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Article history:
Received 12 October 2014
Accepted 19 February 2015
Available online xxx
Isolated left prefrontal, left squamosal and atlas of titanosaur dinosaurs are described and compared.
They come from the Late Cretaceous Serra da Galga Member of the Marlia Formation at the Serra do
polis (Uberaba County, Minas Gerais State, Brazil). Due to the sparse cranial eleVeadinho region, Peiro
ments of titanosaurs already known from Brazil, these specimens are noticeable to be presented. In
addition, the atlas vertebra is described for the rst time for Brazilian titanosaurs. The morphology of the
cranial bones closely resembles lithostratian titanosaurs, such as Rapetosaurus, rather than basal titanosaurs. The atlas is similar to that of other titanosaurs, suggesting that the anatomy of this element
seems to be more conservative than other vertebral elements, in which vertebral laminae play an
important rule in titanosaur taxonomy.
2015 Elsevier Ltd. All rights reserved.
Keywords:
Titanosauria
Lithostratia
polis
Peiro
Prefrontal
Squamosal
1. Introduction
Titanosaur sauropods achieved a wide taxonomic diversity
mainly in Gondwanan landmasses during the Late Cretaceous (e.g.,
Powell, 2003; Wilson, 2006; Novas, 2009). The fossil record of this
group is particularly well-documented and studied in South
America, with about forty named species (e.g., Bonaparte, 1996;
Powell, 2003; Novas, 2009). Remains of titanosaurs occur in
almost all Late Cretaceous faunal associations of South America,
exhibiting a broad spatial as well as temporal distribution (e.g.,
Bonaparte, 1996; Santucci and Bertini, 2001; Powell, 2003; Wilson,
2006; Salgado and Bonaparte, 2007; Novas, 2009; Bittencourt and
Langer, 2011). With regard to the Brazilian fossil record, titanosaurs are particularly notorious integrant of post-Cenomanian
Cretaceous continental assemblages of southeastern Brazil (e.g.,
http://dx.doi.org/10.1016/j.jsames.2015.02.009
0895-9811/ 2015 Elsevier Ltd. All rights reserved.
Please cite this article in press as: Martinelli, A.G., et al., Cranial bones and atlas of titanosaurs (Dinosauria, Sauropoda) from Late Cretaceous
(Bauru Group) of Uberaba, Minas Gerais State, Brazil, Journal of South American Earth Sciences (2015), http://dx.doi.org/10.1016/
j.jsames.2015.02.009
A.G. Martinelli et al. / Journal of South American Earth Sciences xxx (2015) 1e7
Campos and Kellner, 1999; Kellner and Campos, 2000; Bertini et al.
2001; Santucci and Bertini, 2001; Marinho and Candeiro, 2005;
Lopes and Buchmann, 2008; Santucci, 2008; Bittencourt and
Langer, 2011; Martinelli et al., 2011).
Of the aforementioned Brazilian species, cranial elements are
only known in Maxakalisaurus, Brasilotitan, and Tapuiasaurus. The
holotype of Maxakalisaurus includes a fragment of right maxilla
with teeth (Kellner et al., 2006), whereas the holotype of Brasilotitan includes a partial lower jaw with some teeth (Machado et al.,
2013). Conversely, the holotype of Tapuiasaurus includes an almost
complete skull and lower jaws, with teeth, which constitutes one of
the most complete titanosaur skull ever found (Zaher et al., 2011);
its full description will bring new information on cranial titanosaur
morphology.
In this note, two isolated cranial bones (Figs. 1e2) and an isolated atlas (Fig. 3) from Late Cretaceous Serra da Galga Member of
polis
the Marlia Formation at the Serra do Veadinho region, Peiro
(Uberaba, Minas Gerais State, Brazil), are described. Due to the
sparse cranial elements of titanosaurs already known from Brazil,
these specimens are noticeable to be presented and compared. In
addition, the atlas vertebra is described for the rst time for Brazilian titanosaurs (the atlas of Tapuiasaurus is known but not
described yet; Zaher et al., 2011).
2. Geological settings
The sediments of the Bauru Basin were deposited in an inland
depression formed by themomechanical subsidence related to the
opening of the South Atlantic Ocean. It comprises roughly an area of
s,
370,000 km2, including portions of the Brazilian states of Goia
Mato Grosso, Mato Grosso do Sul, Minas Gerais (represented by the
^ngulo Mineiro), Parana
and Sa
~o Paulo
region known as Tria
(Fernandes and Coimbra, 1996). Bauru Basin is divided in two
Group, with Goio
^ Ere
^, Rio Parana
and Santo Anasta
cio
groups: Caiua
formations, and Bauru Group, with Uberaba, Adamantina (Vale do
~o Jose
do Rio Preto, Presidente Prudente
Rio do Peixe, Araatuba, Sa
sensu Fernandes, 2004) and Marlia formations (Fernandes and
Coimbra, 1996). The age of the rocks of the Bauru Group is
considered by some authors as ranging from the Turonian to
Maastrichtian (Dias-Brito et al., 2001), but contrarily GobboRodrigues et al. (1999) consider a shorter chronostratigraphic
range, suggesting it is Campanian-Maastrichtian based on the os~o Paulo State. Magnetracods of the Adamantina Formation of Sa
tostratigraphic studies by Tamrat et al. (2002) suggest that Uberaba
Formation could not be older than Campanian, strengthening the
hypothesis by Gobbo-Rodrigues et al. (1999).
