Cretaceous Research 80 (2017) 38e55

Contents lists available at ScienceDirect

Cretaceous Research
journal homepage: www.elsevier.com/locate/CretRes

A new Brachiosauridae Sauropod dinosaur from the lower Cretaceous

of Europe (Soria Province, Spain)
Rafael Royo-Torres a, *, Carolina Fuentes b, Manuel Meijide b, Federico Meijide-Fuentes b,
Manuel Meijide-Fuentes b
a
n Conjunto Paleontolo
Fundacio
gico de Teruel-Dino

polis/Museo Aragon

es de Paleontología, Av. Sagunto s/n, Teruel, E-44002, Spain
b
C/ Almaza
n 17, 2º C, Soria, E-42004, Spain

a r t i c l e i n f o a b s t r a c t

Article history: A new dinosaur sauropod from the Golmayo Formation (upper Hauterivian-lower Barremian) in Soria
Received 21 March 2017 province (Spain) is described as Soriatitan golmayensis gen. et sp. nov. The new material consists of one
Received in revised form tooth, three dorsal vertebrae with ribs, a partial sacrum, five caudal centra, two caudal vertebral spines,
8 August 2017
one chevron, a humerus, an ulna, a radius, two partial ilia, two ischia, a fragment of pubis, and a partial
Accepted in revised form 28 August 2017
Available online 31 August 2017
femur. Cladistic analysis identified the material as belonging to Brachiosaruidae within Titanosaur-
iformes. Sauropod material from the Golmayo Formation shares a diagnostic feature with Abydosaurus,
Cedarosaurus, Tastavinsaurus and Venenosaurus including anteriorly deflected anterioremiddle caudal
Keywords:
Soriatitan golmayensis
neural spines and with Cedarosaurus a proximodistally straight lateral margin between the proximal
Golmayo Formation head and the shaft of the humerus. Eight characters are potential autapomorphies in the sauropod from
Hauterivian-Barremian Spain, including the presence of a large rectangular ventral ridge below the preacetabular process of the
Brachiosauridae ilium, the same length of the pubic and ischiadic blades in the ischium, and two lateral ridges in the
Sauropod lateral area of the deltopectoral crest of the humerus. The presence of Early Cretaceous brachiosarids in
Europe both, North America and Europe, give support to the hypothesis of a connection between the tectonic
North America plates of these continents at some point during the Early Cretaceous.
© 2017 Elsevier Ltd. All rights reserved.

1. Introduction
Sauropoda have been widely recorded in the Mesozoic, from
their origin in the Late Triassic until their extinction at the end of
the Cretaceous (McIntosh, 1990; Upchurch et al., 2004). During
Hauterivian-Aptian transition (Early Cretaceous) they were com-
mon megaherbivores in the Iberian Peninsula, with the fossil evi-
dence revealing a high diversity of sauropods across the Iberian
Plate. To date, specimens identified as rebachisaurids and titano-
sauriforms have been recovered (Ortega et al., 2006; Pereda-
Suberbiola, 2006; Canudo et al., 2008; Royo-Torres, 2009; Torcida
Ferna
ndez-Baldor et al., 2011; Royo-Torres et al., 2012). In this work
we evaluate a new dinosaur sauropod from Soria province (Spain)
and place it in its stratigraphical context. We present the descrip-
tion and illustrations of a new genus and species. Comparisons
with other sauropods result in the recognition of several
* Corresponding author.
E-mail addresses: royo@dinopolis.com (R. Royo-Torres), carolfuentes@ono.com
(C. Fuentes).
autapomorphies for this new taxon. We also analyze the phyloge-
netic relationships of this specimen based on recent analysis of
titanosauriform interrelationships (D'Emic et al., 2016; Mannion
et al., 2017).
The specimen described herein was discovered in the province

n, Spain). The area, Los Can
of Soria (Castilla y Leo ~ os site at Golmayo,
has a history of dinosaur discovery going back to 1927 (Royo y
Go
mez, 1926a,b,c; Sanz et al., 1992). The Fuentes-Meijide team
have worked in this area, focusing on prospecting and excavating in
the Golmayo Formation and recovering remains in the area around
the towns of Soria: Golmayo and Villaciervos (Fig. 1AeC). In 2000,
the Zorralbo I site was discovered and was followed in 2006 and
2007 by Zorralbo II and III (Fuentes-Vidarte et al., 2005, 2008,
2009a,b). Between 2000 and 2004, Zorralbo I was excavated and
the remains of several dinosaurs were found over a 100 m2 area;
their positions were mapped using a square-meter grid system
(Fig. 1D). The site has yielded dinosaurs such as Polacanthus sp
(Pereda-Suberbiola et al., 2007), the iguanodontian Magnamanus
soriaensis (Fuentes Vidarte et al., 2016), Hypsilophodontidae indet.
and dromeosaurid theropods, as well as turtles and crocodiles

http://dx.doi.org/10.1016/j.cretres.2017.08.012
0195-6671/© 2017 Elsevier Ltd. All rights reserved.
 R. Royo-Torres et al. / Cretaceous Research 80 (2017) 38e55 39

Fig. 1. Geographical and geological location of Zorralbo I site (Soria, Spain). A, Geographic position of Golmayo with the location of Zorralbo I. B, General stratigraphic section of
Golmayo area indicating the position of the vertebrate fossil site studied in this work. C, Geological map of the Zorralbo I site at Golmayo, modified from Pereda-Suberbiola et al.
(2007) and Martín-Closas and Alonso (1998). D, And excavation map of Soriatitan golmayensis for the Zorralbo I site.

(Fuentes Vidarte et al., 2005, 2009a,b). The new sauropod specimen
studied here is the fifth dinosaur from the Zorralbo I site. It was
initially described as a Brachiosauridae indet (Fuentes Vidarte et al.,
2005), afterwards being related to the Titanosauriformes clade
(Royo-Torres et al., 2009). All material is housed in the Museo
Numantino de Soria (MNS; exp. 334.06-So).
Teruel, Spain (CPT and MAP for the Museo Aragone
s de Paleon-
tología); MNS, Museo Numantino de Soria, Soria, Spain; MFN,
Museum für Naturkunde eLeibniz Institute for Research on Evo-
lution and Biodiversity at the Humboldt University Berlin, Berlin,
Germany; NHMUK, Natural History Museum, London, UK (previ-
ously BMNH).

1.1. Institutional abbreviations 2. Geographical and geological setting of the Zorralbo I site

DMNH, Denver Museum of Natural History, Denver, USA; The Zorralbo I site is located near the village of Zorralbo, in the

n Conjunto Paleontolo
FCPTD, Fundacio
gico de Teruel-Dino

polis, municipality of Golmayo, to the west of the city of Soria in northern
40 R. Royo-Torres et al. / Cretaceous Research 80 (2017) 38e55
Spain (UTM coordinates: 30TWM394238, Mapa Topografico
Nacional de Espan ~ a, 1:50000, sheet 342) (Fig. 1BeC). Geologically,
the Zorralbo I site lies within the eastern sector of the Cameros
Basin, on the north-western margin of the Iberian Range. The
Cameros Basin has been interpreted as an intraplate rift basin that
developed during the Late Jurassic and Early Cretaceous. It was
filled by a thick succession of alluvial and lacustrine deposits, with
only rare marine incursions (Alonso and Mas, 1993). The sedi-
mentary succession can be divided into six main depositional se-
quences, which correlate with marine sequences (Mas et al., 1993;
Mas and Salas, 2002). The Zorralbo I site lies within the Golmayo
Formation (Fig. 1A) (Fuentes Vidarte et al., 2005, 2009a; Pereda-
Suberbiola et al., 2007). The latter is 800 m thick and consists of
channel sandstones, micritic limestones, red lutites and, to a lesser
extent, siliciclastic and calcareous conglomerates. These sedimen-
tary rocks were deposited in a fluvio-lacustrine environment
(Clemente and Alonso, 1990). The Golmayo Formation is limited to
the Soria region, extending from the eastern foothills of the Sierra
de Cabrejas to the Sierra de San Marcos (both Upper Cretaceous) in
a narrow strip west of the city of Soria. The Cuevas de Soria fault
underlies this formation along the NE-SW axis. The Golmayo For-
mation overlies the Hoya del Moro Formation (Tithonian-Berria-
sian) and is itself overlain by the Pantano de la Cuerda del Pozo
Formation (middle Barremian). The charophyte association found
in the Golmayo Formation is characteristic of the Triqueta Biozone
and Triqueta Sub-Biozone, which is considered to be late Hau-
terivian to early Barremian in age (Martín-Closas and Alonso, 1998).
3. Systematic palaeontology
Saurischia Seeley, 1887
Sauropoda Marsh, 1878
Neosauropoda Bonaparte, 1986
Titanosauriformes Salgado, Coria & Calvo, 1997a
Brachiosauridae Riggs, 1904
Soriatitan gen. nov.
Etymology. From the Spanish province Soria where the dinosaur
was found and from the Greek “titan” meaning giant in Greek
mythology.
Type species: Soriatitan golmayensis sp. nov.
Diagnosis. As for the type, and only known species.
Soriatitan golmayensis sp. nov.
Figs. 2e10; Table 1
Etymology. In honor of the Golmayo village where the Zorralbo I site
is located.
Site and horizon. Zorralbo I (Golmayo, Soria, Spain) is situated in the
Golmayo Formation (depositional sequence III of Clemente and
Pe
rez-Arlucea [1993] and Mas et al., [1993], and the K1.4
sequence of Mas and Salas [2002]) in the eastern Cameros Basin,
NW Iberian Range. The site is late Hauterivian-early Barremian
(Triqueta-Triqueta charophyte Biozone [Martín-Closas and Alonso,
1998]) in age.
Holotype. The sauropod material recovered at Zorralbo I consists of
cranial and postcranial material (deposited at the Museo Numan-
tino de Soria) (MNS 2001/122), including: teeth (MNS 2001/122),
three dorsal vertebrae (MNS 2001/122.741, MNS 2001/122.743,
MNS 2001/122.745), three fragments of the sacrum (MNS 2001/
122.700, MNS 2001/122.701, MNS 2001/122.703), five caudal
vertebrae (MNS 2001/122.696, MNS 2001/122.1237, MNS 2001/
122.1238, MNS 2001/122.1236, MNS 2001/122.695) and two caudal
vertebral spines (MNS 2001/122.799, MNS 2001/122.793), frag-
ments of ribs (MNS 2001/122.784), one anterior chevron (MNS
2001/122.132), the right humerus (MNS 2001/122.613), the right
radius (MNS 2001/122.1056), the right ulna? (MNS 2001/122.405),
the left and right ilium (MNS 2001/122.1232, MNS 2001/122.1233),
one pubis? (MNS 2001/122.586), the left and right ischium (MNS
2001/122.1240, MNS 2001/122.285), and the left femur (MNS 2001/
122.611).
Diagnosis. Soriatitan golmayensis gen. et sp. nov. is characterized by
eight autapomorphies: i) the caudal ribs develop a pointed anterior
ridge right in the middle; ii) posterior caudal vertebra with large
postzygapophyses: the length of the postzygapophyses is longer
than the anteroposterior length of the spine; iii) presence of a
lateral fossa in every postzygapophyses in the posterior caudal
vertebra; iv) a large, ventral, rectangular ridge located below the
preacetabular process of the ilium; v) ischiadic peduncle in the
ilium three times longer lateromedially than anteroposteriorly; vi)
the same length of the pubic peduncle and the distal shaft in the
ischia; vii) two lateral ridges in the lateral area of the deltopectoral
crest; viii) and two ridges in the medial projection of the humerus
head.
4. Description and comparison
Tooth. An isolated tooth (MNS 2001/122) was discovered along
with the postcranial material. This tooth is semi-spatulate or semi-
peglike (Fig. 2). The root is circular in cross-section. Its crown is a
compressed cone with no constriction with respect to the root, D-
shaped cross-section, with a flat-to-gently concave lingual face and
a strongly convex labial face. Its long axis is skewed through 45

