly on the centrum; slender “neck”
at the pubic peduncle of the ischium; ischium with trapezoidal (in lateral view)
obturator process; small notch at the distal end of the ischial obturator process
and aliform lesser trochanter. This specimen differs from all other allosauroids
because all vertebrae are longer than tall, the neural spines of the anterior
caudals vertebrae with a well-developed spike-like anterior process, posterior
end of ilium is pointed not truncated, and the pubic blade perforated by a large
vertical ellipsoid fenestra.
Three genera have been reported to the Allosauridae: Allosaurus,
Neovenator and Saurophaganax (CHURE, 1995; HUTT et al., 1996). Recently,
SMITH (1998) reclassified Saurophaganax maximus into the genus Allosaurus
as the new species combination Allosaurus maximus. Acrocanthosaurus is
claimed to be Allosaurid by HOLTZ (1994) and Carcharodontosaurid by
SERENO (1996). The family Sinraptoridae (with genera Sinraptor,
Szechuanosaurus, and Yangchuanosaurus) and Carcharodontosauridae (with
genera Carcharodontosaurus, Bahariasaurus and Giganotosaurus) are related
to Allosauridae (CURRIE & ZHAO, 1993; HOLTZ, 1996; SERENO et al., 1996).
The taxonomic position of Chilantaisaurus, Cryolophosaurus and
Monolophosaurus remains uncertain.
Neovenator salerii is the only Allosaurid known in Europe so far (HUTT
et al., 1996). However Lourinhanosaurus is distinct from Neovenator by the
last dorsal vertebrae which is not pleurocoelous, and by the position of the
fourth trochanter on the posterior surface of the femoral shaft in
Lourinhanosaurus antunesi. Several features are similar to those found in
Allosaurus (ischial obturator process trapezoidal and aliform lesser trochanter,
MADSEN, 1976) but it differs by proportionally longer vertebrae, cnemial crest
of tibia less developed and by the others diagnostic characters cited above. The
chevron of Saurophaganax differs from those in Lourinhanosaurus by the
distal expansion. In Saurophaganax the shape of the ischial and pubic
obturator processes doesn’t resemble the Portuguese dinosaur.
The reduced cervical diapophyses are more similar to Sinraptor than to
Allosaurus fragilis. The neural spine is much less developed than in
Acrocanthosaurus atokensis. The obturator process is placed proximally and it
is trapezoidal-shaped as in Allosaurus fragilis (Madsen, 1976), or in
Acrocanthosaurus atokensis (Stovall & Langston, 1950).

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The pubic blade gap in Lourinhanosaurus could be the fusion of the
pubic fenestra with the pubic obturator foramen (the two gaps, pubic fenestra
and obturator foramen, are present in Coelophysis (Colbert, 1989) and
Syntarsus (Raath, 1980)), or an enlargement of the obturator foramen present
in non avetheropoda as Piatnitzkysaurus, Torvosaurus and Carnotaurus as well
in Monolophosaurus (Zhao & Currie, 1993), in the sinraptorid
Yangchuanosaurus shangyouensis (Dong et al., 1983). Considering the
taxonomic approach of these last two species to allosaurids (see Holtz, 1996),
the second hypothesis (enlargement of the obturator foramen) is more
probable. Sinraptor bears such similar character but with a small opening
which forms an obturator notch. This similarity suggests that
Lourinhanosaurus is more primitive than allosaurids and it is probably a
sinraptorid. In conclusion this is the only sinraptorid known in Europe so far.
Acknowledgements
Specials thanks are due to Prof. Miguel Telles Antunes for critical
review and support, as well as Prof. Philippe Taquet for scientific discussions
and help. Thanks are also to the followings Institutions and persons: GEAL-
Museu da Lourinhã (José Filipe, Isabel Mateus, , Horácio Mateus, and Vasco
Ribeiro); Departamento de Ciências da Terra, Faculdade de Ciências e
Tecnologia da Universidade Nova de Lisboa; Laboratoire de Paléontologie du
Muséum National d’Histoire Naturelle (Denis Serrette, Philippe Richir, and
France de Lapparent); Mr. Luís Mateus, who found this dinosaur specimen and
so kindly donated it to the GEAL- Museum of Lourinhã.
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