The specimens here studied come from the Serra do Veadinho,
polis, from levels of the Serra da Galga Member of the
near Peiro
Marlia Formation. This member is a highly fossiliferous unit,
composed of conglomerates and coarse-grained sandstones to negrained sandstones in ning-upwards cycles (Novas et al., 2008;
Salgado and Carvalho, 2008). Within Serra do Veadinho region,
CPPLIP 1241 (prefrontal bone) comes from the outcrop known as
Ponto 1 do Price or Caieira Site whereas CPPLIP 296 (squamosal
Please cite this article in press as: Martinelli, A.G., et al., Cranial bones and atlas of titanosaurs (Dinosauria, Sauropoda) from Late Cretaceous
(Bauru Group) of Uberaba, Minas Gerais State, Brazil, Journal of South American Earth Sciences (2015), http://dx.doi.org/10.1016/
j.jsames.2015.02.009
A.G. Martinelli et al. / Journal of South American Earth Sciences xxx (2015) 1e7
bone) and CPPLIP 247 (atlas vertebra) from the outcrop Ponto 2 do
Price.
Both sites present tabular geometry, planar cross-stratications
and channel cross-stratications (Novas et al., 2008). The ningupwards cycles that occur on this outcrops are characterized by
ne sandstones interbedded with pelites, clayish sandstones and
coarse sandstones with mud intraclasts (Novas et al., 2008). These
features suggest that these sediments were deposited by braided
uvial systems that produced wide alluvial plains with small ponds
and lakes in semi-arid to arid climate, and seasonality marked by
long dry periods intercalated with heavy rains and ash-ood
events (Garcia et al., 1999; Goldberg and Garcia, 2000; Novas
et al., 2008). Despite the evidences of aridity, this paleoenvironment was humid enough to support water dependent organisms
such as shes, anurans, turtles and semi-aquatic crocodyliforms,
found as autochthonous or parauthochthonous remains.
3. Systematic paleontology
DINOSAURIA Owen, 1842
Please cite this article in press as: Martinelli, A.G., et al., Cranial bones and atlas of titanosaurs (Dinosauria, Sauropoda) from Late Cretaceous
(Bauru Group) of Uberaba, Minas Gerais State, Brazil, Journal of South American Earth Sciences (2015), http://dx.doi.org/10.1016/
j.jsames.2015.02.009
A.G. Martinelli et al. / Journal of South American Earth Sciences xxx (2015) 1e7
County, Minas Gerais State, Brazil. All specimens come from Serra
da Galga Member, Marlia Formation (Campanian-Maastrichtian),
Bauru Group.
4. Description and comparisons
4.1. Prefrontal
The prefrontal is a small bone which forms part of the dorsal
edge of the orbit. CPPLIP 1241 consists of an almost complete left
prefrontal, lacking part of the anterior process (Fig. 1). It is a rather
stout bone, fairly rhomboidal (considering the broken anterior
process) in dorsal and ventral views and gently convex dorsally in
lateral view, due to its correspondence with the dorsal orbital edge.
This edge is rounded and, medially and anteriorly the prefrontal
becomes thicker. In the anterior portion of the orbital edge, there
are two small nutritious foramina and distally, there is a notch for
another foramen, which was enclosed between this bone and the
frontal. In dorsal view, the prefrontal is convex, with a small
nutritious foramen near the base of the anterior process (Fig. 1). The
surface of CPPLIP 1241 lacks ornamentation. In contrast, the anterodorsal margin of the orbit, on the prefrontal, of Nemegtosaurus
mongoliensis (Wilson, 2005) is heavily ornamented by several small
pointed projections. Rapetosaurus krausei, also possess ornamentation, but much more subtle near its contact with the frontal
(Wilson, 2005). The only partially visible facets in this view are for
the frontal and the nasal. These facets are better observed in medial
and ventral views. The facet for the nasal occupies two-third of the
prefrontal's antero-medial edge. It is almost vertical, relatively high,
with small furrows, and encloses a ventrally opened vascular
groove, as in Rapetosaurus (Curry Rogers and Forster, 2004). On the
other hand, the facet for the frontal is restricted to the one-third
distal medial edge of the prefrontal in medial view, but with a
great development under this bone. As such, the prefrontal overlaps the frontal, leaving a large articular surface. Hence, in ventral
view, the concave surface of the prefrontal has two main surfaces:
the smooth roof of the orbital cavity and the rough articular facet
for the frontal (Fig. 1).