toward the apical end. The crown has an elliptical wear facet on the
labial face just at the apical position. The crown lacks ornamenta-
tion in general, although the lingual crown face shows some slight
apicobasal lines parallel. The mesial and distal edges are sharp,
without denticles. The slenderness index value, which is the length
of the tooth crown divided by its maximum mesiodistal width
(Upchurch, 1998), is 2.3. The general morphology and wear facets
differ from those seen in basal eusauropods and neosauropods. For
example, in Turiasaurus and Camarasaurus, overlapping, inter-
locking dentitions meet in a one-to-two fashion to produce
V-shaped wear facets (Madsen et al., 1995; Royo-Torres et al., 2006;
Chure et al., 2010; Royo-Torres and Upchurch, 2012; Mocho et al.,
2017). The crown of MNS 2001/122 is more compressed than the
tooth of Oplosaurus armatus (NHMUK R964 and CPT-678) (Martill
and Naish, 2001; Royo-Torres and Cobos, 2007) and that of Sau-
roposeidon (¼Paluxysaurus, Rose, 2007) (D'Emic, 2012). The tooth
has similar morphology to the teeth of Astrodon Leidy 1865
(Carpenter and Tidwell, 2005) and the upper teeth of Abydosaurus
(Chure et al., 2010). The tooth also shares characters with the teeth
described as “Pleurocoelus valdensis” from the Valanginian-
Barremian from the United Kingdom (NHMUK R3562, R35354,
R647, R647a, R4403-4407, R2528, 26034, R10093-R10100, R.1898,
R.1896 and R.649) (Naish and Martill, 2001; Upchurch et al., 2011;
Royo-Torres pers. obs. 2010 in NHMUK), from the upper
Hauterivian-lower Barremian of Spain (Sanz et al., 1987, 1992; Ruiz-
Omen ~ aca and Canudo, 2005) and from lower Barremian of Portugal
(Lapparent and Zbyszewsky, 1957; Antunes and Mateus, 2003).
Dorsal vertebrae. Three dorsal centra are available (MNS 2001/
122.7143, MNS 2001/122.741, MNS 2001/122.745). In general these
are poorly preserved. MNS 2001/122.741 is the longest of the three
(Fig. 3); it is an anterior dorsal centrum, whereas the other two are
probably middle or posterior dorsals. All three are slightly opis-
thocoelous. The centra have a subcircular cross section. The ventral
surface is concave anteroposteriorly and gently convex trans-
versally. There is a large, deep, oval pleurocoel on the sides of the
centra, set within associated fosse. The caudal ends of the pleuro-
coels are acute in MNS 2001/122.745, as in the anterior dorsal
 R. Royo-Torres et al. / Cretaceous Research 80 (2017) 38e55 41

Fig. 2. Tooth (MNS 2001/122) of Soriatitan golmayensis: A, in labial view; B, in mesial view; C, in distal view; D, and in lingual view; W (wear facet).

Fig. 3. Dorsal vertebra (MNS 2001/122.745) of Soriatitan golmayensis: in lateral view, A; in dorsal view, B; anterior view, C; and a dorsal pleurocoel in detail with internal laminae, D.

vertebrae of macronarians (Upchurch et al., 2004). The general
features and shape of the dorsal centra of Soriatitan are similar to
the sauropod dorsal vertebrae from the Lower Cretaceous of
Europe: Xenoposeidon (Taylor and Naish, 2007), Eucamerotus
(Blows, 1995) and Tastavinsaurus (Canudo et al., 2008) and of North
America: Cedarosaurus (Tidwell et al., 1999) and Sauroposeidon
(¼Paluxysaurus Rose, 2007). The suture of the neural arch partially
covers the dorsal surface of the centrum and it is different to that of
Xenoposeidon (Taylor and Naish, 2007; Upchurch et al., 2011). The
internal bone texture is spongy, with large, open, “camellate” in-
ternal cells similar to those seen in Giraffatitan and Brachiosaurus
(Upchurch, 1998; Wilson, 2002; Taylor, 2009) and different to the
honeycomb pattern from Eucamerotus (Royo-Torres pers. obs. 2010
in NHMUK) and Tastavinsaurus (Royo-Torres et al., 2012) and the
42 R. Royo-Torres et al. / Cretaceous Research 80 (2017) 38e55
Fig. 4. Anterior caudal spine (MNS 2001/122.799) of Soriatitan golmayensis: in posterior view, A; left lateral view, B; anterior view, C; and right lateral view, D.
somphospondylous structure of titanosaurs (Wilson, 2002; D'Emic,
2012; Mannion et al., 2013).
Sacrum. Three fragments of the sacrum are available (MNS 2001/
122.700, MNS 2001/122.701, MNS 2001/122.703). The sacrum re-
tains two fused vertebral centra and a sacral rib. The centra lack a
ventral groove or keel and their surface is convex. The rib is short
and robust with an S-shaped blade. A small lateral fossa can be seen
in some of the centra. No further features can be observed.
Caudal vertebrae. Five caudal vertebrae are available (MNS 2001/
122.696, MNS 2001/122.1237, MNS 2001/122.1238, MNS 2001/
122.1236, MNS 2001/122.695) and two caudal vertebral spines
(MNS 2001/122.799, MNS 2001/122.793) are preserved. MNS 2001/
122.696 is the most anterior caudal vertebra and, together with the
proximal caudal spine MNS 2001/122.799 displays several potential
diagnostic features (Figs. 4 and 5). The most proximal caudal
centrum is axially short compared to its diameter and is com-
pressed dorsoventrally. The caudal centra MNS 2001/122.1237,
MNS 2001/122.1238, MNS 2001/122.1236 are middle centra while
MNS 2001/122.695 represents a posterior vertebra (Fig. 5). The
series becomes more elongated axially. The centra have a plesio-
morphic amphicoleous articulation, as do all basal eusauropods and
macronarian sauropods (Upchurch et al., 2004). The proximal
caudal vertebrae possess deeply concave cranial faces, while the
posterior faces are between flat and concave. This condition is
similar to that of Cedarosaurus weiskopfae (Tidwell et al., 1999) but
different to those of Giraffatitan (Taylor, 2009), Sonorasaurus
(Ratkevitch, 1998) and Astrodon nanus (Tidwell et al., 1999). In the
proximal caudals the ventral surface of each centrum has a narrow
groove extending proximo-distally, and becoming a more trans-
versal convex surface in the middle and distal caudals. The prox-
imal and middle caudals have an elongated, oval depression just
below the caudal rib. This character is different from that of the true
pleurocoel of Diplodocus and Barosaurus (Upchurch et al., 2004), but
resembles that of the Cedarosaurus (Tidwell et al., 1999), Pelor-
osaurus (Upchurch et al., 2011), Venenosaurus (D'Emic, 2012),
Giraffatitan (Royo-Torres pers. obs. in Berlin, MB.R. 2921.1, MB.R.
2921.2, MB.R. 2921.3; Wedel and Taylor, 2013) and Padillasaurus
(Carballido et al., 2015). The same condition is also seen in the first
caudal of Tastavinsaurus (Canudo et al., 2008; Royo-Torres, 2009).
The arch is located strongly forward so that it occupies the cranial
half of each centrum, a condition of Titanosauriformes (Salgado
et al., 1997a). The posterior caudal prezygapophyses project
beyond the anterior edge of the centrum and are horizontal such as
in Tastavinsaurus and different to those of Giraffatitan. The posterior
caudal postzygapophyses also project posteriorly. Both, pre-
zygapophyses and postzygapophyses are parallel to center. There is
a potential autopomorphie in the posterior caudal vertebra with
large postzygapophyses: the length of the postzygapophyses is
longer than the anteroposterior length of the spine. And a new
potential diagnostic character fossa occurs lateral to every post-
zygapophyses offering for this taxa. The caudal neural spines are
short, single, and rise from the neural arch anterodorsally and then
project strongly backward. This latter feature MNS 2001/122 is
different to that in Cedarosaurus. Mid-caudal neural spines project
anterodorsally from the neural arch arch as in the anterior to mid-
caudals of Cedarosaurus, Venenosaurus (Tidwell et al., 1999; Tidwell
et al., 2001; Rose, 2007; Woodruff, 2012; D'Emic, 2012) and Tasta-
vinsaurus (Canudo et al., 2008; Royo-Torres, 2009). It is a potential
diagnostic character for the Laurasiformes (Royo-Torres et al.,
2012). Spinoprezygapofisial laminae are present only in the base
of the spine. The postzygapophyses in the most anterior spine
preserved are inclined 45
with a triangular hyposphene between
them. It is smaller than the postzygapophyses, and both, hypo-
sphene and postzygapophyses are planar and meet at an angle of
about 90
. This latter character is similar to that found in Astro-
phocaudia (Tidwell et al., 1999; D'Emic, 2013). Posteriorly in the
 R. Royo-Torres et al. / Cretaceous Research 80 (2017) 38e55 43

Fig. 5. Anterior chevron and caudal vertebrae of Soriatitan golmayensis: anterior chevron (MNS 2001/122,132) in proximal view, A; anterior caudal vertebra (MNS 2001/122.696) in
proximal view, B; dorsal view, C; lateral view, D; distal view, E; and ventral view, F; middle-anterior caudal vertebra (MNS 2001/122.1237) in lateral view, G; posterior caudal
vertebra (MNS 2001/122.1238) in right lateral view, H; proximal view, I; dorsal view, J; ventral view, K; and left lateral view, L.

