Journal of South American Earth Sciences xxx (2015) 1e7

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Journal of South American Earth Sciences

journal homepage: www.elsevier.com/locate/jsames

Cranial bones and atlas of titanosaurs (Dinosauria, Sauropoda) from

Late Cretaceous (Bauru Group) of Uberaba, Minas Gerais State, Brazil
Agustn G. Martinelli a, b, *, Thiago da Silva Marinho a, c, Leonardo S. Filippi d,
Luiz Carlos Borges Ribeiro a, Mara Lcia da Fonseca Ferraz a, Camila Lourencini Cavellani a,
Vicente de Paula Antunes Teixeira a
a

gicas L. I. Price, CCCP/UFTM, 38001-970, BR-262, Km 784, Peiro

polis, Uberaba, MG, Brazil
Centro de Pesquisas Paleontolo
Departamento de Paleontologia e Estratigraa, Instituto de Geoci^
encias, Universidade Federal de Rio Grande do Sul, Av. Bento Gonalves 9500, Agronomia,
91501-970, Porto Alegre, RS, Brazil
c
~o (ICENE), UFTM, Av. Randolfo Borges Jr. 1700, Univerdecidade, 38064-200, Uberaba, Minas Gerais, Brazil
Instituto de Ci^
encias Exatas, Naturais e Educaa
d

n de los Sauces, Neuqu

Museo Municipal Argentino Urquiza, Jujuy y Chaco s/n, (Q8319BFA), Rinco
en, Argentina
b

a r t i c l e i n f o

a b s t r a c t

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.,

* Corresponding author. Departamento de Paleontologia e Estratigraa, Instituto

^ncias, Universidade Federal de Rio Grande do Sul, Av. Bento Gonalves
de Geocie
9500, Agronomia, 91501-970, Porto Alegre, RS, Brazil.
E-mail address: agustin_martinelli@yahoo.com.ar (A.G. Martinelli).

Kellner and Azevedo, 1999; Kellner and Campos, 2000; Santucci

and Bertini, 2001, 2006; Kellner et al., 2005, 2006; Salgado and
Carvalho, 2008; Bittencourt and Langer, 2011). The Bauru Group
(Bauru Basin) includes the species Gondwanatitan faustoi (Ada~o Paulo; Kellner and Azevedo, 1999), Adamantina Formation, Sa
~o Paulo; Santucci
mantisaurus mezzalirai (Adamantina Formation, Sa
and Bertini, 2006), Aeolosaurus maximus (Adamantina Formation,
~o Paulo; Santucci and Arruda-Campos, 2011), Brasilotitan nemSa
~o Paulo; Machado et al., 2013),
ophagus (Adamantina Formation, Sa
Maxakalisaurus topai (Adamantina Formation, Minas Gerais;
Kellner et al., 2006), Baurutitan britoi (Marlia Formation, Minas
Gerais; Kellner et al., 2005), Trigonosaurus pricei (Marlia Formation,
Minas Gerais; Campos et al., 2005), and Uberabatitan ribeiroi
(Marlia Formation, Minas Gerais; Salgado and Carvalho, 2008).
Outside this basin, Gondwanatitan sp. was also recognized in the
Cambambe Formation, Parecis Basin, in Mato Grosso State (FrancoRosas et al., 2004) and Tapuiasaurus macedoi is known from the

Formation, Sanfranciscana Basin, Minas Gerais State (Zaher
Quirico
et al., 2011). Beside the already described species, there is a huge
fossil record based upon isolated remains that could be indicating a
noticeably, still poorly known, higher diversity of titanosaurs (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).

1.1. Institutional abreviations

gicas Llewellyn Ivor
CPPLIP, Centro de Pesquisas Paleontolo

polis/UFTM, Peiro

pPrice, Complexo Cultural e Cientco de Peiro
olis, Uberaba, MG, Brazil.

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

polis, Uberaba County, Minas Gerais State

Fig. 1. Isolated left prefrontal CPPLIP 1241 from the Late Cretaceous Marlia Formation, site Ponto 1 do Price at Serra do Veadinho, Peiro
(Brazil) in lateral (A), medial (B), dorsal (C), and ventral (D) views. Abbreviations: b, orbital border; ffr, facet for the frontal; fn, facet for the nasal. Gray areas indicate broken surfaces.

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

polis, Uberaba County, Minas Gerais State

Fig. 2. Isolated left squamosal CPPLIP 296 from the Late Cretaceous Marlia Formation, site Ponto 2 do Price at Serra do Veadinho, Peiro
(Brazil) in lateral (A), medial (B), anterior (C), and posterior (D) views. Abbreviations: bif, border of the infratemporal fenestra; bsf, border of the supratemporal fenestra; fpa, facet
for parietal; fpo, facet for postorbital; fpp, facet for the paroccipital process; fq, facet for quadrate; vp, ventral process. Gray areas indicate broken surfaces.

