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A new Titanosaur genus (Dinosauria, Sauropoda) from the Late Cretaceous of southern France and its paleobiogeographic implications


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A new titanosaur, Atsinganosaurus velauciensis, gen. and sp. nov. is described from well-preserved remains from the new Upper Cretaceous locality of Velaux-La Bastide Neuve (Aix-en-Provence Basin, France). This taxon is mainly diagnosed by a combination of characters, which differentiates it without ambiguity from other European Late Cretaceous taxa (Lirainosaurus, Ampelosaurus and Magyarosaurus). Atsinganosaurus confirms the presence in western Europe during the latest Cretaceous of a third titanosaurian species, slender and less derived which allows us to better understand the evolutionary and paleobiogeographical history of this group during the Cretaceous.
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A new Titanosaur genus (Dinosauria, Sauropoda) from the Late Cretaceous
of southern France and its paleobiogeographic implications
Key-words. Sauropoda, Titanosauria, Upper Cretaceous, Southern France, Paleobiogeography.
Abstract.–Anewtitanosaur, Atsinganosaurus velauciensis, gen. and sp. nov. is described from well-preserved remains
from the new Upper Cretaceous locality of Velaux-La Bastide Neuve (Aix-en-Provence Basin, France). This taxon is
mainly diagnosed by a combination of characters, which differentiates it without ambiguity from other European Late
Cretaceous taxa (Lirainosaurus, Ampelosaurus and Magyarosaurus). Atsinganosaurus confirms the presence in western
Europe during the latest Cretaceous of a third titanosaurian species, slender and less derived which allows us to better
understand the evolutionary and paleobiogeographical history of this group during the Cretaceous.
Un nouveau genre de titanosaure (Dinosauria, Sauropoda) provenant du Crétacé supérieur
du Sud de la France et ses implications paléobiogéographiques
Mots-clés. Sauropoda, Titanosauria, Crétacé supérieur, Sud de la France, Paléobiogéographie.
Résumé. Un nouveau titanosaure, Atsinganosaurus velauciensis, gen. et sp. nov. est décrit à partir de restes bien pré
servés provenant d’une nouvelle localité fossilifère du Crétacé supérieur, nommée Velaux-La Bastide Neuve (bassin
d’Aix-en-Provence, France). Ce taxon est surtout défini par une combinaison de caractères qui le différencie sans am-
bigüité des autres taxons européens du Cretacé supérieur (Lirainosaurus, Ampelosaurus and Magyasaurus). Atsingano-
saurus confirme la présence en Europe de l’Ouest durant la fin du Crétacé d’une troisième espèce de titanosaurien, plus
élancée et moins dérivée qui nous permet de mieux comprendre l’histoire évolutive et paléobiogéographique de ce
groupe durant le Crétacé.
Titanosaur remains are abundant but sometimes fragmen
tary in the latest Cretaceous European archipelago [Tyson
and Funnell, 1987]. Only two valid titanosaur species are
currently recognized on the Iberian-Armorican island [Le
Lœuff, 1993; Wilson and Upchurch, 2003]: Ampelosaurus
atacis [Le Lœuff, 1995; 2005a] and Lirainosaurus astibiae
[Sanz et al., 1999].
The presence of a third species has been consistently as
sumed, but never demonstrated [Le Lœuff, 1995, 2005a;
Laurent et al., 2001; Allain and Pereda Suberbiola, 2003].
In the Transylvanian island (eastern Europe), another taxon
has been named, Magyarosaurus dacus, which clearly re
quires revision [Le Lœuff, 1993; Jianu and Weishampel,
1999; Wilson and Upchurch, 2003; Csiki et al., 2007].
The Late Cretaceous European dinosaur fauna raises
several paleobiogeographic problems, especially the inter-
relationships among the islands of the European archipel-
ago, and their possible connections with the Gondwanan
and Laurasian landmasses [Le Lœuff, 1991b; Rage, 2002;
Garcia et al., 2003; Allain and Pereda Suberbiola, 2003;
Gheerbrant and Rage, 2006; Pereda-Suberbiola, 2009]. Fur
thermore, it is still unclear whether the apparent absence of
sauropod skeletal remains from the Cenomanian to the
Campanian in Europe is due to an extinction event followed
by immigration from Africa [Le Lœuff, 1991b; 1993], or by
a bias in the fossil record [Wilson and Sereno, 1998;
Wilson, 2005]. In addition, titanosaur-like footprints have
been described in the Turonian-Santonian deposits of the
Periadriatic region [Mezga et al., 2006; Nicosia et al.,
Atsinganosaurus velauciensis, gen. et sp. nov. is de
scribed from well preserved titanosaur remains, previously
studied by Thouand [2004], from Velaux-La Bastide Neuve,
Bull. Soc. géol. Fr., 2010, t. 181, n
3, pp. 269-277
Bull. Soc. géol. Fr., 2009, n
1. IPHEP, UMR CNRS 6046, Université de Poitiers, 40 Avenue du Recteur Pineau, 86022 Poitiers cedex, France,,
2. Direction de l’Environnement, Conseil Général des Bouches-du-Rhône, Hôtel du Département, 52 Avenue St Just, 13256 Marseille cedex 20,
3. Géologie des Systèmes Carbonatés, EA 4234, Université de Provence Aix-Marseille 1, 3 place Victor Hugo, 13331 Marseille cedex 03,
4. 9, rue Buzelin 75018 Paris, France
Manuscrit déposé le 30 mars 2009; accepté après révision le 8 octobre 2009.
in the Aix-en-Provence Basin (Bouches-du-Rhône, southern
France). The bones were recovered in partial anatomical
connexion and close to some other vertebrate remains (tur
tle plates, crocodilian, theropod, nodosaurid and
rhabdodontid teeth), from a single horizon over an area of
(fig. 1). The fusion of the neural arches and cervical
and sacral ribs with the centra suggests that these sauropod
remains belong to mature individuals. Furthermore, the dif
ferent sizes of some bones could correspond to at least two
Saurischia Seeley, 1888
Sauropoda Marsh, 1878
Titanosauria Bonaparte and Coria, 1993
Atsinganosaurus velauciensis, gen. et sp. nov.