caudal vertebral series, the hyposphene is represented by a single
crest, as in other sauropods (Upchurch, 1998). Finally, their dorsal
ends terminate in a rugose club similar to that seen in several
sauropods: Losillasaurus (Casanovas et al., 2001), Camarasaurus
(Ikejiri et al., 2005), Sauroposeidon (¼Paluxysaurus, Rose, 2007) and
Tastavinsaurus, but without the three rounded bumps characteristic
of the latter taxon (Canudo et al., 2008; Royo-Torres, 2009).
Chevrons. One Y-shaped chevron (Fig. 5A) was recovered (MNS
2001/122,132). The proximal end is formed from two laterally
compressed but dorsally open rami, different to those seen in basal
eusauropods and diplodocids (Wilson, 2002; Upchurch et al.,
2004). The distal blade is similar in length to the dorsal rami but
more robust. The length of the hemal canal is shorter than in the
distal blade (44%), this character differing from that of titanosaurs
chevrons (Wilson, 2002; Royo-Torres, 2009) with 50% of the length
of the hemal canal (Curry Rogers, 2005).
A value over 40% is a derived character state of titanosaur-
iformes and it is a feature shared by Soriatitan, Tastavinsaurus and
Giraffatitan (Upchurch et al., 2004; Royo-Torres, 2009).
Forelimb. The humerus (MNS 2001/122.613) is a long straight
bone that is medially inclined (Fig. 6). The proximal and distal ends
are almost the same width and are not particularly expanded with
respect to the diaphysis. The proximal end is straight, as in Giraf-
fatitan (Janensch, 1961) and Cedarosaurus (Tidwell et al., 1999), and
the diaphysis is elliptical in section; the distal end is straight at the
cranial border, with two smooth condyles on the caudal surface. On
the anterior surface, the two condyles are separated by a small
vertical groove. The deltopectoral crest is prominent and
perpendicular to the diaphysis (Fig. 6D). It begins below the head of
the humerus, and extends down to the midshaft. On the postero-
lateral surface of the deltopectoral crest there is a double ridge and
a groove. In accordance with Borsuk-Bialynicka (1977), this area has
a scar from the muscular scapulohumeralis anterior. The features,
general morphology and proportions are similar to the humerus
(DMNH 39045) of the Cedarosaurus weiskopfae. They share a
proximodistally straight lateral margin between the proximal head
and the shaft of the humerus, the latter being a new potential
diagnostic character for the Laurasiformes (Fig. 7). The humerus is
also similar to that of Giraffatitan differing in that the deltopectoral
crest in the present specimen is at right angles to the cranial surface
of the diaphysis while in Giraffatitan the angle is acute and the
lateral margin in the shaft is concave like in all other sauropods. The
length of the left femur is probably almost the same as that of the
humerus, although it is impossible to be sure because the proximal
end of the femur is lost. The ulna (MNS 2001/122.405) is seriously
damaged and no features can be observed. The radius (MNS 2001/
122.1056) is only seriously damaged at its proximal and distal ends.
They are slightly expanded with respect to the diaphysis (Fig. 8) and
in general it is compressed anteroposteriorly. The dorsoventral
developmental radius rotates in a manner similar to that seen in
Tastavinsaurus and Cedarosaurus (Tidwell et al., 1999). The diag-
nostic ridges seen in the holotype of C. weiskopfae (Tidwell et al.,
1999; D'Emic, 2012) are not developed in Soriatitan, but the bone
is quite slender, similar to that of Cedarosaurus (Tidwell et al., 1999)
and Tastavinsaurus (Royo-Torres et al., 2012) in contrast to that of
Camarasaurus and Giraffatitan.
44 R. Royo-Torres et al. / Cretaceous Research 80 (2017) 38e55
Fig. 6. Right humerus (MNS 2001/122.613) of Soriatitan golmayensis: in posterior view, A; in ventral view, B; and in anterior view, C; the deltopectoral crest in lateral view, D.
Pelvic girdle. Partial left and right ilia are available (MNS 2001/
122.1232, MNS 2001/122.1233). Only the lower parts of these bones
can be described, i.e., the ventral surface of the preacetabular
process, the pubic process and the ischial peduncle (Fig. 9). The
pubic process is prominent and projects anteroventrally; the ischial
peduncle is reduced, as in the Sauropoda clade (Salgado et al.,
1997a; Wilson and Sereno, 1998) and the ischiadic peduncle is
three times longer lateromedially than anteroposteriorly which is a
potencial autopomorphie for this taxa. In general the ilium is ori-
ented in a parasagittal plane with the preacetabular process
directed anteriorly and laterally, which is a character of the Neo-
sauropoda clade (Wilson and Sereno, 1998). The main character of
both ilia is the presence of a large, rectangular ventral ridge below
the preacetabular process and anterior to the pubic process. This is
the first time that this character has been described in a sauropod;
it is therefore a potential diagnostic character for Soriatitan gol-
mayensis. This ridge is not present in other basal Titanosauriformes,
such as Giraffatitan (Taylor, 2009), Brachiosaurus (Riggs, 1903;
Janensch, 1961), Tastavinsaurus (Canudo et al., 2008) or Brontome-
rus (Taylor et al., 2011). The pubis? (MNS 2001/122.586) is severely
damaged and no characters can be described. Both ischia from
Soriatitan are, however, well preserved (Fig. 10), including a
complete left bone (MNS 2001/122.1240) and a partial right one
(MNS 2001/122.285). The ischia are slender bones with three parts:
a short iliac peduncle, a rectangular pubic blade, and an equi-
dimensional ischiadic blade. The acetabulum is smooth and short.
The pubic blade is short anteroposteriorly and the ischial blade
directed ventrally. The ischiadic blade is relatively narrow medi-
olaterally, with no expanded distal end. There is a vertical ridge on
the lateral surface near the posterior boundary. The general
morphology of the Soriatitan is quite similar to that of Cedarosaurus,
Tastavinsaurus, Venenosaurus and Giraffatitan. The ratio between
the length of the pubic peduncle and the distal ischiadic blade is 1:1
for both the right and left ischia, which is characteristic of this taxa,
but different to that of other basal Titanosauriformes such as Ven-
enosaurus, Giraffatitan, Tastavinsaurus (see Table 4.52 in Royo-
Torres, 2009) and Sauroposeidon (¼Paluxysaurus Rose, 2007) in
which the ischial blade is larger than the pubic blade. This feature is
not known in Cedarosaurus (Tidwell et al., 1999), making it another
potential diagnostic character for Soriatitan golmayensis.
Hindlimb. The femur (MNS 2001/122.611) is straight, with the
femoral shaft compressed anteroposteriorly and elliptical in cross
section (Fig. 10). The fourth trochanter is reduced to a small,
rounded groove, a possible synapomorphie for Cedarosaurus,
 R. Royo-Torres et al. / Cretaceous Research 80 (2017) 38e55 45

Fig. 7. Comparison between humerus of Macronaria. Humerus with a concave lateral margin in the shaft: Tehuelchesaurus, A (Carballido et al., 2011); Giraffatitan, B (Janensch, 1961);
Chubutisaurus, C (McIntosh, 1990); Alamosaurus, D (McIntosh, 1990); Ligabuesaurus, E (Bonaparte et al., 2006); Fukuititan, F (Azuma and Shibata, 2010); Vouivria (Mannion et al.,
2017), G. Humerus with a straight lateral margin in the shaft: Angolatitan, H (Mateus et al., 2011); Soriatitan, I (this study); and Cedarosaurus, J (Tidwell et al., 1999), scale bar
10 cm.

Abydosaurus and Venenosaurus clade (D'Emic, 2012) rather than a
reduced rounded ridge as is usual in most sauropods (Upchurch
et al., 2004). It is situated on the posteromedial margin in the
more proximal area of the femur, as in the holotype of Cedar-
osaurus. The distal end of the femur has condyles that articulate
with the fibula and the tibia, and which are separated by a distinct
large intercondylar groove. The condyles show good development
anteriorly and posteriorly, unusually lateromedially narrow such as
in Cedarosaurus (Tidwell et al., 1999), but different to those of
Brachiosaurus and Giraffatitan which are compressed ante-
roposteriorly (Royo-Torres, 2009). Two important characteristics
can be described in this bone. The first is the presence of a later-
odistal ridge in the distal shaft (Fig. 10). It is 15 cm long and is
developed dorsoventrally featuring a groove in the anterior view. A
small rounded ridge has been seen in similar area in Tastavinsaurus
and Lourinhasaurus (Royo-Torres pers. obs. 2013; Mocho et al.,
2014) although in both of these examples they are very smooth.
In general, sauropods lack a ridge in this area of the femur. The well
developed ridge in Soriatitan is due to taphonomic deformation.
The second character is the fact that the epicondyle is located in a
clear lateral position rather than in lateroposterior point in the
fibular condyle as is usual in all other sauropods (Upchurch et al.,
2004). The epicondyle is just below the lateral supracondylar
ridge described above.
5. Phylogenetic analysis
In order to further understand the phylogenetic position and
relationships of Soriatitan we have employed a specific analysis
of the Titanosauriformes based on the D'Emic et al. (2016) matrix
with some modifications for Tastavinsaurus (Royo-Torres et al.,
2012, 2014), as well as the last specific analysis of the Titano-
sauriformes based on the Mannion et al., 2017 matrix. Data
matrices were analyzed using TNT 1.1 (Goloboff et al., 2003) in
order to find the most parsimonious trees (MPTs). For the D'Emic
et al. (2016) analysis, we implemented a heuristic tree search by
performing 1000 replications of Wagner trees (using random
addition sequences) followed by tree bisection reconnection
(TBR) as a swapping algorithm, and saving 10 trees per replicate.
In order to test the support of the phylogenies, the Bremer
support and the bootstrap (absolute frequencies based on 10000
replicates) values were obtained (Fig. 11). For the Mannion et al.
(2017) analysis the pruned data matrix was analyzed using the
‘Stabilize Consensus’ option in the ‘New Technology Search’ in
TNT (Goloboff et al., 2003). Searches were carried out using
sectorial searches, drift, and tree fusing, with the consensus
stabilized five times prior to using the resultant trees as the
starting trees for a ‘Traditional Search’ using tree Bisection-
Reconection (Fig. 12).
46 R. Royo-Torres et al. / Cretaceous Research 80 (2017) 38e55

Fig. 8. The right ulna? (MNS 2001/122.405) in anterior view, AeB; and the right radius (MNS 2001/122.1056) in medial view, CeD of Soriatitan golmayensis.

Fig. 9. Pelvic girdle bones of Soriatitan golmayensis: left ischia (MNS 2001/122.1240) in lateral view, A; right ilium (MNS 2001/122.1232) in lateral view, B; left ilium (MNS 2001/
122.1233) in lateral view, C; the narrow indicates an autapomorphie of Soriatitan golmayensis: a rectangular ridge bellow the pubic process and below the preacetabular process.