The anterior process of the prefrontal is broken off but based on
its sub-triangular cross-section it should have been elongated and
slender, as in titanosaurs (Rapetosaurus, Tapuiasaurus; Curry Rogers
and Forster, 2004; Zaher et al., 2011), for contact to the lacrimal.
Contrarily, in Nemegtosaurus the anterior process is reduced
(Wilson, 2005) and in basal macronarians, such as Camarasaurus
(Madsen et al., 1995), this process is absent. Although the general
shape of this bone is similar to that of Rapetosaurus, the articular
facet for the frontal is better developed than in the African species
and the posterior end is not transversely straight.
4.2. Squamosal
The isolated squamosal CPPLIP 296 corresponds to the left side
of the skull. It is an L-shaped bone in lateral view, with a main body
and a cranio-ventrally descending process (Fig. 2). This element has
Please cite this article in press as: Martinelli, A.G., et al., Cranial bones and atlas of titanosaurs (Dinosauria, Sauropoda) from Late Cretaceous
(Bauru Group) of Uberaba, Minas Gerais State, Brazil, Journal of South American Earth Sciences (2015), http://dx.doi.org/10.1016/
j.jsames.2015.02.009
A.G. Martinelli et al. / Journal of South American Earth Sciences xxx (2015) 1e7
Please cite this article in press as: Martinelli, A.G., et al., Cranial bones and atlas of titanosaurs (Dinosauria, Sauropoda) from Late Cretaceous
(Bauru Group) of Uberaba, Minas Gerais State, Brazil, Journal of South American Earth Sciences (2015), http://dx.doi.org/10.1016/
j.jsames.2015.02.009
A.G. Martinelli et al. / Journal of South American Earth Sciences xxx (2015) 1e7
still poorly known, we could not conclude with condence that the
squamosal CPPLIP 296 belongs to nemegtosaurid. At present, it is
more parsimonious to consider CPPLIP 296 as a derived lithostratian titanosaur, with close resemblance with some nemegtosaurids. Nonetheless, CPPLIP 296 is clearly differentiated from
the squamosal of Tapuiasaurus, indicating it belongs to a different
taxon.
With regard to the prefrontal, CPPLIP 1241 is more similar to
that of Rapetosaurus and Tapuiasaurus (Curry Rogers and Forster,
2004; Zaher et al., 2011) than to that of Nemegtosaurus (Wilson,
2005). Because the conguration of this bone among titanosaurs
is also poorly known, CPPLIP 1241 could only be referred as a
lithostratian titanosaur (see comparisons in the description).
Although based on sparse data, the prefrontal and squamosal
polis
here described are indicative that the titanosaurs from Peiro
region had a skull conguration similar to nemegtosaurids, such as
Rapetosaurus. Also, the difference in size between the squamosal
(smaller) and the prefrontal (larger) indicates different ontogenetic
stages or two different taxa with different skull sizes. Only more
complete and associated specimens will permit to elucidate these
problems.
The atlas, although incomplete, has a similar pattern that of
other titanosaurs, such as Futalongkosaurus (Calvo et al., 2007),
Rapetosaurus (Curry Rogers, 2009), and Mongolosaurus (Mannion,
2010). Therefore, the anatomy of this element seems to be more
conservative than other vertebral elements, in which vertebral
laminae play an important rule for titanosaur taxonomy (e.g.,
Bonaparte, 1999; Wilson, 2012).
Acknowledgments
We thank the continuous support of all the staff of the Complexo
polis (CCCP) of the Universidade FedCultural e Cientico de Peiro
eral do Tri^
angulo Mineiro (UFTM), Uberaba, MG, Brazil. This
contribution was beneciated by the nancial support of the
~o de Amparo a
Pesquisa do Estado de Minas Gerais
Fundaa
(FAPEMIG), the Conselho Nacional de Desenvolvimento Cientco e
gico (CNPq), the Coordenaa
~o de Aperfeioamento de PesTecnolo
soal de Nvel Superior (CAPES), the Funda~
ao de Ensino e Pesquisa
rio de Cie
^ncia Tecnologia e
de Uberaba (FUNEPU) and the Ministe
~o (MCTI). The reviewers M. Ezcurra, F. Agnolin and the the
Inovaa
Editor J. N. Kellogg and C. R. Candeiro provided useful comments
that greatly improved the Ms.
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Please cite this article in press as: Martinelli, A.G., et al., Cranial bones and atlas of titanosaurs (Dinosauria, Sauropoda) from Late Cretaceous
(Bauru Group) of Uberaba, Minas Gerais State, Brazil, Journal of South American Earth Sciences (2015), http://dx.doi.org/10.1016/
j.jsames.2015.02.009