Holotype. VBN.93.01.a, b, c and d: four articulated poste
rior dorsal vertebrae, housed in the collections of the
Université de Poitiers, France.
Referred specimens. Université de Poitiers, France:
VBN.93.12a, b and c three cervical vertebrae; VBN.93.11
scapula; VBN.93.10 metatarsal; VBN.93.03-08 caudal ver
tebra. Musée de Paléontologie, Université de Provence,
France: VBN.02.03, 22 and 53 teeth; VBN.02.78 a and b
scapulo-coracoid; VBN.02.99 isolated dorsal vertebra;
VBN.02.82 sacrum; VBN.02.90 metacarpal; VBN.02.109
tibia; VBN.02.110 caudal vertebra. Musée archéologique de
Velaux, France: VBN.00.12 humerus; VBN.00.01-03 caudal
vertebrae. Muséum d’Histoire Naturelle d’Aix-en-Provence,
France: VBN.93.MHNA.99.21 tooth; VBN.93.MHNA.99.52
humerus; VBN.93.MHNA.99.32-34 caudal vertebrae.
Etymology. Generic name taken from the Byzantine Greek
atsinganoj, gypsy, in reference to the existence of Late
Cretaceous migrations between western and eastern Europe
revealed by these remains. The specific name is from the
Latin Velaucio, the name of the city –Velaux– where the
material was collected.
Type locality. La Bastide Neuve, Velaux; Aix-en-Provence
Basin, Bouches-du-Rhône, France. This locality was dis
covered by one of us (X. V.) in 1992.
Type horizon. Glauconitic grainstone limestone, Upper
Campanian (Begudian continental local stage) based on
charophytes (pers. comm., M. Feist) and dinosaur eggshells
according the local biostratigraphical context [Garcia and
Vianey-Liaud, 2001]. The presence of decapod claws asso
ciated with the bones strongly suggests a coastal environ
ment [Schweitzer et al., 2003].
Diagnosis. A slender titanosaurid which is characterized
by a combination of characters that differentiates it from
other European Late Cretaceous taxa (Lirainosaurus,
Ampelosaurus and Magyasaurus). Shares with
Malawisaurus the anterior cervical vertebrae without
marked pleurocoels, the posterior dorsal vertebrae with an
accessory lamina connecting the posterior centro
diapophyseal and postzygodiapophyseal laminae, and the
amphiplatyan to slightly opisthocoelous distal caudals.
Other features include teeth with cylindrical root and
slightly spatulate crown, with the labial face slightly convex
and the lingual face concave; absence of hyposphene-
hypantrum articulation in posterior dorsal vertebrae;
coracoid foramen equidistant from the dorsal and ventral
margins of the coracoid; sacrum with five opisthocoelous
fused vertebrae, with pleurocoels present only on the first
vertebra; strongly procoelous anterior caudal vertebrae;
dorsal edge of the scapular blade perpendicular to the proxi-
mal expansion of the scapula; distal end of the humerus
very slightly transversely expanded.
Bull. Soc. géol. Fr., 2010, n
270 GARCIA G. et al.
FIG. 1. Simplified geological map of the Aix-en-Provence Basin and stratigraphical column of the Velaux site. Arrow indicates the location of the site.
IG.1.–Carte géologique simplifiée du bassin d’Aix-en-Provence et colonne stratigraphique du site de Velaux. La flèche indique la localisation du site.
Anatomical abbreviations. a, acromion; acdl, anterior
centrodiapophyseal lamina; acpl, anterior centropara
pophyseal lamina; cc, cnemial crest; cpol, centropostzy
gapophyseal lamina; di, diapophysis; dpc, deltopectoral
crest; fc, coracoid foramen; ns, neural spine; pc, pleurocoel;
pcdl, posterior centrodiapophyseal lamina; pcpl, posterior
centroparapophyseal lamina; podl, postzygodiapophyseal
lamina; poz, postzygapophysis; prz, prezygapophysis; spol,
spinopostzygapophyseal lamina; sprl, spinoprezy
gapophyseal lamina; L, lamina connecting the pcdl and the
podl; S1, first sacral vertebra; S5, fifth sacral vertebra. Ab
breviations for vertebral laminae follow Wilson [1999].
Teeth (figs. 2A and B)
Some isolated teeth of different sizes were discovered at the
site. Most of the teeth are straight and massive and present a
cylindrical root, whereas the apical part of the crown is
labiolingually flattened. There are apical wear facets on
some specimens. The convex labial face does not show
grooves or ridges, and the lingual face is slightly concave,
contrary to Lirainosaurus [Sanz et al., 1999]. The teeth do
not display the diagnostic mesial and distal expansions of
Ampelosaurus [Le Lœuff, 1995].
Cervical vertebrae (fig. 3A)
Three poorly preserved adjacent anterior cervical vertebrae
have been found in quasi-connexion. They have elongate (23
to 27.5 cm long) and opisthocoelous centra, without marked
pleurocoels on their lateral sides, as in Malawisaurus [Jacobs
et al., 1993; Gomani, 2005] and contrary to Ampelosaurus
atacis which presents deep pleurocoel cavities [Le Loeuff,
2005]. The ventral face of the centrum is concave
transversally to slightly flat in its posterior part. The neural
spine of the specimen VBN.93.12 c is high and not bifid, and
the postzygapophyses do not project beyond the posterior ar
ticulation. Prezygapophyses are anteriorly elongate. Deep
prespinal and postspinal fossae are delimited by thick
spinoprezygapophyseal and spinopostzygapophyseal laminae,
respectively. The angle between the posterior centrodia
pophyseal and postzygodiapophyseal laminae is less than
Dorsal vertebrae (fig. 3B)
Four articulated posterior dorsal vertebrae (holotype) and
one isolated (probably more anterior) dorsal vertebra are
preserved. Their centra are strongly opisthocoelous, not
very short, and with well-developed pleurocoels. The ven
tral face, which is concave laterally, lacks a sagittal ridge.