6. Results resulting in 31 taxa and 122 morphological characters was used.

Characters 14, 18, 32, 69, 81 and 88 were treated as ordered in the
D'Emic et al., 2016 data set: The updated matrix of D'Emic et al. D'Emic et al. dataset (D'Emic, 2012; D'Emic et al., 2016). The analysis
(2016) including the new scoring for Tastavinsaurus and Soriatitan yielded 5 MPTs of length 220 placing Soriatitan is in
 R. Royo-Torres et al. / Cretaceous Research 80 (2017) 38e55
Fig. 10. Left femur (MNS 2001/122.611) in anterior view, A; ventral view, B; lateral view (only distal femur), C; and posterior view, D.
Brachiosaruidae along with Abydosaurus, Venenosaurus, Tasta-
vinsaurus and Cedarosaurus. The topology (Fig. 11) is similar to that
obtained by D'Emic (2012) and D'Emic et al. (2016).
Mannion et al., 2017 data set: The matrix of Mannion et al.
(2017) including the Tastavinsaurus correction and Soriatitan
resulting in 69 taxa and 415 morphological characters was used.
Characters 11, 14, 15, 27, 40, 51, 104, 122, 147, 148, 177, 195, 205 and
259 were treated as ordered in the Mannion et al. dataset, nine
unstable and highly incomplete taxa were excluded a priori
(Astrophocaudia, Australodocus, Brontomerus, Fukuititan, Fusuisau-
rus, Liubangosaurus, Malarguesaurus, Mongolosaurus and Tenda-
guria). The analysis yielded 20 MPTs of length 1669, where
Soriatitan is a Brachiosauridae within a clade with Cedarosaurus and
Abydosaurus. The topology (Fig. 12) is similar to that obtained by
Mannion et al. (2017). When three taxa are excluded a posteriori
(Cloverly titanosauriform, Chubutisaurus and Angolatitan) Tasta-
vinsaurus is included in the same clade of Soriatitan, with 10 MPTs
of length 1664 (Fig. 13). The Laurasiformes would be inside of
Brachiosauridae along with Cedarosaurus, Abydosaurus, Soriatitan
and Tastavinsaurus.
7. Discussion
Soriatitan had been initially described as a Brachiosauridae indet
(Fuentes Vidarte et al., 2005), but was afterwards related to the
Titanosauriformes clade (Royo-Torres et al., 2009).
From the available Soriatitan material it is possible to observe
the absence of synapomorphies of Diplodocoidea, one of the most
47
important evolutionary lineages within Neosauropoda (Upchurch,
1995; Wilson & Sereno, 1998; Wilson, 2002; Upchurch et al.,
2004; Whitlock, 2011; Carballido & Sander, 2014; Tschopp et al.,
2015), as well as, of Rebachisauridae, the only diplodocoid group
that seems to be represented in the Iberian Lower Cretaceous
(Torcida Ferna
ndez-Baldor et al., 2011). On the other hand, Sor-
iatitan possesses a suite of characters supporting its inclusion in
Macronaria: Firstly, anterior caudal chevrons with open proximal
articulations (Upchurch, 1995; Wilson & Sereno, 1998). The absence
of bridged anterior chevrons was considered a synapomorphy of
the Macronaria clade (Upchurch, 1995; Wilson & Sereno, 1998;
Upchurch et al., 2004; Carballido et al., 2011) or Camarasaur-
omorpha (Salgado et al., 1997a). Secondly, Soriatitan possesses an
ischium with a dorsoventrally long pubic peduncle, similar to
Camarasaurus and Titanosauriformes (Salgado et al., 1997a; Wilson
& Sereno, 1998; Wilson, 2002). Finally, the ischial distal shafts are
nearly coplanar (Wilson & Sereno, 1998; Wilson, 2002) which is
considered a diagnostic character of the Macronaria clade (Wilson
& Sereno, 1998) or Camarasauromorpha (Salgado et al., 1997a).
Soriatitan also displays several characters that have traditionally
been optimized as supporting a more derived placement within
Titanosauriformes. It has tooth crowns that show no overlap
(Wilson, 2002; Upchurch et al., 2004; D'Emic, 2012; Mannion et al.,
2013). It has a spongy camellate presacral bone texture (Wilson,
2002) and the caudal vertebra arch occupies the cranial half of
each centrum (Salgado et al., 1997a), this being a synapomorphy of
Titanosauriformes (Salgado et al., 1997a; Upchurch, 1998). However
some eusauropod non-neosauropods such as Cetiosaurus also have
48 R. Royo-Torres et al. / Cretaceous Research 80 (2017) 38e55

Table 1
Measurements of the bones of Soriatitan golmayensis.

Element Feature measured Measurements (mm)

Tooth Length of crown from apical tip to base 18

2001/122 Labiolingual width of the crown at its base 6
Mesiodistal width of the crown at its base 6
Labiolingual width of the crown at the point where it is widest mesiodistally 7
Mesiodistal width of the crown at the point where it is widest mesiodistally 7
Dorsal vertebra Height of posterior surface of the centrum 130
2001/122.7143 Length of centrum 150ca
Width of centrum across its posterior surface 150
Anterior caudal vertebra Height of the anterior articular surface of the centrum 126
2001/122.696 Width of centrum across its anterior surface 211
Height of the posterior articular surface of the centrum 113
Width of centrum across its posterior surface 208
Length of the centrum 102
Middle caudal vertebra Height of the anterior articular surface of the centrum /
2001/122.1238 Width of centrum across its anterior surface 113
Height of the posterior articular surface of the centrum 111
Width of centrum across its posterior surface 122
Length of the centrum 122
Posterior caudal vertebra Height of the anterior articular surface of the centrum 100
2001/122.784 Width of centrum across its anterior surface 95
Height of the posterior articular surface of the centrum 92
Width of centrum across its posterior surface 92
Length of the centrum 145
Humerus Dorsovental length 1250
2001/122.613 Transverse width of proximal end 370
Anteroposterior width of proximal end 130
Distance from the proximal end to the lateral margin of the deltopectoral crest 370
Anterior projection of the deltopectoral crest above the anterior surface of the shaft 140
Minimum mid-shaft circumference 170 and 60
Transverse width of the distal end 340
Maximum anteroposterior width of the distal end 100
Radius Dorsovental length 611ca
2001/122.1056
Ulna? Dorsovental length 630ca
2001/122.405
Ischium Total length 750
2001/122.1240 Length from the most dorsal point in the pubic branch to distal end 740
Length from the most ventral point in the pubic branch to distal end 390
Anteroposterior width of iliac symphysis 110
Mediolateral width of iliac symphysis 60
Max. width of the distal end 11
Dorsoventral length for pubic branch from acetabulum to distal end 390
Anteroposterior length of the pubic branch 190
Length from the most anterior point in the iliac symphysis to the point between 178
the acetabulum and pubic branch
Length of the acetabulum in the ischia 140
Femur Total length 1020ca
2001/122.611 Distal anteroposterior width 220
Distal mediolateral width 260
Minimum anteroposterior width of the shaft 78
Width perpendicular to the minimum anteroposterior width of the shaft 170
Length of 4th trochanter 140

the derived character in some caudal vertebrae (Upchurch &
Martin, 2002, 2003; Royo-Torres and Upchurch, 2012). Soriatitan
has an anterior and middle caudal vertebrae with shallow fosse in
the lateral face of the centrum (D'Emic, 2012; Mannion et al., 2013;
Carballido and Sander, 2014), it also has a gracile humerus with a
length/midshaft width ratio of more than 7.5 (D'Emic, 2012). Sor-
iatitan also possesses an ischium with a raised tubercle on the
dorsolateral face without an associated groove (Wilson, 2002;
D'Emic, 2012). This feature has a wide distribution in Titanosaur-
iformes being present in the basal titanosauriforms Giraffatitan,
Venerosaurus and Sauroposeidon (D'Emic, 2012) but it is also seen in
titanosaurs as Diamantinasaurus and Huabeisaurus (D'Emic, 2013;
Poropat et al., 2015b). In the Iberian Peninsula, this condition is
present in the Late Jurassic Lusotitan atalaiensis (Mannion et al.,
2013; Mocho et al., 2016), Galveosaurus herreroi (Barco, 2009),
and in the Early Cretaceous, it is present in Tastavinsaurus sanzi
(Canudo et al., 2008; Royo-Torres et al., 2012).
The present work compares this specimen with the titano-
sauriform sauropod dinosaurs and the sauropods described in the
Lower Cretaceous in Europe and North America. Soriatitan has
tooth crowns that show no overlap and a compressed cone similar
to the titanosauriform teeth of “Pleurocoelus valdensis” from the
Barremian-Aptian of Europe (United Kingdom and in Iberian
Peninsula) (Sanz et al., 1987; Naish and Martill, 2001; Ruiz-
Omen ~ aca and Canudo, 2005; Upchurch et al., 2011).
The proximal caudal vertebrae (excluding the first) of Soriatitan
have a concave proximal face and a flat-convex distal surface with a
small depression in the middle, similar to Cedarosaurus, Tasta-
vinsaurus and Venenosaurus. This condition is shared with a wider
number of sauropods, i.e., it might exist in Wintonotitan (Hocknull
et al., 2009; Poropat et al., 2015a) and also in Andesaurus; several
authors indicate this genus to lack a true procoelous condition
(Calvo and Bonaparte, 1991; Upchurch, 1998). Also the caudal
vertebrae in Soriatitan (MNS 2001/122) are compressed
 R. Royo-Torres et al. / Cretaceous Research 80 (2017) 38e55 49

Fig. 11. Strict consensus tree of the MPTs produced by cladistic analysis using the D'Emic et al. (2016) data set with Soriatitan golmayensis. Bootstrap values are shown as percentages
(nodes lacking percentages have bootstrap values of less than 50%). Bremer supports are shown in square brackets.

dorsoventrally, similar to those of Brachiosaurus, Pelorosaurus,
Cedarosaurus and Lusotitan. Anterior caudals share the lateral
pneumatic fosse with brachiosaurid titanosauriformes (Royo-
Torres, 2009; D'Emic, 2012; Mannion et al., 2013) and also with
some titanosaurs such as Savannasaurus (Poropat et al., 2016).
Soriatitan has a well-developed hyposphene, a character shared
with Astrophocaudia (SMU 61732) from early Albian (D'Emic, 2013)
and different to the hyposphenal ridge seen between of the post-
zygapophisis in several sauropods (Upchurch, 1998) including
Venenosaurus (Tidwell et al., 2001), Cedarosaurus (Tidwell et al.,
1999) and Tastavinsaurus (Canudo et al., 2008). Soriatitan also
shares a humerus with a “squared” proximolateral corner (Wilson,
2002; D'Emic, 2012) with the Titanosauriformes. Anterior to mid-
caudal neural spines project anterodorsally from the neural arch.
This has been considered a potential diagnostic character for the
Laurasiformes clade (Royo-Torres, 2009; Royo-Torres et al., 2012) as
seen in Cedarosaurus, Venenosaurus (D'Emic, 2012) and Tasta-
vinsaurus, but different in to that Pelorosaurus (Mantell, 1850;
Upchurch et al., 2011) and Giraffatitan (D'Emic, 2012). This last
derived character is also shared with several titanosaurs such as
Epachthosaurus (Martínez et al., 2004) and Aeolosaurus (Salgado
et al., 1997b). The ischium has an ischial blade orientated poster-
oventrally as in Giraffatitan and Venenosaurus, but the ratio be-
tween the pubic pedicel and ischial blade is 1:1. This differs to that
recorded for other titanosauriforms such as Giraffatitan, Ven-
enosaurus and Tastavinsaurus. Soriatitan shares a number of fea-
tures with the Cedarosaurus, including anterior caudal vertebrae
with deeply concave proximal faces but flat, concave distal faces; a
deltopectoral crest on the humerus closer to the midshaft (Tidwell
et al., 1999; Upchurch et al., 2004); and a proximodistally straight
lateral margin between the proximal head and the shaft of the
humerus. In addition, Soriatitan shares the following characters
with the Cedarosaurus: the dorsal centra with their elliptical pleu-
rocoels, the caudal vertebrae with their long lateral processes, a
slender radius and the slender morphology of the femur. On the
other hand, Soriatitan lacks the autopomorphies of the radius of
Cedarosaurus and it is different because Soriatitan has a prox-
imodistally shorter spines in the posterior caudal vertebrae.
50 R. Royo-Torres et al. / Cretaceous Research 80 (2017) 38e55

Fig. 12. Strict consensus tree of the MPTs produced by cladistic analysis using the Mannion et al. (2017) data set with Soriatitan golmayensis. Bootstrap values are shown as
percentages (nodes lacking percentages have bootstrap values of less than 50%). Bremer supports are shown in square brackets.