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

SAURISCHIA Seeley, 1888

SAUROPODA Marsh, 1878
TITANOSAURIA Bonaparte and Coria, 1993
LITHOSTROTIA Upchurch, Barrett, and Dodson, 2004
Gen. et sp. indet.
3.1. Referred material
CPPLIP 247, isolated partial atlas vertebra (Fig. 3); CPPLIP 0296,
isolated left squamosal (Fig. 2); CPPLIP 1241, isolated left prefrontal
(Fig. 1).
3.2. Locality and horizon
CPPLIP 1241 comes from the site Ponto 1 do Price (19 430 2400 S,
at Serra do Veadinho, located approximately 2 km

polis, Uberaba County, Minas Gerais
north of the rural town of Peiro
State, Brazil. CPPLIP 247 and CPPLIP 296 come from the site known
as Ponto 2 do Price (19 4301300 S, 47450 0600 W), at Serra do Vea
polis, Uberaba
dinho, situated approximately 2.5 km north of Peiro
47440 4500 W),

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

polis, Uberaba County, Minas Gerais

Fig. 3. Isolated partial atlas vertebra CPPLIP 247 from the Late Cretaceous Marlia Formation, site Ponto 2 do Price at Serra do Veadinho, Peiro
State (Brazil) in posterior (A), left lateral (B), anterior (C), and right lateral (D) views. Abbreviations: ic, intercentrum; np, neural spine.

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

several articular facets to accommodate others cranial bones (i.e.,

postorbital, parietal, exoccipital opisthotic, and quadrate). The
body of the squamosal has a lateral, vertical and almost at surface,
with a laminar descending process (or ventral process), and a
dorsal, slightly convex surface, which participates of the skull roof.
In lateral view, the squamosal bears a well-delimited, deeply
concave scar for contact with the postorbital (Fig. 2). This concavity
faces latero-anteriorly and its ventral edge coincides with the
beginning of the postero-dorsal border of the infratemporal
fenestra (Fig. 2). The descending process is thin, long, slender and
D-shape in cross-section with the outer surface slightly convex and
the inner surface slightly convex to straight. The ventral and
antero-ventral edges of this process are broken off. As preserved, it
seems that anterior and posterior borders are sub-parallel but
possibly the anterior edge slightly diverges ventrally. The anterior
edge of the descending process delimits the posterior border of the
infratemporal fenestra. On the other hand, its posterior edge contacts the quadrate through an articular surface which extends onto
most of the inner surface (Fig. 2).
In posterior view, the main features are the concave articular
facet for the paroccipital process of the opisthotic and the inner
articular facet for the quadrate (Fig. 2). The scar of the paroccipital
process is postero-medially projected and not visible in lateral
view. In medial view, the facet for the parietal is oval shaped, with a
rugose texture, located in the dorsal-most portion of the squamosal
body. Next to this facet, there is a small portion of the posterior
edge of the supratemporal fenestra. The articular facet for the
quadrates is best seen in medial view; it has some shallow, subparallel dorso-ventrally oriented ridges (Fig. 2). This facet is anteriorly delimited by a medial process that becomes taller upward.
The long and slender descending process of the squamosal is
similar to that of Rapetosaurus (Curry Rogers and Forster, 2001,
2004) and Nemegtosaurus (Wilson, 2005) and rather different to
that of Tapuiasaurus, which has a more robust and dorsoventrally
shorter process (Zaher et al., 2011), and Phuwiangosaurus sirindhornae, in which it is very short and hook-shaped (Suteethorn
et al., 2009). In CPPLIP 296 there is clear evidence that the squamosal participates of the posterior edge of the supratemporal
fenestra as in Rapetosaurus (Curry Rogers and Forster, 2004),
differing from Nemegtosaurus and Quaesitosaurus orientalis
Kurzanov and Bannikov, 1983, in which this bone is excluded from
this fenestra (Nowinski, 1971; Upchurch, 1995; Wilson, 2005). In
Quaesitosaurus the squamosal is a massive and elongated bone,
with dorsal and posterior surface intensely roughened (Kurzanov
and Bannikov, 1983), differing from the smooth surface of CPPLIP
296.
The taxonomic referral of the squamosal to Titanosauria indet. is
supported by its resemblance to the squamosal of the derived
lithostrotian Rapetosaurus, from the Late Cretaceous of Madagascar
(Curry Rogers and Forster, 2001, 2004). Although the specimen
CPPLIP 296 cannot be referred to any particular taxon, its
morphology is reminiscent of lithostratian titanosaur rather than to
basal forms. For example, in more basal taxa, such as the basal
macronarian Camarasaurus sp. (Madsen et al., 1995) and nontitanosaur titanosauriform Abydosaurus mcintoshi (Chure et al.,
2010), the squamosal is a stout element, quite different from the
morphology of CPPLIP 296. Also, this element is quite different from
the squamosal of the diplodocid Diplodocus, in which there is a
prominent medial process (e.g., Yu, 1993) not observed in CPPLIP
296 and other titanosaurs (Curry Rogers and Forster, 2004).
4.3. Atlas vertebra
CPPLIP 247 consists of an almost complete atlas (Fig. 3). The
fusion of the neurapophyses to the intercentrum indicates that the