All elements have characteristic cancellous bone in the
centra and the neural arch. The features of laminae are ob
served especially on the third vertebra of the holotype,
which is the most complete preserved (fig. 3B). There is no
hyposphene-hypantrum articulation. The neural arch is an
teriorly directed. In lateral view, the slender acpl and pcpl
are nearly fused; the latter is restricted to the anterior part
of the thick neural arch. The acdl and pcdl are fused and
form a ventrally widened “infradiapophyseal lamina”, with
out deep fossa on both sides. An accessory lamina, well de
fined on the third vertebra connects the pcdl to the podl
(L in fig. 3B), not observed in Ampelosaurus and
Lirainosaurus. The diapophysis is well expanded and
slightly slanted. The poorly preserved neural spines are di
rected posteriorly. They are reinforced by spinodiapo
physeal, spinopostzygapophyseal and prespinal laminae,
but postspinal laminae are not distinguishable.
Sacrum (figs. 4A and B)
The nearly complete sacrum is composed of five fully co-os-
sified vertebrae. Prezygapophyses and postzygapophyses of
adjacent centra are fused, and neural spines are linked by the
fusion of adjacent prespinal and postspinal laminae. Al-
though it is not preserved, the presence of an unfused sixth
sacral cannot be ruled out. The anterior articular surface (of
the first centrum) is strongly convex; the posterior articular
facet (of the fifth) is concave. The third to fifth centra are
strongly transversely narrow. Fused ossified tendons are
present above the neural spines, as in Malawisaurus,
Epachthosaurus [Martínez et al., 2004], and a titanosaurid
from Brazil [Baurutitan britoi from the Peirópolis locality,
Campos and Kellner, 1999; Kellner et al., 2005]. Unlike
many titanosaurs, Atsinganosaurus, Epachthosaurus and
Baurutitan display well developed pleurocoels in the first sa
cral, but pleurocoels are absent in the remaining sacrals.
These shared features may reflect close affinities between
these taxa, as proposed by Martínez et al. [2004]. As in the
Peirópolis sacrum, a pneumatic opening is present ventral to
each parapophysis of the first sacral vertebra. The prespinal
lamina is not bifurcate at its base, unlike Epachthosaurus.
The opening between the infradiapophyseal lamina and the
tuberculum and capitulum of the ribs is absent in the two last
sacrals, whereas it is present in Malawisaurus.
Caudal vertebrae (figs. 4C-G)
Six (VBN.93.03-08) out of about ten vertebrae, with identi
cal morphology and decreasing sizes were recovered close
to each other and could belong to a single individual. The
centra of all vertebrae are dorsoventrally or laterally com
pressed due in part to post-mortem deformation. The
dorsoventral flattening ratio (centrum l: width /h: height)
varies between 0.8 and 2.3 cm.
All neural arches are situated on the anterior half of the
centrum. The anterior caudal vertebrae, which are smaller
in size (diameters of the centra around 4.5 cm) than those
Bull. Soc. géol. Fr., 2010, n
FIG. 2. Teeth of Atsinganosaurus velauciensis. VBN.93.MHNA.99.21:
tooth in mesial (A) and labial (B) views.
IG.2.Dents d’Atsinganosaurus velauciensis. VBN.93.MHNA.99.21 :
dent en vues mésiale (A) et labiale (B).
generally observed in titanosaurs from southern Europe
(around 6 cm), are procoelous, but the posterior condyle is
not as prominent as in many titanosaurs (fig. 4C). As in
Malawisaurus [Gomani et al., 1999] and Lirainosaurus, the
apex is dorsally located. On the ventral face, ridges link the
anterior and posterior hemapophyseal facets and border a
wide sagittal depression. Only one middle caudal vertebra
has been discovered. Its longer centrum is strongly amphi
coelous with a sagittal depression in ventral view. Its neural
arch is not preserved. Distal caudals, except for two
proceolous types, are generally amphiplatyan (fig. 4D-F) to
slightly opisthocoelous, unlike Lirainosaurus astibiae and
Bull. Soc. géol. Fr., 2010, n
272 GARCIA G. et al.
FIG. 3. Presacrals. A, VBN.93.12a, b and c: cervical
vertebrae in left lateral view. B, VBN.93.01a, b, c, and
d : four articulated dorsal vertebrae in right lateral
view (holotype of Atsinganosaurus velauciensis). C,
detail and schematic drawing of the 2
dorsal vertebra
showing the major structures.
IG.3.Pré-sacrées. A, VBN.93.12a, b et c : vertè
bres cervicales en vue latérale gauche. B, VBN.93.01a,
b, c, et d : quatre vertèbres dorsales articulées en vue
latérale droite (holotype d’Atsinganosaurus velaucien
sis). C, détail et dessin schématique de la 2
dorsale montrant les principales structures.
Bull. Soc. géol. Fr., 2010, n
Ampelosaurus atacis, in which the distal caudals are de-
scribed as procoelous [Sanz et al., 1999; Le Loeuff, 2005a].
The prezygapophyses protrude anteriorly; the postzy-
gapophyses extend over the posterior half of the centra
(fig. 4F). The centra and prezygapophyses of the six caudal
vertebrae become progressively longer and lower.
Appendicular skeleton (figs. 5 and 6)
A left scapulo-coracoid and a poorly preserved larger scap-
ula have been discovered. There is no medial ridge on the
ventral edge of the scapular blade. Its long axis is perpen
dicular to the proximal expansion of the scapula and to the
FIG. 4. Sacrum and caudal vertebrae. A and B,
VBN.02.82: sacrum in right lateral and ventral
views. C, VBN.02.110: middle caudal vertebra in left
lateral view. D-G, VBN.93.MHNA.99.33: posterior
caudal vertebra in dorsal (D), posterior (E), lateral
(C) and cranial (G) views with their schematic outli
IG.4.Sacrum et vertèbres sacrées. A et B,
VBN.02.82 : sacrum en vues latérale droite et ven
trale. C, VBN.02.110 : vertèbre caudale moyenne en
vue latérale gauche. D-G, VBN.93.MHNA.99.33 :
vertèbre caudale postérieure en vues dorsale (D),
postérieure (E), latérale (C) and craniale (G) avec
leurs contours schématisés.