Soriatitan is unlikely to be a somphospondylan sauropod 8. Taxonomic status of Soriatitan golmayensis

because it lacks the internal somphospondylous bone texture of
titanosaurs (Wilson, 2002; Upchurch et al., 2004; D'Emic, 2012;
Mannion et al., 2013), the posterior articular surfaces of middle-
posterior caudal centra are concave rather than convex (Mannion
et al., 2013), and an ilium with pointed and low preacetabular
process (Wilson, 2002; D'Emic, 2012). Also, the lack of a robust 4th
trochanter located near the midline separate Soriatitan of the
somphospondylian sauropod Euhelopus (Wiman, 1929; Wilson &
Upchurch, 2009). Finally, Soriatitan also lacks the following two
characteristics of Titanosauria: anterior and middle caudal verte-
brae with a longitudinal hollow, and the plate-like ischium, with no
emargination distal to the pubic peduncle (Wilson, 2002;
Carballido et al., 2011; D'Emic, 2012; Mannion et al., 2013). We
can conclude in this discussion that Soriatitan is placed between
Titanosauriformes, within Brachiosauridae and outside of
Titanosauria.
Comparisons between Soriatitan and other contemporaneous
europea taxa, such as Tastavinsaurus, Demandasaurus, Eucamer-
otus, Oplosaurus, Pelorosaurus, and North American taxa, such as
Cedarosaurus, Venenosaurus, Abydosaurus, Sauroposeidon and
Brontomerus, reveal numerous differences, some of which have
been noted in the ‘Description and comparisons’ and ‘Discussion’
sections. Eight characters are here considered potential autapo-
morphies for Soriatitan: The caudal ribs develop a pointed anterior
ridge in the middle which is absent in Tastavinsaurus, Cedar-
osaurus or Giraffatitan. Posterior caudal vertebra with large post-
zygapophyses: the length of the postzygapophyses is longer than
the anteroposterior spine length which is absent in other sauro-
pods included the Titanosauriformes i.e Tastavinsaurus. The lateral
fossa in every postzygapophyses and a large ventral rectangular
ridge located below the preacetabular process of the ilium is
 R. Royo-Torres et al. / Cretaceous Research 80 (2017) 38e55 51

Fig. 13. Strict consensus tree of the MPTs produced by cladistic analysis using the Mannion et al. (2017) data set with Soriatitan golmayensis after exclude a posteriori three taxa
(Cloverly titanosauriform, Chubutisaurus and Angolatitan). Bootstrap values are shown as percentages (nodes lacking percentages have bootstrap values of less than 50%). Bremer
supports are shown in square brackets.

described for the first time in sauropods. The ischiadic peduncle in
the ilium is three times longer lateromedially than ante-
roposteriorly which is different to Giraffatitan, Brontomerus and
Tastavinsaurus. The 1:1 ratio between the length of the pubic
peduncle and distal blade in the ischia is different to the ratio of
the basal Titanosauriformes, i.e., in Giraffatitan it is 0.44, in Tas-
tavinsaurus it is between 0.65 and 0.77, while in Venenosaurus it is
0.65 (Royo-Torres, 2009). Two lateral ridges in the lateral area of
the deltopectoral crest and two ridges in the medial projection of
the humerus head differentiate Soriatitan from Giraffatitan and
Cedarosaurus, and the epicondyle of the femur located in a lateral
position is also different to those found in Tastavinsaurus and
Giraffatitan.
According to the analyses in the present paper with the matrices
of D'Emic et al. (2016) and Mannion et al. (2017), Soriatitan is
included within Brachiosaridae, but also in a more inclusive clade
which may be the Laurasiformes. This clade was defined by Royo-
Torres (2009) as a stem-based clade containing sauropods more
closely related to Tastavinsaurus than Saltasaurus and defended by
Royo-Torres et al. (2012, 2014). The main problem for this clade is
the variation of the phylogenetic position of Tastavinsaurus in the
most recent studies: in Macronaria no Titanosauriformes (Royo-
Torres, 2009; Carballido and Sander, 2014), in Titanosauriformes
no Titanosauria (Royo-Torres et al., 2012, 2014) and in Sompho-
spondyli (Canudo et al., 2008; D'Emic, 2012; Mannion et al., 2013,
2017; Poropat et al., 2015a,b). The results depend on the data ma-
trix, on the codification of characters for Tastavinsaurus, and also on
the inclusion in some taxa. For example, with the data matrix of
Mannion et al. (2017) when used without some incomplete OTUs
such as ‘Cloverly titanosauriform’, Angolatitan and Chubutisaurus,
Tastavinsaurus is located in a clade with Venenosaurus, Cedar-
osaurus, Abydosaurus and Soriatitan within Brachiosaridae (Fig. 13).
In this way, the Laurasiformes could be a valid clade, but at the
moment new studies are necessary in order to confirm or reject its
52 R. Royo-Torres et al. / Cretaceous Research 80 (2017) 38e55
validity. Duly, in this paper we prefer classify Soriatitan as a
Brachiosauridae.
9. Brachiosauridae and its palaeobiogeographical
implications during the Early Cretaceous in Europe
The oldest member of Brachiosaruidae is represented by
Vouivria of the middle-late Oxfordian from Europe. Currently, Late
Jurasic members of Brachiosauridae are known from East Africa
with Giraffatitan (Janensch, 1914), western Europe with Lusotitan
(Mannion et al., 2013; Mocho et al., 2016), the USA, with Brachio-
saurus (Riggs, 1903) and with doubts in South America (Rauhut,
2006) (see discussion in Mannion et al., 2017).
After the Jurassic, Early Cretaceous brachiosaurid remains are
known of the USA, of southern Africa (McPhee et al., 2016), and
possibly from Afro-Arabia too (Buffetaut et al., 2006), while Padil-
lasarusu from South America has been considered a sompho-
spondylan (Mannion et al., 2017). The described taxa came only
from Early Cretaceous of the USA: Abydosaurus, Cedarosaurus,
Sonorasaruus and Venenosaurus (D'Emic et al., 2016; Mannion et al.,
2017). And at the moment, no cretaceous sauropod remains from
Europe has been assigned to Brachiosauridae (Mannion et al., 2013;
Royo-Torres et al., 2014; Mannion et al., 2017) so Soriatitan gol-
mayensis is the first and the only brachiosarid described from the
Early Cretaceous of Europe. This evidence discards the hypothesis
of a regional extinction for brachiosarids during the Early Creta-
ceous in Europe.
The presence of similar Early Cretaceous brachiosaurid sauropod
dinosaurs common to North America and Europe (Cedarosaurus and
Soriatitan) is consistent with previous data describing similar
dinosaur fauna, i.e., polacanthid ankylosaur and iguanodontid, for
both continents during the Barremian (Kirkland, 1992; Carpenter
et al., 2002; Tidwell and Carpenter, 2002; Kirkland and Madsen,
2007; Canudo et al., 2009). The general fauna of Golmayo Forma-
tion (Zorralbo I site) is very similar to that of North America in the
Upper Yellow Cat member of the Cedar Mountain Formation
(Kirkland et al., 2016). In addition to Titanosauriformes with
Cedarosaurus, Venenosaurus, Abydosaurus, Tastavinsaurus and Sor-
iatitan, the findings of polacanthid ankylosaurs with Gastonia burgei
(Kirkland, 1998) in Utah, and a Polacanthus sp in Spain (Pereda-
Suberbiola et al., 2007), iguanodontids with Hippodraco scutodens
(McDonald et al., 2010) in Utah, and a Magnamanus soriaensis in
Spain (Fuentes Vidarte et al., 2016), “Hypsilophodonts” and small
dromeosaurid theropods in both formations support the hypothesis
(Carpenter et al., 2002; Fuentes-Vidarte et al., 2005, 2009a; Pereda-
Suberbiola et al., 2007; Kirkland et al., 2016) that a connection could
have existed between North America and Europe at some point
previous to the HauterivianeBarremian (Brikiatis, 2016).
In the Early Cretaceous, North America and Europe were part of
the same palaeobiogeographic unit of Laurasia, which was
composed of small tectonic plates (Hay et al., 1999). According to
the available biogeographic data, these units may have formed
continuous land masses at certain times during the Early Creta-
ceous (Russell, 1993; Smith et al., 1994; Le Loeuff, 1997; Milner et al.,
2000; Kirkland and Madsen, 2007). It is also possible that members
of these taxa could have walked freely between North America,
Greenland, Baltica and Europe (Golonka et al., 2003). Between
Berriasian-Hauterivian the northernmost Atlantic Ocean had not
yet opened up (Golonka et al., 2003), thereby permitting the ex-
change of animals between these two regions prior to the forma-
tion of an Alaskan land bridge (Cifelli et al., 1997; Kirkland et al.,
1997, 1998, 1999; Kirkland and Madsen, 2007). The combined ef-
fect of the movement of the plates in conjunction with constant
changes in sea level during the Early Cretaceous may well have
allowed this (Skelton et al., 2003). The presence of the basal
therizinosauroid Falcarius (Kirkland et al., 2005) in Utah supports
the proposal that the animals may have been tied to Asia through
Europe at some point during the Early Cretaceous (Kirkland and
Madsen, 2007). These data corroborate the hypothesis suggested
in previous studies (Royo-Torres, 2009; Royo-Torres et al., 2012,
2014) that titanosauriformes sauropods were present at least in
Europe and North America in the Early Cretaceous.
10. Conclusions
This study describes a new species of sauropod, Soriatitan gol-
mayensis (MNS 2001/122) from the Golmayo Formation (upper
Hauterivianelower Barremian) located in Zorralbo I site in Gol-
mayo village (Soria, Spain). This sauropod is classified as a Bra-
chiosauridae within Titanosauriformes supported by two different
phylogenetic analyses. Soriatitan shares the presence of anteriorly
deflected anterioremiddle caudal neural spines with Cedarosaurus,
Venenosaurus and Tastavinsaurus, and a proximodistally straight
lateral margin between the proximal head and the humerus shaft
with Cedarosaurus. Eight characters are newly identified and are
considered potential diagnostic criteria for the new possible genus
and species: the caudal ribs develop a pointed anterior ridge in the
middle; posterior caudal vertebra with large postzygapophyses;
presence of a lateral fossa in every postzygapophysis in the poste-
rior caudal vertebra; the presence of a rectangular ventral ridge
cranial to the pubic process and below the preacetabular process in
the ilium; ischiadic peduncle in the ilium three times longer in
lateromedially than anteropospestiorly; the same length of the
pubic peduncle and distal blade in the ischia; two lateral ridges in
the lateral area of the deltopectoral crest; and the presence of the
two ridges in the medial projection of the humerus head. At the
moment, Soriatitan golmayensis is the first and the only brachio-
sarid described from the Early Cretaceous of Europe. The presence
of the Early Cretaceous brachiosarid sauropods common to North
America and Europe support to the presence of a possible bridge
between these plates, prior to the late HauterivianeBarremian.
Acknowledgements
This work was supported by Ministerio de Economía y Com-
~ a under project CGL2013-41295-P, co-
petitividad, Gobierno de Espan
financed with FEDER funds, the Departamento de Innovacio
n,

n y Universidad, the Fondo Social Europeo (FOCONTUR,
Investigacio
Ref. E62), the Departamento de Educacio
n, Cultura y Deporte del
Gobierno de Arago
n and the Fundacio
n Conjunto Paleontolo
gico de
Teruel-Dino
polis. The authors thank the Museo Numantino de Soria