specimen belongs to an adult individual. The atlantal intercentrum

is robust, much wider transversely than it is high, such as Mongolosaurus (Mannion, 2010) and Futalongkosaurus (Calvo et al.,
2007). The anterior face of the intercentrum, for the occipital
condyle, is concave and the posterior one is at, such as in other
titanosaurian atlas. The lateral surface of the intercentrum is mildly
antero-posteriorly concave, such as in Quetecsaurus rusconii
(Gonz

alez Riga and Ortiz David, 2014), Mongolosaurus, and Rapetosaurus (Curry Rogers, 2009). Similar to Rapetosaurus and Futalognkosaurus, CPPLIP 247 lacks the two well-developed
posteroventral processes which are present in Quetecsaurus and
very reduced in Erketu ellisoni. The neurapophysis is proximally
higher than in Mongolosaurus, Rapetosaurus, Futalongkosaurus, and
Erketu ellisoni (Ksepka and Norell, 2006). The upper parts of the
neurapophyses are thin, and posterodorsally projected, but do not
approach one another on the midline, similar to Mongolosaurus,
Futalongkosaurus and Rapetosaurus. In CPPLIP 247, the parts of the
neurapophyses are more separated medially that other titanosaur
atlas. The atlas of Erketu shows the neurapophyses fused at the
midline. The neurapophyses in CPPLIP 247 are posterodorsally
projected in an angle of 45 similar to Mongolosaurus. In Futalongkosaurus the angle is about 50 while in Rapetosaurus and
Erketu, between 30 and 40 .
5. Discussion and conclusion
The titanosaur context at the paleontological quarries located

polis town (Uberaba, Minas
along the Serra do Veadinho, near Peiro
Gerais State, Brazil), is quite diverse including the titanosaurs

rez Valieri and Daz,
Trigonosaurus pricei (Campos et al., 2005; Jua
2013) and Baurutitan britoi (Kellner et al., 2005). In addition,
there is an isolate caudal vertebra of an Aeolosaurini indet.
(Santucci and Bertini, 2001; Martinelli et al., 2011; Filippi et al.,
2013) and several specimens of indeterminate titanosaurs (e.g.,
Campos and Kellner, 1999; Santucci and Bertini, 2001; Marinho and
Candeiro, 2005). The specimens here described constitute the rst
cranial bones of titanosaurs from this region and unfortunately
cannot be referred to any particular taxon rather than indeterminate lithostrotian titanosaurs. The skull anatomy of titanosaurs is
largely based upon only a few, almost complete, skulls (e.g., Nemegtosaurus, Rapetosaurus, Tapuiasaurus; Nowinski, 1971; Curry
Rogers and Forster, 2001, 2004; Wilson, 2005; Zaher et al., 2011)
and isolated and/or non-articulated remains (such as, mandibles,
braincases, isolated bones) (e.g., Powell, 2003; Apestegua, 2004;
Martinelli and Forasiepi, 2004; Kellner et al., 2006; Garca et al.,
2008; Filippi and Garrido, 2008; Filippi et al., 2011; Gallina and
Apestegua, 2011; Machado et al., 2013; Curry Rogers and Wilson,
2014), which in comparison with the total number of recognized
species, this information is still quite limited.
The prefrontal and squamosal have some features that are
reminiscent of derived lithostratian titanosaurs, such as Rapetosaurus from Madagascar and Tapuiasaurus from Brazil. The squamosal CPPLIP 296 has a long and slender descending process as in
Rapetosaurus (Curry Rogers and Forster, 2001, 2004) and Nemegtosaurus (Wilson, 2005). Contrarily, this process is more robust
and dorsoventrally shorter in Tapuiasaurus (Zaher et al., 2011). According to the phylogenetic hypothesis of Zaher et al. (2011),
Nemegtosaurus
is
the
sister
taxon
of
the
Rapetosaurus Tapuiasaurus clade, all constituting a monophyletic
Nemegtosauridae. Therefore, two congurations of the descending
process of the squamosal are recognized into this family. As a result,

polis has a character-state similar to the
the specimen from Peiro
non-South American Rapetosaurus and Nemegtosaurus rather than
to Tapuiasaurus from Brazil. Because the distribution of this character among others non-nemegtosaurid advance titanosaurs is

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