FIG. 5. Scapulocoracoid and limbs. A and B, VBN.02.78: left scapulo-coracoid in medial and lateral views with its schematic drawing. C, VBN.00.12:
right humerus in anterior view. D, VBN.02.109: left tibia in postero-lateral view.
IG.5.Scapulo-coracoide et membres. A et B, VBN.02.78 : scapulo-coracoide gauche en vues médiale et latérale avec son dessin schématique.
C, VBN.00.12 : humérus droit en vue antérieure. D, VBN.02.109 : tibia gauche en vue postero-latérale.
articulation with the quadrangular coracoid. The cranial
margin of the coracoid is well rounded. The coracoid fora
men is close to the suture between the scapula and the
coracoid, and equidistant from the ventral and dorsal mar-
gins of the bone (fig. 5A and B).
The limb bones are all very slender, particularly the hu-
merus. On the two right humeri (the well-preserved speci-
men is VBN.00.12), the prominent deltopectoral crest does
not extend to the distal half of the bone (fig. 5C), unlike in
saltasaurids [Salgado et al., 2005]. The distal end of the hu-
merus (medio-lateral width is 10 cm for the preserved spec-
imen) is much less expanded than in most other titanosaurs.
On the other hand, the proximal end is very large
(medio-lateral width of 20 cm) giving a concave shape to
the medial border. In caudal view, we observe a
supracondylar depression distally edged by two ridges. The
distal condyles are not well developed anteriorly.
On the left tibia (length: 53 cm), the cnemial crest is
strong and orientated slightly anterolaterally (fig. 5D), and
its distal part is equally expanded transversely and
anteroposteriorly, as in Lirainosaurus. Only two isolated
metapodials without any conspicuous feature (one metacar
pal fig. 6A and one metatarsal fig. 6B) have been found.
Atsinganosaurus is assigned to Titanosauria [sensu Sereno,
1998: a stem-based taxon defined as all Somphospondyly
more closely related to Saltasaurus than to Euhelopus]on
the basis of the following characters: presence of
centroparapophyseal and ventrally widened or slightly
forked centrodiapophyseal laminae on posterior dorsal ver
tebrae; acuminate pleurocoels in dorsal vertebrae [Salgado
et al., 1997; Calvo and González-Riga, 2003; Bonaparte
et al., 2006]. Atsinganosaurus differs from the basal-most
titanosaurs Andesaurus, Phuwiangosaurus, and Ligabuesaurus
in having undivided cervical pleurocoels, procoelous ante
rior caudals with a ventral longitudinal hollow, and in lack
ing a hyposphene-hypantrum articulations in dorsals.
Atsinganosaurus exhibits several features which are
here interpreted as plesiomorphic conditions retained in
basal lithostrotians [Upchurch et al., 2004]: perpendicular
angle between the scapular blade and the coracoid articula
tion; amphiplatyan distal caudals [Wilson, 2002];
parapophysis not lying directly below the diapophysis in
posterior dorsals; absence of prominent process at the junc
tion of the proximal and lateral edges of the humerus
[Upchurch et al., 2004]; dorsal margin of the coracoid lying
below the dorsal margin of the proximal end of the scapula
[Upchurch, 1998]. Although the number of fused sacral ver
tebrae is subject to both phylogenetic and ontogenetic varia
tion [Upchurch, 1998] the sacrum of Atsinganosaurus does
not exhibit juvenile features. Its five fused vertebrae are
thus more reminiscent of the condition in plesiomorphic
Camarasauromorpha [Tidwell et al., 2005] than that of the
most derived titanosaurians Saltasaurus and Neuquen
saurus, in which the sacrum consists of six fused sacrals
with the possible incorporation of a seventh [Powell, 2003;
Salgado et al., 2005]. Apart from Atsinganosaurus, the
combination of the absence of hyposphene-hypantrum artic
ulations in dorsals and procoely restricted to the anterior
part of the tail (two features which have been consistently
considered to be of phylogenetic interest [Jacobs et al.,
1993; Bonaparte, 1996; Upchurch, 1998; Wilson and
Sereno, 1998; Wilson, 2002; Calvo and González Riga,
2003; O’Leary et al., 2004; Salgado and Coria, 2005]), is
found only in the basal lithostrotian Malawisaurus. This ge-
nus also shares with the titanosaur from Velaux the reduc-
tion of cervical pleurocoels, fused ossified ligaments in the
sacrum, and the dorsally located apex of the posterior
condyle in caudals. All these features suggest the basal po-
sition of Atsinganosaurus within Lithostrotia, which has
been confirmed by preliminary works [Tortosa, 2006; Gar-
cia et al., in prep].
The tooth morphology, the presence of an accessory
lamina between the pcdl and the podl in posterior dorsals,
amphiplatyan distal caudals, the coracoid foramen far from
the dorsal margin of the bone and the weak distal expansion
of the humerus are features that distinguish Atsingano
saurus from Lirainosaurus astibiae and Ampelosaurus
atacis [Le Loeuff, 1995, 2005a; Sanz et al., 1999]. More
over, the new genus differs from Ampelosaurus in the ab
sence of deep pleurocoels in anterior cervicals; poor
development of the pcpl, fusion of the acdl and pcdl, and
absence of an accessory lamina between the postzyga
pophysis and pcdl in posterior dorsals and concave poste
rior articulation of the fifth sacral. Unlike Lirainosaurus,
the distal caudals of Atsinganosaurus do not have a poste
rior condyle with a sagittal groove, and their
spinopostzygapophyseal structure is posteriorly projected;
also, in the new genus, the dorsal centra lack an axial keel,
the podl is well developed, and the angle between the axis
of the scapular blade and the proximal expansion is perpen
dicular. Magyarosaurus differs from Atsinganosaurus in
having ridges on its scapular blade, by the position of its
coracoid foramen, and by the orientation of its tibial distal
end. However, it shares with Atsinganosaurus non-procoelous
distal caudals [Nopcsa, 1915; Le Lœuff, 1992].