n (exp. 334.06-
for their collaboration and the Junta de Castilla y Leo
So) and a private company M.R. for their economic support in the
excavation. We want to thank the help of Fe
lix Sanz Aldea in the
field, and Dr. Pedro Mocho, Javier Verdú and the staff of the Museo
Numantino for their help in the museum. Dr. James Kirkland sup-
plied us with information about the Cedar Mountain Formation and
Dr. Jeffrey A. Wilson and Dr. Mike D'emic about Cedarosaurus.
Ciar
an Rowe translated and edited the text in English. The authors
finally thank the anonymous reviewers for their attempts to help to
improve the paper.
References
Alonso, A., Mas, R., 1993. Control tecto
nico e influencia del eustatismo en la sed-
imentacio
n del Creta
cico Inferior de la Cuenca de los Cameros. Cuadernos de
Geología Ibe
rica 17, 285e310.
Antunes, M.T., Mateus, O., 2003. Dinosaurs of Portugal. Comptes Rendus Palevol 2,
77e95.
 R. Royo-Torres et al. / Cretaceous Research 80 (2017) 38e55
Azuma, Y., Shibata, M., 2010. Fukuititan nipponensis, a new titanosauriform
sauropod from the Early Cretaceous Tetori Group of Fukui Prefecture, Japan.
Acta Geologica Sinica e English Edition 84 (3), 454e462.
Barco, J.L., 2009. Sistema
tica e implicaciones filogene
ticas y paleobiogeogra
ficas del
sauro
podo Galvesaurus herreroi (Formacio
n Villar del Arzobispo, Galve,
Espan ~ a) (Unpubl. PhD thesis). Universidad de Zaragoza, pp. 1e389.
Blows, W.T., 1995. The early cretaceous Brachiosaurid dinosaurs Ornithopsis and
Eucamerotus from the isle of Wight, England. Paleontology 38, 187e197.
Bonaparte, J.F., 1986. Les dinosaures (Carnosaures, Allosaurides, Sauropodes,
Cetiosaurides) du Jurassique moyen de Cerro Condor (Chubut, Argentine).
Annales de Paleontologie 72, 325e386.
Bonaparte, J.F., Gonza
lez Riga, B., Apesteguía, S., 2006. Ligabuesaurus leanzai gen. et
sp. nov. (Dinosauria, Sauropoda), a new titanosaur from the Lohan Cura For-
mation (Aptian, Lower Cretaceous) of Neuque
n, Patagonia, Argentina. Creta-
ceous Research 27, 364e376.
Borsuk-Bialynicka, M., 1977. A new camarasaurid sauropod Opisthocoelicaudia
skarzynskii, gen. n. sp. from the Upper Cretaceous of Mongolia. Paleontología
Polonica 37, 1e64.
Brikiatis, L., 2016. Late Mesozic North Atlantic land bridges. Earth-Science Reviews
159, 47e57.
Buffetaut, E., Azar, D., Nel, A., Ziade, K., Acra, A., 2006. First nonavian dinosaur from
Lebanon: a brachiosaurid sauropod from the Lower Crtaceous of the Jezzine
District. Die Naturwissenschaften 93, 440e443.
Calvo, J.O., Bonaparte, J.F., 1991. Andesaurus delgadoi gen. et sp. nov. (Saurischia-
Sauropoda) dinosaurio Titanosauridae de la Formacio
n Rio Limay (Albiano-
Cenomaniano), Neuquen, Argentina. Ameghiniana 28 (3e4), 303e310.
Canudo, J.I., Royo-Torres, R., Cuenca-Besco
s, G., 2008. A new sauropod: Tasta-
vinsaurus sanzi gen. et sp. nov. from the Early Cretaceous (Aptian) of Spain.
Journal of Vertebrate Palaeontology 28, 712e731.
Canudo, J.I., Barco, J.L., Pereda-Suberbiola, X., Ruiz-Omen ~ aca, J.I., Salgado, L., Torcida
Ferna
ndez-Baldor, F., Gasulla, J.M., 2009. What Iberian dinosaurs reveal about
the bridge said to exist between Gondwana and Laurasia in the Early Creta-
ceous. Bulletin de la Socie
te Ge
ologique de France 180 (1), 5e11.
Carballido, J.L., Sander, P.M., 2014. Postcranial axial skeleton of Europasaurus holgeri
(Dinosauria, Sauropoda) from Upper Jurassic of Germany: implications for
sauropod ontogeny and phylogenetic relationships of basal Macronaria. Journal
of Systematic Paleontology 12, 335e387.
Carballido, J.L., Rauhut, O.W.M., Pol, D., Salgado, L., 2011. Osteology and phylogeny
relationships of Tehuelchesaurus benitezii (Dinosauria, Sauropoda) from the
Upper Jurassic of Patagonia. Zoological Journal of the Linnean Society 163,
605e662.
Carballido, J.L., Pol, D., Parra Ruge, M.L., Padilla Bernal, S., P
aramo-Fonseca, M.E.,
Etayo-Serna, F., 2015. A new early Cretaceous Brachiosaurid (Dinosauria, Neo-
sauropoda) from Northwestern Gondwana (Villa de Leiva, Colombia). Journal of
Vertebrate Paleontology 35 (5), e980505.
Carpenter, K., Tidwell, V., 2005. Reassessment of the Early Cretaceous Sauropod
Astrodon johnsoni Leidy 1865 (Titanosauriformes). In: Tidwell, V., Carpenter, K.
(Eds.), Thunder-Lizards, the sauropodomorph Dinosaurs. Indiana University
press, Bloomington and Indianapolis, pp. 78e114.
Carpenter, K., DiCroce, T., Gilpin, D., Kinner, B., Sanders, F., Tidwell, V., 2002. Origins
of the Early and “Middle” Cretaceous Dinosaurs of North America: Implications
for Plate Tectonics. Proceedings of the Iternational Symposium on New Con-
cepts in Global Tectonics 289e308.
Casanovas, M.L., Santafe
, J.V., Sanz, J.L., 2001. Losillasaurus giganteus, un nuevo
sauro
podo del tra
nsito Jur
asico-Cret

acico de la cuenca de “Los Serranos”
(Valencia, Espan ~ a). Paleontologia i Evolucio
32e33, 99e122.
Chure, D., Britt, B.B., Whitlock, J.A., Wilson, J.A., 2010. First complete sauropod
dinosaur skull from the Cretaceous of the Americas and the evolution of
sauropod dentition. Naturwissenschaften 97, 379e391.
Cifelli, R.L., Kirkland, J.I., Weil, A., Deinos, A.R., Kowallis, B.J., 1997. High precision
40Ar/39Ar geochronology and the advent of North America's Late Cretaceous
terrestrial fauna. Proceedings National Academy of Science 94, 11163e11167.
Clemente, P., Alonso, A., 1990. Estratigrafía y sedimentología de las facies con-
tinentales del Cret
acico Inferior en el borde meridional de la Cuenca de los
Cameros. Estudios Geolo
gicos 46, 257e276.
Clemente, P., Pe
rez-Arlucea, M., 1993. Depositional architecture of the Cuerda del
Pozo Formation, Lower Cretaceous of the extensional Cameros Basin, notrh-
central Spain. Journal of Sedimentary Petrology 63, 437e452.
Curry Rogers, K., 2005. Titanosauria. In: Curry Rogers, K., Wilson, J.A. (Eds.), The
Sauropods, evolution and paleobiology. University of California Press, Berkeley,
pp. 50e103.
D'Emic, M.D., 2012. The early evolution of titanosauriform sauropod dinosaurs.
Zoological Journal of the Linnean Society 166, 624e671.
D'Emic, M.D., 2013. Revision of the sauropod dinosaurs of the Lower Cretaceous
Trinity Group, southern USA, with the description of a new genus. Journal of
Systematic Palaeontology 11, 707e726.
D'Emic, M.D., Foreman, B.Z., Jud, N.A., 2016. Anatomy, systematics, paleoenvir-
onment, growth, and age of the sauropod dinosaur Sonorasaurus thompsoni
from the Cretaceous of Arizona, USA. Journal of Paleontology 90 (1),
102e132.
Fuentes Vidarte, C., Meijide Calvo, M., Meijide Fuentes, F., Meijide Fuentes, M., 2005.
Fauna de vertebrados del Creta
cico Inferior del Yacimiento de “Zorralbo” en
Golmayo (Soria, Espan ~ a). Revista Espan~ ola de Paleontología nº extraordinario X
83e92.
53
Fuentes Vidarte, C., Meijide Calvo, M., Meijide Fuentes, F., Meijide Fuentes, M., 2008.
Coprolitos de cocodrilo del yacimiento de “Zorralbo I” (Cret
acico inferior) de
Golmayo (Soria, Espan ~ a). XXIV Jornadas de la Sociedad Espan ~ ola de Paleon-
tología. Museo del Jur
asico de Asturias, Colunga, Libro de resúmenes, 29.
Fuentes Vidarte, C., Meijide Calvo, M., Meijide Fuentes, F., Meijide Fuentes, M.,
2009a. A new Iguanodon species (Dinosauria, Ornithischia) from the Lower
Cretaceous of Soria (Spain), 40. Abstracts Darwin-Bernissart Meeting, Brussels.
Fuentes Vidarte, C., Meijide-Calvo, M., Meijide-Fuentes, F., Meijide-Fuentes, M.,
2009b. Skeletal remains of a theropod dinosaur from the Lower Cretaceous of
Soria (Spain). In: Delgado-Buscalioni, A., Fregenal-Martínez, M. (Eds.), Tenth
International Symposium on Mesozoic Terrestrial Ecosystems and Biota. Uni-
versidad Auto
noma de Madrid, p. 145.
Fuentes Vidarte, C., Meijide-Calvo, M., Meijide-Fuentes, F., Meijide-Fuentes, M.,
2016. Un nuevo dinosaurio estiracosterno (Ornithopoda: Ankylopollexia) del
Cret
acico Inferior de Espan ~ a. Spanish Journal of Palaeontology 31 (2), 407e446.
Goloboff, P.A., Farris, J.S., Nixon, K., 2003. TNT: tree analysis using new technology.
Systematic Biology 54, 176e178.
Golonka, J., Bocharova, N.Y., Ford, D., Edrich, M.E., Bednarczyk, J., Wildharber, J.,
2003. Paleogeographic reconstructions and basins development of the Arctic.
Marine and Petroleum Geology 20, 211e248.
Hay, W.H., De Conto, R.M., Wold, C.N., Wilson, K.M., Voigt, S., Schulz, M., Rossby
Wold, A., Dullo, W.-C., Ronov, A.B., Balukhovsky, A.N., So € ding, E., 1999. Alter-
native global Cretaceous paleogeography. Geological Society of America, Special
Paper 332, pp. 1e47.
Hocknull, S.A., White, M.A., Tischler, T.R., Cook, A.G., Calleja, N.D., Sloan, T.,
Elliot, D.A., 2009. New mid-Cretaceous (latest Albian) dinosaurs from Winton,
Queensland, Australia. PLoS One 4 (7), e6190. http://dx.doi.org/10.1371/
journal.pone.0006190.
Ikejiri, T., Tidwell, V., Trexler, D.L., 2005. New adult specimens of Camarasaurus
lentus highlight ontogenetic variation within the species. In: Tidwell, V.,
Carpenter, K. (Eds.), Thunder-Lizards The sauropodomorph Dinosaurs. Indiana
University Press, pp. 154e179.
Janensch, W., 1914. Übersicht über der Wirbeltierfauna der Tendaguru-Schichten
nebst einer kurzen Charakterisierung der neu aufgefuhrten Arten von Sau-
ropoden. Archiv für Biontologie 3, 81e110.
Janensch, W., 1929. Material und Formengehalt der Sauropoden in der Ausbeute der
Tendaguru-Expedition. Palaeontographica 2 (Suppl. 7), 1e34.
Janensch, W., 1961. Die gliedmaszen und gliedmaszengurtel der sauropoden der
Tendaguru-schichten. Palaeontographica 3 (Suppl. 7), 177e235.
Kirkland, J.I., 1992. Dinosaurs define a two-fold Lower Cretaceous zonation of the
Cedar Mountain Formation, central Utah: Geological Society of America Ab-
stracts and Program, 24, p. 22.
Kirkland, J.I., 1998. A polacanthid ankylosaur from the Early Cretaceous of eastern
Utah. In: Lucas, S.G., Kirkland, J.I., Estep, J.W. (Eds.), Lower to Middle Cretaceous
non-marine Cretaceous faunas, 14. New Mexico Museum of Natural History and
Science Bulletin, pp. 271e281.
Kirkland, J.I., Madsen, S.K., 2007. The Lower Cretaceous Cedar Mountain Formation,
eastern Utah: the view up an always interesting learning curve. Geological
Society of America Rocky Mountain Section. Annual Meeting, St. George, Utah,
May 4e6, 2007, pp. 1e108.
Kirkland, J.I., Cifelii, R.L., Elder, W.P., 1997. Land-bridge between Asia and North
Amercia: dating its Latest Albian (Cretaceous) origins and its migration induced
extinctions, 29. Geological Society of America Abstracts and program, p. A462.
Kirkland, J.I., Lucas, S.G., Estep, J.W., 1998. Cretaceous dinosaurs of the Colorado
Plateau. In: Lucas, S.G., Kirkland, J.I., Estep, J.W. (Eds.), Lower to Middle Creta-
ceous non-marine Cretaceous faunas: New Mexico Museum of Natural History
and Science Bulletin 14, pp. 67e89.
Kirkland, J.I., Cifelli, R., Britt, B.B., Burge, D.L., De Courten, F., Eaton, J., Parrish, J.M.,
1999. Distribution of vertebrate faunas in the Cedar Mountain Formation, east-
central Utah. In: Gillette, D. (Ed.), Vertebrate paleontology in Utah: Utah
Geological Survey Miscellaneous Publication, 99e1, pp. 201e217.
Kirkland, J.I., Zanno, L.E., Sampson, S.D., Clark, J.M., DeBlieux, D.D., 2005. A primitive
therizinosauroid dinosaur from the Early Cretaceous of Utah. Nature 435
(7038), 84e87.
Kirkland, J.I., Suarez, M., Suarez, C., Hunt-Foster, R., 2016. The Lower Cretaceous in
east-central Utahdthe Cedar Mountain Formation and its bounding strata.
Geology of the Intermountain West 3, 101e228.
Lapparent, A.F., Zbyszewski, G., 1957. Les Dinosauriens du Portugal. Memoire Ser-
vices Ge
ologiques du Portugal 2, 1e61.
Le Loeuff, J., 1997. Biogeography. In: Currie, P.J., Padian, K. (Eds.), Encyclopedia of
Dinosaurs. Academic Press, San Diego, pp. 51e56.
Mantell, G.A., 1850. On the Pelorosaurus: an undescribed gigantic terrestrial reptile,
whose remains are asociated with those of the Iguanodon and other saurians in
the strata of the Tilgate Forest, in Sussex, 140. Philosophical Transactions of the
Royal Society, London, pp. 379e390.
Madsen, J.H., McIntosh, J.S., Berman, D.S., 1995. Skull and atlas-axis complex of the
Upper Jurassic sauropod Camarasaurus Cope (Reptilia: Saurischia). Bulletin of
the Carnegie Museum of Natural History 31, 1e115.
Mannion, P., Upchurch, P., Barnes, R.N., Mateus, O., 2013. Osteology of the Late
Jurassic Portuguese sauropod dinosaur Lusotitan atalaiensis (Macronaria) and
the evolutionary history of basal titanosauriformes. Zoological Journal of the
Linnean Society 168, 98e206.
Mannion, P., Allain, R., Moine, O., 2017. The earliest known titanosauriform
sauropod dinosaur and the evolution of Brachiosauridae. PeerJ 5, e3217.
54 R. Royo-Torres et al. / Cretaceous Research 80 (2017) 38e55
Marsh, O.C., 1878. Principal characters of American Jurassic Dinosaurs Part I.
American Journal of Science 16 (series 3), 411e416.
Martill, D., Naish, D., 2001. Dinosaurs of the Isle of Wight. The Paleontological As-
sociation, Field Guides to Fossils 10, 1e424.
Martín-Closas, C., Alonso, A., 1998. Estratigrafía y bioestratigrafía (Charophyta) del
Creta
cico Inferior en el sector occidental de la Cuenca de Cameros (Cordillera