Several southern European lithostrotian remains, which
are still generically indeterminate [Allain and Pereda
Suberbiola, 2003] might be phylogenetically close to the
new titanosaur from Velaux. For example, the teeth from the
Bull. Soc. géol. Fr., 2010, n
274 GARCIA G. et al.
FIG. 6. Hindlimbs. A, VBN.02.90 : metacarpal in lateral view. B,
VBN.93.10 : metatarsal in lateral view.
IG.6.Membres. A, VBN.02.90 : métacarpien en vue latérale. B,
VBN.93.10 : métatarsien en vue latérale.
Campanian of Saint-Estève-Janson (pers. obs. Garcia) and
from the late Maastrichtian of Cassagnau [Laurent et al.,
2002] closely resemble those of Atsinganosaurus. The
amphiplatyan to slightly amphicoelous distal caudals,
which have been described from the Maastrichtian of
Cubilla [Pereda Suberbiola and Ruiz-Omeñaca, 2001] and
in the Late Campanian of Tercis-les-Bains [Laurent et al.,
2001] are reminiscent of those of Atsinganosaurus. Isolated
strongly amphicoelous middle caudals have been identified
in the Maastrichtian of Mas d’Azil [Le Lœuff, 1991a] and
in the Albian Macrurosaurus [Le Lœuff, 1992, 1993].
The description of Atsinganosaurus confirms that at least
three titanosaurian species were present in western Europe
during the latest Cretaceous. Such a large diversity of her
bivorous animals on an island is not inconceivable, because
the co-occurrence of several sauropod taxa in the same area
is well documented [Dodson et al., 1980; Sander, 2000; Day
et al., 2002; Gomani, 2005], even in island ecosystems
[Curry Rogers and Forster, 2001; Dalla Vecchia, 2005; Wil
son, 2005]. Moreover, the Iberian-Armorican island was
quite large, up to three times larger than modern Madagas
car, and it was occasionally connected to other central Euro
pean landmasses [Le Lœuff, 2005b]. The differences in
body size and tooth morphology among the three western
European titanosaurs may reflect their occupation of dis-
tinct ecological niches [Upchurch et al., 2004; Salgado and
Coria, 2005].
Atsinganosaurus confirms without ambiguity that ti-
tanosaurs with procoely restricted to the anterior part of the
tail were still present, at least in Europe, during the latest
Cretaceous, whereas this condition was previously thought
restricted to Early and mid-Cretaceous titanosaurs [Jacobs
et al., 1993; O’Leary et al., 2004]. In the European Late
Cretaceous, this plesiomorphic feature is shared only with
Magyarosaurus. Therefore, we consider this taxon closer to
Atsinganosaurus than to the more derived Lirainosaurus
and Ampelosaurus. The persistence of basal lithostrotians in
the Ibero-Armorican and Transylvanian islands, while more
derived titanosaurs were restricted to western Europe, im
plies the existence of filters or sweepstakes dispersal routes
between various European regions during the Late Creta
ceous. Although a land bridge with topographic barriers has
been assumed [Jianu and Boekschoten, 1999], these routes
were more likely discontinuous, comprised of temporarily
emergent areas, which have been recently proved more nu
merous than previously thought [Buffetaut et al., 2002;
Rage, 2002]. Moreover, these links probably allowed
sauropod migrations [Weishampel et al., 2003; Dalla
Vecchia, 2005].
The new titanosaurian from Velaux appears close to
Malawisaurus from the Early Cretaceous of southern Af
rica, and could demonstrate the survival of basal
lithostrotians into the latest Cretaceous. The proposed affin
ities of the new French form suggest the existence of a dis
persal route from Africa to Europe, possibly via the
Mediterranean Tethyan Sill, which was an important dis
persal route during the Cretaceous [Gheerbrant and Rage,
2006; Pereda-Suberbiola, 2009] especially for sauropods
such as rebbachisaurids [Pereda Suberbiola et al., 2003;
Dalla Vecchia, 2005]. The Late Cretaceous survival of
primitive titanosaurians in Europe argues in favor of an
early migration event, likely by the Cenomanian, followed
by insular endemism until the Late Cretaceous. Based on
the presence of advanced characters in taxa such as
Ampelosaurus and Lirainosaurus [Curry Rogers, 2005], we
suggest that more derived titanosaurs probably also oc
curred in the latest Cretaceous of Europe, along with relict
forms, as proposed for European ankylosaurs and
hadrosaurids [Casanovas et al., 1999; Garcia and Pereda
Suberbiola, 2003; Õsi, 2005]. In that case, their occurrence
would indicate a second dispersal event. Nevertheless, the
phylogenetic affinities of European titanosaurs remain un-
clear. Although Lirainosaurus [Sanz et al., 1999; Wilson,
2002; Calvo and González-Riga, 2003; Powell, 2003;
Upchurch et al., 2004] and Ampelosaurus [Wilson, 2002;
Upchurch et al., 2004] are often proposed to be advanced
lithostrotians, the only published cladistic analysis in which
they are both included recovers them as basal titanosaurs
[Curry Rogers, 2005]. The revision of Magyarosaurus,
the description of new material of Ampelosaurus, and the
inclusion of these taxa in a cladistic analysis with
Atsinganosaurus is needed in order to assess these
paleobiogeographic hypotheses.
Acknowledgements. We gratefully acknowledge the Velaux Municipality
(Mayor J.-P. Maggi, Mr. and Mme Nicolini, V. Attale and local volunteer
excavators, especially E. Turini and S. Rafay), the personnel of the envi
ronment department from CG 13 (including M. Fronteri), our colleagues
M. Feist, J.-F. Babinot, J. Hennuy and G. Tronchetti for their assistance
and S. Riffaut for the photographs. We also thank D. Roggero who gave us
the material collected in 1993, J. Le Loeuff (Musée des dinosaures d’Espé
raza, Aude), J. Philip and L. Villier (Université de Marseille, Musée de Pa
léontologie), the Muséum d’Histoire Naturelle d’Aix-en-Provence and the
Musée archéologique de Velaux (Bouches-du-Rhône). This work was sup
ported by funds from the Conseil Général des Bouches-du-Rhône and by
the UMR-CNRS 6046 (University of Poitiers).
SGF associate editor: D. Grosheny.
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... The Upper Cretaceous fossil record from Europe is rich in sauropod occurrences, spanning from the Cenomanian to the Maastrichtian, which have been remarkably increased in the last three decades with new discoveries and the establishment of new taxa (e.g. Le Loeuff 1993Loeuff , 2005Sanz et al. 1999;Csiki et al. 2010a;Díez Díaz et al. 2016García et al. 2010;Ortega et al. 2015;Ősi et al. 2017;Mocho et al. 2019a;Vila et al. 2022). In addition, the redescription of some previously established taxa (e.g. ...
... Csiki-Sava et al. 2015, and references herein;Ősi et al. 2017;Nikolov et al. 2018). The Campanian-Maastrichtian sauropod fauna of the continent is diverse and characterized by eight valid titanosaurian taxa probably belonging to Lithostrotia (Nopcsa 1915;Huene 1932;Le Loeuff 1995;Sanz et al. 1999;Csiki et al. 2010a;García et al. 2010;Díez Díaz et al. 2016Vila et al. 2022). However, the phylogenetic relationships of these European forms within Lithostrotia are still poorly understood. ...
... In recent years, the number of studies about the European titanosaurs has increased, mainly based on the titanosaurian remains found in Southeastern Europe (France and Spain) (e.g. García et al. 2010;Díez Díaz et al. 2016;Vila et al. 2022). By contrast, our understanding of the titanosaurs from the Upper Cretaceous of Romania is still poor, being necessary to provide a full systematic reassessment of the historical material, as well as the description of new specimens. ...
The Upper Cretaceous sauropod record from Romania is abundant, but mainly composed by isolated and poorly preserved specimens. They were found in Maastrichtian units cropping out in both the western-southwestern margin of the Transylvanian Basin, and in the Hațeg and Rusca Montană basins. Two small-sized taxa, Magyorasaurus dacus and Paludititan nalatzensis, were considered as valid; however, the presence of other forms is not ruled out. An important sample of sauropod caudal vertebrae is described in detailed to provide new data about the diversity of the sauropods of the Hațeg Island during the Maastrichtian. Distinct morphologies are recognized for the caudal vertebrae, including four morphologies for the anterior-most elements; at least three for the anterior and middle ones, and at least two for the posterior caudals, including sauropods with opisthocoelous condition. The possibility of four different tail morphologies is suggested, which might belong to four different taxa, including Paludititan nalatzensis, Magyarosaurus dacus (the material previously referred to it seems to be composed by more than one taxon) and two other species. A new character for morphological datasets describing the position of the intraprezygapophyseal lamina in the middle and posterior caudals is proposed, which may be characteristic of some European taxa.
... (2) the Mannion et al. (2013) dataset family (e.g., González Riga et al., 2018Poropat et al., 2021); (3) the Carballido and Sander (2014) dataset family (e.g., Cerda et al., 2021;Gallina et al., 2021); and (4) the Gorscak and O'Connor (2016) dataset family (e.g., Sallam et al., 2018;Gorscak and O'Connor, 2019;Vila et al., 2022;current study). For example, European and/or African representatives of Titanosauria tend to be underrepresented or even nearly absent in many phylogenetic analyses despite the fact that these continents have produced numerous new titanosaurian taxa in recent decades (e.g., Smith et al., 2001;Gomani, 2005;Csiki et al., 2010;Garcia et al., 2010;Curry Rogers and Wilson, 2014;Gorscak et al., 2014Gorscak et al., , 2017Díez Díaz et al., 2016, 2021Sallam et al., 2018;Gorscak and O'Connor, 2019;Vila et al., 2022). A similar phenomenon has occurred with the surge in Australian titanosauriform discoveries within the past ∼15 years (e.g., Australotitan cooperensis, Diamantinasaurus, Savannasaurus, and Wintonotitan wattsi), which have typically been analyzed using datasets within the Mannion et al. (2013) family (e.g., Hocknull et al., 2021;Poropat et al., 2021). ...
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Dinosaur fossils from the latest Cretaceous (Campanian–Maastrichtian) of Africa and the Arabian Peninsula are rare. Most discoveries to date have consisted of limited fossils that have precluded detailed phylogenetic and paleobiogeographic interpretations. Fortunately, recent discoveries such as the informative Egyptian titanosaurian sauropod dinosaur Mansourasaurus shahinae are beginning to address these long-standing issues. Here we describe an associated partial postcranial skeleton of a new titanosaurian taxon from the Upper Cretaceous (Campanian) Quseir Formation of the Kharga Oasis, Western Desert of Egypt. Consisting of five dorsal vertebrae and 12 appendicular elements, Igai semkhu gen. et sp. nov. constitutes one of the most informative dinosaurs yet recovered from the latest Cretaceous of Afro-Arabia. The relatively gracile limb bones and differences in the coracoid and metatarsal I preclude referral of the new specimen to Mansourasaurus. Both model-based Bayesian tip-dating and parsimony-based phylogenetic analyses support the affinities of Igai semkhu with other Late Cretaceous Afro-Eurasian titanosaurs (e.g., Mansourasaurus, Lirainosaurus astibiae, Opisthocoelicaudia skarzynskii), a conclusion supported by posterior dorsal vertebrae that lack a postzygodiapophyseal lamina, for example. Igai semkhu strengthens the hypothesis that northern Africa and Eurasia shared closely related terrestrial tetrapod faunas at the end of the Cretaceous and further differentiates this fauna from penecontemporaneous assemblages elsewhere in Africa, such as the Galula Formation in Tanzania, that exhibit more traditional Gondwanan assemblages. At present, the specific paleobiogeographic signal appears to vary between different dinosaur groups, suggesting that Afro-Arabian Cretaceous biotas may have experienced evolutionary and paleobiogeographic histories that were more complex than previously appreciated.