rica). Revista de la Sociedad Geolo
Ibe
gica de Espan ~ a 11, 253e269.
Martínez, R.D., Gime
nez, O., Rodriguez, J., Luna, M., Lamanna, M.C., 2004. An arti-
culated specimen of the basal titanosaurrian (Dinosauria: Sauropoda) Epach-
tosaurus sciuttoi from the early Late Cretaceous Bajo Barreal Formation of
Chubut province, Argentina. Journal of Vertebrate Paleontology 24, 107e120.
Mas, J.R., Salas, R., 2002. Lower Cretaceous of the Iberian basin. In: Gibbons, W.,
Moreno, T. (Eds.), The Geology of Spain. The Geological Society, London,
pp. 284e288.
Mas, J.R., Alonso, A., Guimera, J., 1993. Evolucio
n tectonosedimentaria de una
cuenca extensional intraplaca: La cuenca finijur
asica-eocret
acica de Los
Cameros (La Rioja-Soria). Revista de la Sociedad Geolo
gica de Espan ~ a 6,
129e144.
Mateus, O., Jacobs, L., Schulp, A.S., Polcyn, M.J., Tavares, T.S., Buta Neto, A.,
Morais, M.L., Antunes, M.T., 2011. Angolatitan adamastor, a new sauropod
dinosaur and the first record from Angola. Anais da Academia Brasileira de
^ncias 83, 1e13.
Cie
McDonald, A.T., Kirkland, J.I., Deblieux, D.D., Madsen, S.K., 2010. New basal igua-
nodonts from the Cedar Mountain Formation of Utah and the evolution of
thumb-spiked dinosaurs. PLoS One 5 (11), e14075.
McIntosh, J.S., 1990. Sauropoda, p.345-401. In: Weishampel, D.B., Dodson, P.,
Osmo
lska, H. (Eds.), The Dinosauria. University of California Press, Berkeley, CA,
pp. 345e390.
McPhee, B.W., Mannion, P.D., De Klerk, W.J., Choiniere, J.N., 2016. High diversity in
the sauropod dinosaur fauna of the Lower Cretaceous kirkwood Formation of
South Africa: implications for the Jurassic-Cretaceous transition. Cretaceous
Research 59, 151e205.
Milner, A.C., Milner, A.R., Evans, S.E., 2000. Amphibians, reptiles and birds: a
biogeographical review. In: Culver, S.J., Rawson, P.F. (Eds.), Biotic Response to
Global Change e The last Million 145 Million Year. Cambridge University Press,
Cambridge, pp. 316e332.
Mocho, P., Royo-Torres, R., Ortega, F., 2014. Phylogenetic reassessment of Lour-
inhasaurus alenquerensis, a basal Macronaria (Sauropoda) from the Upper
Jurassic of Portugal. Zoological Journal of the Linnean Society 170, 875e916.
Mocho, P., Royo-Torres, R., Ortega, F., 2016. New data for the Portuguese brachio-
saurid Lusotitan atalaiensis (Sobral Formation, Upper Jurassic). Historical
Biology 168, 98e206. http://dx.doi.org/10.1080/08912963.2016.1247447.
Mocho, P., Royo-Torres, R., Malafaia, E., Escaso, F., Ortega, F., 2017. Sauropod tooth
morphotypes from the Upper Jurassic of the Lusitanian Basin (Portugal). Papers
in Palaeontology 3, 259e295. http://dx.doi.org/10.1002/spp2.1075.
Naish, D., Martill, D.M., 2001. Saurischian dinosaurs 1: Sauropoda. In: Martill, D.M.,
Naish, D. (Eds.), Dinosaurs of the Isle of Wight. The Paleontological Association,
London, pp. 185e241.
Ortega, F., Escaso, F., Gasulla, J.M., Dantas, P., Sanz, J.L., 2006. Dinosaurios de las
Península Ibe
rica. Estudios Geolo
gicos 62, 219e240.
Pereda-Suberbiola, X., 2006. El estudio de los dinosaurios en Espan ~ a. In: Fundacio
n
del Patrimonio Histo
rico de Castilla y Leo
n (Ed.), Huellas que perduran. Icnitas
de dinosaurios: patrimonio y recurso, pp. 54e83.
Pereda-Suberbiola, X., Fuentes, C., Meijide, M., Meijide-Fuentes, F., Meijide-
Fuentes, M., 2007. New remains of the ankylosaurian dinosaur Polacanthus from
the Lower Cretaceous of Soria, Spain. Cretaceous Research 28, 583e596.
Poropat, S.F., Mannion, P.D., Upchurch, P., Hocnull, S.A., Kear, B.P., Elliot, D.A., 2015a.
Reassessment of the non-titanosaurian somphospondylan Wintonotitan wattsi
(Dinosauria: Sauropoda: Titanosauriformes) from the mid-Cretaceous Winton
Formation, Queensland, Australia. Papers in Paleontology 1, 59e106.
Poropat, S.F., Upchurch, P., Mannion, P.D., Hocnull, S.A., Kear, B.P., Sloan, T.,
Sinapius, G.H.K., Elliot, D.A., 2015b. Revision of the sauropod dinosaur Dia-
mantinasaurus matildae Hocknull et al., 2009 from the middle Cretaceous of
Australia: implications for Gondwanan titanosauriform dispersal. Gondwana
Research 27, 995e1033.
Poropat, S.F., Mannion, P.D., Upchurch, P., Hocnull, S.A., Kear, B.P., Kundr
at, M.,
Tischler, T.R., Sloan, T., Sinapius, G.H., Elliot, J.A., Elliot, D.A., 2016. New
Australian sauropods shed light on Cretaceous dinosaur palaeobiogeography.
Scientific Report 6, 34467. http://dx.doi.org/10.1038/srep34467.
Ratkevitch, R., 1998. New Cretaceous brachiosaurid dinosaur, Sonorasaurus thomp-
soni gen. Et sp. Nov. from Arizona. Journal of the Arizona-Nevada Academy of
Science 31, 71e82.
Rauhut, O.W.M., 2006. A brachiosaurid sauropod from the Late Jurassic Can ~ ado
n
Calcareo Formation of Chubut, Argentina. Fossil Record 9, 226e237.
Riggs, E.S., 1903. Brachiosaurus altithorax, the largest kown dinosaur. American
Journal of Science 15, 299e306.
Riggs, E.S., 1904. Structure and relationships of opisthocoelian dinosaurs. Part II, the
Brachiosauridae. Field Columbian Museum. Geological Series 2 (6), 229e247.
Rose, P., 2007. A new titanosauriform sauropod (Dinosauria: Saurischia) from the
Early Cretaceous of central Texas and its phylogenetic relationships. Paleon-
tologia Electronica 10, 1e65.
Royo y Go
mez, J., 1926a. Nuevos vertebrados de la facies wea
ldica de Los Can ~ os
(Soria) y Benage
ber (Valencia) y algunos moluscos cuaternarios de Villavieja
(Castello
n). Boletín de la Real Sociedad Espan ~ ola de Historia Natural 26,
317e318.
Royo y Go
mez, J., 1926b. Los vertebrados del Cret
acico espan ~ ol de facies wea
ldica.
Boletín del Instituto Geolo
gico de Espan ~ a 47, 170e176.
Royo y Go
mez, J., 1926c. Los descubrimientos de reptiles gigantescos en Levante.
Boletín de la Sociedad Castellonense de Cultura 7, 147e162.
Royo-Torres, R., 2009. El sauro
podo de Pen ~ arroya de Tastavins, 6. Instituto de
Estudios Turolenses-Fundacio
n Conjunto Paleontolo
gico de Teruel-Dino
polis,
Monografías Turolenses, pp. 1e548.
Royo-Torres, R., Cobos, A., 2007. Teeth of Oplosaurus armatus (Sauropoda) from El
Castellar (Teruel, Spain). In: 5th Meeting of the European Association of
Vertebrate Palaeontologists-12th European Workshop of Vertebrate Palae-
ontology, Carcassone-Espe
raza, abstracts volume, pp. 52e54.
Royo-Torres, R., Upchurch, P., 2012. The cranial anatomy of the sauropod Turiasaurus
riodevensis and implications for its phylogenetic relationships. Journal of Sys-
tematic Palaeontology 10, 553e583.
Royo-Torres, R., Cobos, A., Alcal
a, L., 2006. A Giant European Dinosaur and a New
Sauropod Clade. Science 314, 1925e1927.
Royo-Torres, R., Meijide Calvo, M., Fuentes Vidarte, C., Meijide-Fuentes, F., Meijide-
Fuentes, M., 2009. The Iberian Titanosauriformes of Zorralbo (Soria, Spain) from
Lower Cretaceous. In: Delgado-Buscalioni, A., Fregenal-Martínez, M. (Eds.),
Tenth International Symposium on Mesozoic Terrestrial Ecosystems and Biota.
Universidad Auto
noma de Madrid, pp. 143e144.
Royo-Torres, R., Alcala
, L., Cobos, A., 2012. A new specimen of the Cretaceous
sauropod Tastavinsaurus sanzi from El Castellar (Teruel, Spain), and a phyloge-
netic analysis of the Laurasiformes. Cretaceous Research 34, 61e83.
Royo-Torres, R., Upchurch, P., Mannion, P.D., Mas, R., Cobos, A., Gasco
, F., Alcala