... Open Sci. 10: 221618 Ibiricu et al. [279]Machado et al.[164] Garcia et al.[280]; Díez Díaz et al.[13] R. Soc. Open Sci. ...
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Titanosaurian sauropod dinosaurs were diverse and abundant throughout the Cretaceous, with a global distribution. However, few titanosaurian taxa are represented by multiple skeletons, let alone skulls. Diamantinasaurus matildae, from the lower Upper Cretaceous Winton Formation of Queensland, Australia, was heretofore represented by three specimens, including one that preserves a braincase and several other cranial elements. Herein, we describe a fourth specimen of Diamantinasaurus matildae that preserves a more complete skull—including numerous cranial elements not previously known for this taxon—as well as a partial postcranial skeleton. The skull of Diamantinasaurus matildae shows many similarities to that of the coeval Sarmientosaurus musacchioi from Argentina (e.g. quadratojugal with posterior tongue-like process; braincase with more than one ossified exit for cranial nerve V; compressed-cone–chisel-like teeth), providing further support for the inclusion of both taxa within the clade Diamantinasauria. The replacement teeth within the premaxilla of the new specimen are morphologically congruent with teeth previously attributed to Diamantinasaurus matildae, and Diamantinasauria more broadly, corroborating those referrals. Plesiomorphic characters of the new specimen include a sacrum comprising five vertebrae (also newly demonstrated in the holotype of Diamantinasaurus matildae), rather than the six or more that typify other titanosaurs. However, we demonstrate that there have been a number of independent acquisitions of a six-vertebrae sacrum among Somphospondyli and/or that there have been numerous reversals to a five-vertebrae sacrum, suggesting that sacral count is relatively plastic. Other newly identified plesiomorphic features include: the overall skull shape, which is more similar to brachiosaurids than ‘derived' titanosaurs; anterior caudal centra that are amphicoelous, rather than procoelous; and a pedal phalangeal formula estimated as 2-2-3-2-0. These features are consistent with either an early-branching position within Titanosauria, or a position just outside the titanosaurian radiation, for Diamantinasauria, as indicated by alternative character weighting approaches applied in our phylogenetic analyses, and help to shed light on the early assembly of titanosaurian anatomy that has until now been obscured by a poor fossil record.
... Díez Díaz et al. 2016). In the Ibero-Armorican domain (present-day areas of the Iberian Peninsula and France), the titanosaurian fossil record comprises bones, teeth, tracks, eggs and even skin impressions, with up to five species described: Ampelosaurus atacis (Le Loeuff, 1995), Atsinganosaurus velauciensis (Garcia et al., 2010), Lirainosaurus astibiae (Sanz et al., 1999), Lohuecotitan pandafilandi (Díez Díaz et al., 2016), and Garrigatitan meridionalis (Díez-Díaz et al., 2021). Nevertheless, several authors estimated a higher diversity in the region (e.g. ...
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Titanosauria is the only sauropod taxa present in the Late Cretaceous of Europe. One of the most diverse localities in the Southern Pyrenees is the lower Maastrichtian site of Els Nerets (Tremp, Catalonia, Spain). The present work reviews the new sauropod material discovered in recent years together with the classical specimens from the site in order to establish the Minimum Number of Individuals (MNI) through the relationships between the appendicular elements. The results show the presence of three adult individuals (based on histological observations) of different sizes, suggesting the occurrence of two distinct, indeterminate species. Although they are similar, morphological comparisons with other Ibero-Armorican titanosaurs suggests that the individuals from Els Nerets cannot be confidently assigned to any of them given the fragmentary nature of the material and the lack of recognisable autapomorphies. However, this work improves on our understanding of the Els Nerets titanosaur diversity.
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A fragmentary centrum of a dorsal vertebra and a manual ungual phalanx of eusauropod dinosaurs from the Upper Barremian-Lower Aptian of Balve in western Germany are described. The dorsal centrum shares potential synapomorphies with the enigmatic genus Ornithopsis and can probably be referred to a titanosauriform. The Balve record is of significance as it represents the only known direct evidence of this clade from an upland environment in Europe, which is assumed to have reached palaeoelevations of several hundred meters above sea level. Taphonomic signatures at some of the material from this site indicate prolonged surface exposure plus various stages of wear caused by water transport. The remains were finally deposited in sediment traps within a karst cave or fissure system. The fossil record suggests a retreat or shift of sauropod habitat range from coastal plains and lowlands to hinter-and uplands during the Early Cretaceous in northwest Europe. It is hypothesized that this was linked to the faunal replacement of low-and mid-level browsing sauropods (e.g. diplodocoideans) by iguanodontian ornithopods near the Jurassic-Cretaceous boundary in this region. High-browsing titanosauriform sauropods had small overlap in the trophic niche with the ornithopods, facilitating a sympatric coexistence of both groups. However, their range was dependent on high-canopy vegetation that was probably more abundant in uplands. The previously observed "mid-Cretaceous bottleneck" in sauropod diversity may therefore be a result of partial extinction and/or a sampling bias, as the preservation of upland faunas are very exceptional. However, it is be cautioned to interpolate these regional observations to global sauropod diversity patterns.