, L.,
Sanz, J.L., 2014. The anatomy, phylogenetic relationships and stratigraphic po-
sition of the Tithonian-Berriasian Spanish sauropod dinosaur Aragosaurus
ischiaticus. Zoological Journal of Linnean Society 71, 623e655.
Ruiz-Omen ~ aca, J.I., Canudo, J.I., 2005. “Pleurocoelus“ valdensis Lydekker, 1889 (Sau-
rischia, Sauropoda) en el Creta
cico Inferior (Barremiense) de la Península
Ibe
rica. Geogaceta 38, 43e46.
Russell, D.A., 1993. The role of Central Asia in dinosaurian biogeography. Canadian
Journal of Earth Sciences 30, 2002e2012.
Salgado, L., Coria, R.A., Calvo, J.O., 1997a. Evolution of titanosaurid sauropods. I:
Phylogenetic analysis based on the postcranial evidence. Ameghiniana 34, 3e32.
Salgado, L., Coria, R.A., Calvo, J.O., 1997b. Presencia del ge
nero Aeolosarus (Sau-
ropoda, Titanosauridae) en la Formacio
n Los Alamitos, Creta
cico Superior de la
provincia de Río Negro, Argentina. Geocie ^ncias II 6, 44e49.
Sanz, J.L., Buscalioni, A.D., Casanovas, M.L., Santafe
, J.V., 1987. Dinosaurios del
Cret
acico Inferior de Galve (Teruel, Espan ~ a). Estudios Geolo
gicos Extr. Galve -
Tremp 45e64.
Sanz, J.L., Buscalioni, A.D., Pe
rez-Moreno, B., Moratalla, J., Jime
nez García, S., 1992.
Los dinosaurios de Castilla y Leo
n. In: Jime
nez-Fuentes, E. (Ed.), Vertebrados

siles de Castilla y Leo
fo
n. Museo de Salamanca, pp. 47e57.
Seeley, H.G., 1887. On the classification of the fossil animals commonly called
Dinosauria. Proceedings of the Royal Society London 43, 165e171.
Skelton, P.W., Spicer, R.A., Kelley, S.P., Gilmour, I. (Eds.), 2003. The Cretaceous world.
Cambridge University Press, Cambridge, pp. 1e360.
Smith, A.G., Smith, D.G., Funnell, B.M., 1994. Atlas of Mesozoic and Cenozoic
coastlines. Cambridge University Press, Cambridge, pp. 1e99.
Taylor, M.P., 2009. A re-evaluation of Brachiosaurus altithorax Riggs 1903 (Dino-
sauria, Sauropoda) and its generic separation from Giraffatitan brancai
(Janensch 1914). Journal of Vertebrate Paleontology 29, 787e806.
Taylor, M.P., Naish, D., 2007. An unusual new neosauropod dinosaur from the Lower
Cretaceous Hastings Beds Group of East Sussex, England. Palaeontology 50,
1547e1564.
Taylor, M.P., Wedel, M.J., Cifelli, R.L., 2011. A new sauropod from the Lower Cretaceous
Cedar Mountain Formation, Utah, USA. Acta Palaeontologica Polonica 56, 75e98.
Tidwell, V., Carpenter, K., 2002. Bridging the Atlantic: new correlations of Early
Cretaceous Titanosauriformes (Sauropoda) from England and North America.
Journal of Vertebrate Paleontology 22 (S3), 114A.
Tidwell, V., Carpenter, K., Brooks, W., 1999. New sauropod from the Lower Creta-
ceous of Utah, USA. Oryctos 2, 21e37.
Tidwell, V., Carpenter, K., Meyer, S., 2001. New Titanosauriform (Sauropoda) from
the Poison Strip Member of the Cedar Mountain Formation (Lower Cretaceous),
Utah. In: Tanke, D.H., Carpenter, K. (Eds.), Mesozoic Vertebrate Life. Indiana
University Press, pp. 139e165.
Torcida Fern
andez-Baldor, F., Canudo, J.I., Huerta, P., Montero, D., Pereda
Suberbiola, X., Salgado, L., 2011. Demandasaurus darwini, a new rebbachisaurid
sauropod from the Early Cretaceous of the Iberian Peninsula. Acta Palae-
ontologica Polonica 56, 535e552.
Tschopp, E., Mateus, O., Benson, R., 2015. A specimen-level phylogenetic analysis
and taxonomic revision of Diplodocidae (Dinosauria, Sauropoda). PeerJ 3, e857.
http://dx.doi.org/10.7717/peerj.857.
Upchurch, P., 1995. Evolutionary history of sauropod dinosaurs. Philosophical
Transactions of the Royal Society of London B 349, 365e390.
Upchurch, P., 1998. The phylogenetic relationships of sauropod dinosaurs. Zoolog-
ical Journal of the Linnean Society 124, 43e103.
Upchurch, P., Martin, J., 2002. The Rutland Cetiosaurus: The anatomy and relation-
ships of a Middle Jurassic British sauropod dinosaur. Palaeontology 45,
1049e1074.
 R. Royo-Torres et al. / Cretaceous Research 80 (2017) 38e55
Upchurch, P., Martin, J., 2003. The anatomy and taxonomy of Cetiosaurus (Saurischia,
Sauropoda) from the Middle Jurassic of England. Journal of Vertebrate Pale-
ontology 23, 208e231.
Upchurch, P., Barrett, P.M., Dodson, P., 2004. Sauropoda. In: Weishampel, D.B.,
Dodson, P., Osmolska, H. (Eds.), The Dinosauria, second ed. University of Cali-
fornia Press, Berkeley, pp. 259e322.
Upchurch, P., Mannion, P.D., Barrett, P., 2011. Sauropod dinosaurs. In: Batten, D.J.
(Ed.), English Wealden fossils. The Palaeontological Association, London,
pp. 476e525.
Wedel, M.J., Taylor, M.P., 2013. Caudal pneumaticity and pneumatic hiatuses in the
sauropod dinosaurs Giraffatitan and Apatosaurus. PLoS One 8, e78213.
Whitlock, J.A., 2011. A phylogenetic analysis of Diplodocoidea (Saurischia: Sau-
ropoda). Zoological Journal of the Linnean Society 161, 872e915.
Wilson, J.A., 2002. Sauropod dinosaur phylogeny: critique and cladistic analysis.
Zoological Journal of the Linnean Society 136, 217e276.
Wilson, J.A., Sereno, P.C., 1998. Early evolution and higher-level phylogeny of
sauropod dinosaurs. Journal of Vertebrate Palaeontology 18, 1e68.
Wilson, J.A., Upchurch, P., 2009. Redescription and reassessment of the phylogenetic
affinities of Euhelopus zdanskyi (Dinosauria: Sauropoda) from the Early Creta-
ceous of China. Journal of Systematic Palaeontology 7, 199e239.
Wiman, C., 1929. Die Kreide-Dinosaurier aus Shantung. Palaeontologia Sinica Series
C6, 1e67.
Woodruff, D.C., 2012. A new titanosauriform from the Early Cretaceous Cloverly
Formation of Montana. Cretaceous Research 36, 58e66.
Appendix
D'Emic et al. (2016)
55
The following characters have been modified for Tastavinsaurus according to
Royo-Torres et al. (2012) and personal observations (RRT): Character 18, 1 changed to
2; Character 36, 0 changed to 1; Character 56, 0 changed to 1; Character 60,
0 changed to 1; Character 102, 1 changed 0&1; Character 105, 0 changed to 1;
Character 118, 0 changed to 1; Character 119,? changed to 0.
Scoring of Soriatitan golmayensis for the characters of the modified D'Emic et al.
(2016) matrix
?????????????2?011???????????????????11?????????0????001?1110???
1????????????100??00??0??????????111?00011??10??????????00
Mannion et al. (2017)
The following characters have been modified for Tastavinsaurus according to
personal observations (RRT): Character 25, 0 changed to 1; Character 26, 0 changed
to 1; Character 27, 1 changed to 0; Character 60, 0 changed to 0&1; Character 61,
0 changed to 0&1; Character 62, 0 changed to 1; Character 66, 1 changed to 0;
Character 70, 1 changed to 0&1; Character 71, 0 changed to 0&1; Character 177, 1
changed to 0&1; Character 178, 0 changed to 1; Character 179, ? changed to 0;
Character 180, ? changed to 0; Character 187, 1 changed to 0; Character 195, 1
changed to 2; Character 249, ? changed to 0; Character 256, 1 changed to 0; Char-
acter 259, 0 changed to 1; Character 260, 1 changed to 0.
Scoring of Soriatitan golmayensis for the characters of the modified Mannion
et al. (2017) matrix
??????????1??????????1??1101000?101????311010???????????1???
101111??????????????????????????????????????????
001011??????????????????????????20111?2????????????????????????????
00??000000?011??110210001????21?00?????????????0110000001???????????
10?1?0??000??010????????????????????
1???????????????????????????????????????0???????????000???????????????
0?00?000??????0100101???????????10??01?00???????????0?????????????