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Résumé : Les dinosauriens de la Réserve Naturelle de Sainte-Victoire sont surtout représentés par la découverte de leurs oeufs depuis les années 1940. Cependant, des ossements furent également mis au jour à la fin des années 1980, quelques années avant le classement du territoire en réserve naturelle nationale. Cet article fait le point sur l’histoire des découvertes d’ossements fossilisés sur le territoire avec, notamment, Variraptor mechinorum. Puis, sont présentés les résultats des premières fouilles méthodiques menées par le Muséum d’Histoire Naturelle d’Aix-en-Provence dans les années 2010. L’ensemble des restes osseux collectés est attribué à trois taxons, scientifiquement publiés ou non : Variraptor, Rhabdodon et un titanosaurien indéterminé. Ces dinosauriens sont décrits de manière détaillée et sont comparés aux autres représentants de leurs familles respectives. Ces descriptions anatomiques sont accompagnées d’anecdotes et présentent les implications scientifiques liées à leur découverte vis-à-vis des autres spécimens connus en Provence ou des autres régions fossilifères du Sud de l’Europe. Abstract: Dinosaurs from the Réserve Naturelle de Sainte-Victoire are mainly represented by the discoveries of their eggs since the 1940s. However, fossilized bones were also brougth to light at the end of the 1980s, a few years before the designation of the territory as a national natural reserve. This article reviews the history of the discoveries of fossil bones on the territory of the reserve, including those of Variraptor mechinorum. Then, the results of the first systematic excavations conducted by the Muséum d’Histoire Naturelle d’Aix-en-Provence in the 2010s are presented. The whole of the collected remains are attributed to three taxa, scientifically published or not: Variraptor, Rhabdodon and an indeterminate titanosaur. These dinosaurs are described in detail and are compared with other representatives of their respective families. These anatomical descriptions are complemented by anecdotes and present the scientific implications linked to their discovery, with respect to the other specimens known in Provence or the other fossiliferous regions of southern Europe.
After the extinction of rebbachisaurids during the Cenomanian–Turonian interval, titanosaurs were the only group of sauropods to face the K–Pg event. This same global pattern also holds for the end-Cretaceous (Campanian–Maastrichtian) titanosaur record in South America, where their remains can be found from southern Argentina to Ecuador, with more frequent findings in Argentina and Brazil. In this chapter, we review these fossil findings and the main aspects of the taxonomy, systematics, and paleogeographic implications of this record and briefly discuss the importance of these occurrences for the understanding of titanosaur evolution. The diversity and abundance of end-Cretaceous titanosaur taxa in South America represent about 25% of the known Titanosauria species in the world, which makes them the most common group of large terrestrial herbivores of that time. Cretaceous titanosaurs from South America also vary highly in morphology and size, comprising small to large-sized taxa, for example. Their record mainly consists of appendicular and axial remains, including rare skull material, but also comprises eggs, nests, footprints, and coprolites. In South America, by the end of the Late Cretaceous, titanosaurs were generally represented by more derived titanosaurians that are mainly taxonomically assigned to more derived species within Aeolosaurini and Saltasaurinae.
Given that bone microstructure is a very important source of paleobiological information, several paleohistological studies have been conducted on sauropodomorph dinosaurs, possibly making this clade the most studied histologically. Despite these paleohistological studies on sauropodomorph dinosaurs from South America are relatively scarce in comparison with other regions of the world, significant progress on this matter (i.e. paleohistology of South American sauropodomorphs) has been made during the last decade. Following an order from rather specific to more generalized issues, the most important advances are related to the origin of particular skeletal elements (i.e. osteoderms, sacral supraspinous ossifications and extremely elongated cervical ribs), the growth patterns of basal sauropodomorphs and the variation on sauropod growth dynamics and its relationship with gigantism. Regarding the origin of osteoderms and extremely elongated cervical ribs, these structures have been formed by metaplastic ossification of dermal and tendinous tissues, respectively. Their histological characterization has been helpful to discover that the alleged osteoderms of Agustinia ligabuei were actually dorsal and cervical ribs. The long bone histology of basal sauropodomorphs has revealed that the cyclical growth pattern assumed for this group actually showed some degree of variation. Finally, a clade of basal sauropods (i.e. lessemsaurids) was characterized by a cyclical growth pattern, not previously reported for other sauropods, combined with episodes of highly accelerated growth rates. The gigantic body sizes obtained by lessemsaurids were therefore reached through a growth strategy different from that developed by eusauropods (i.e. rapid and continuous growth).
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The rediscovery of the Pouech collection allows us to describe a reptile fauna from the Upper Cretaceous (Maastrichtian) of Le Mas d'Azil area. Most of these forms had never been described in this area [an ankylosaur, a trematochampsid crocodile, an eusuchian crocodile with crushing posterior teeth (alligatorid?) and a chelonian]. Titanosaurid and hypsilophodontid dinosaurs are also represented in the fauna. Variations of the faunal composition between some Upper Cretaceous European localities are discussed for the first time. They can be explained by paleoecological, paleogeographical and stratigraphical differences. -English summary
Brachyuran decapods (crabs) are rarely preserved in coastal environments and have not been documented in close association with dinosaur fossils. A crab referable to the Necrocarcinidae Förster, 1968, is here described from the Cenomanian Bahariya Formation, Bahariya Oasis, Egypt. The occurrence of a crab in a mangrove setting in association with terrestrial vertebrates is extremely unusual in the fossil record. Review of decapod occurrences from the region has resulted in placement of Portunus sp. of Roger (1946) within the Raninidae de Haan, 1839, and Lophoraninella cretacea (Dames, 1886) into the Galatheoidea Samouelle, 1819. The crabs of the Bahariya Formation were probably scavengers, feeding on vegetation and other organic material and were possibly a food source for fish and other vertebrates and invertebrates in the environment.
Provisional global shorelines are presented, on palaeo-positioned continents, for all 12 Cretaceous stages. A stage-by-stage commentary and bibliography of sources for the Cretaceous shorelines of Europe is represented as an example of the degree of spatial and temporal uncertainty inherent in all shoreline reconstructions, even in a well-researched region. -Author