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S0195-6671(16)30151-3
DOI:
10.1016/j.cretres.2016.08.001
Reference:
YCRES 3431
To appear in:
Cretaceous Research
24 July 2016
Please cite this article as: Dez Daz, V., Mocho, P., Pramo, A., Escaso, F., Marcos-Fernndez, F.,
Sanz, J.L., Ortega, F., A new titanosaur (Dinosauria, Sauropoda) from the Upper Cretaceous of Lo
Hueco (Cuenca, Spain), Cretaceous Research (2016), doi: 10.1016/j.cretres.2016.08.001.
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Grupo de Biologa Evolutiva. Facultad de Ciencias. UNED. Senda del Rey 9 28040.
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Madrid, Spain.
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2, 28040 Madrid.
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* Corresponding author
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ABSTRACT
The upper Campanian-lower Maastrichtian site of Lo Hueco (Cuenca, Spain) has
provided a set of well-preserved partial skeletons in anatomical connection or with a
low dispersion of their skeletal elements. One partial skeleton is herein described and a
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short postspinal lamina with a transversely expanded distal end represented by smooth
scars in the dorsal vertebrae; anteriormost caudals with the medial
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having two round and rough structures in the dorsal edge of the posterior articulation,
which extends to the dorsal surface of the centrum; the articular ends of the rami of the
haemal arches are divided in two articular surfaces; and tuberosity between the anterior
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and the lateral trochanter of the fibula. The herein performed phylogenetic analysis
considered L. pandafilandi as a member of Lithostrotia more derived than
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1. Introduction
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present on all continents (e.g. Upchurch et al., 2004; Cerda et al., 2012), and several
new occurrences demonstrate that they were relatively abundant in the European
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territory (e.g. Sanz et al., 1999; Le Loeuff, 2005; Csiki et al., 2010; Garcia et al., 2010;
Ortega et al., 2015). To date, just three titanosaurs are described for the Campanian-
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of teeth (Dez Daz et al. 2012, 2013a, 2014) and appendicular remains (Vila et al.,
2012; Pramo et al., 2014, 2015a, b).
The titanosaurian record from the Upper Cretaceous of southern France has been
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known since the last years of the nineteenth century (Depret, 1899, 1900). The
discovery in the last decades of more than 20 sites with sauropod remains has greatly
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improved the knowledge of the French titanosaurian sauropod faunas (e.g. Le Loeuff,
1995, 2005; Garcia et al., 2010; and see Dez Daz, 2013 for a complete list). One of the
best known titanosaurs from the French Upper Cretaceous record is Ampelosaurus
atacis (Le Loeuff, 1995, 2005), found in Bellevue (Aude Department, southern France).
The second titanosaurian species from the Upper Cretaceous of France is
Atsinganosaurus velauciensis (Garcia et al., 2010) from Velaux (Bouches-du-Rhne
Department, southeastern France).
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In Spain and Portugal more than 15 relevant Upper Cretaceous fossil-sites with
titanosaurian remains are known (e.g. Antunes and Sigogneau-Russell, 1992; Sanz et
al., 1999; Antunes and Mateus, 2003; Ortega et al., 2012; and Dez Daz, 2013 for a
complete list). The first titanosaurian remains found in the Iberian Peninsula were
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recovered from the Maastrichtian of the Tremp Basin (NE Spain) (Lapparent and
Aguirre, 1956, 1957), but, subsequently, a great deal of titanosaur bone remains have
been reported from a suite of sites mainly located in the eastern half of the Iberian
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Peninsula: Barcelona (Sells et al., 2015), Cuenca (Ortega et al., 2008, 2015),
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(Casanovas-Cladellas and Santaf-Llopis, 1993; Casanovas et al., 1987; CasanovasCladellas et al., 1995; Masriera and Ullastre, 1987), Segovia (Prez-Garca et al., 2015;
Sanz and Buscalioni, 1987), Soria (Lapparent et al., 1957; Pereda Suberbiola and RuizOmeaca, 2001), and Valencia (Company et al., 2009; Company and Pereda-
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Suberbiola, 2013).
One of the most outstanding sites is the Lao quarry (upper Campanian-lower
Maastrichtian, northern Spain) that has yielded numerous cranial and postcranial
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elements, and is the type locality of Lirainosaurus astibiae (Sanz et al., 1999).
Titanosaurian remains that have also been ascribed to Lirainosaurus were also found in
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the Upper Cretaceous fossil site of Chera, Valencia (Company et al., 2009). In this site,
two titanosaurian morphotypes were identified and one of them referred to
Lirainosaurus (Dez Daz et al., 2015). L. astibiae is the only described titanosaur taxon
for the Iberian Upper Cretaceous.
In 2007 a singular accumulation of fossils representing individuals of some
lineages of continental tetrapods was found at Lo Hueco (Fuentes, Cuenca) (Fig. 1a), in
the southwestern branch of the Iberian Ranges (central Spain). Abundant remains
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attributed to fishes, amphibians, squamate lizards, turtles, crocodiles, and different
groups of dinosaurs have been recognized (Ortega et al., 2015), but sauropods are the
group of vertebrates more abundantly represented in the site.
The Lo Hueco fossil site is placed in sedimentary deposits belonging to the
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upper part of the Margas, Arcillas y Yesos de Villalba de la Sierra Formation (Vilas et
al., 1982) (Fig. 1b). Its stratigraphic position and palaeontological content indicate that
its sediments were deposited in the late Campanian-early Maastrichtian interval (Ortega
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et al., 2015). A succession of versicolor marly mudstone levels, with a rich and diverse
fossil concentration, modified by sandy channel structures can be observed in the site.
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The palaeoenvironmental interpretation suggests a near coast muddy flood plain crossed
by distributary sandy channels environment, exposed to brackish to fresh water aquatic
influences (Barroso-Barcenilla et al., 2009). The bed in which the described specimen
was collected was identified as G1 (Fig. 1c), corresponding to the proximal part of a
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flooded muddy plain, close to distributary channels (see more data on the geological
context of the fossil site in Barroso-Barcenilla et al., 2009).
In the case of the sauropods, the site has yielded multiple partial skeletons in
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The new specimen herein described as the holotype of Lohuecotitan pandafilandi is the
most complete titanosaurian sauropod so far described for the European Upper
Cretaceous record. In this sense, this specimen will be important to understand the
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For the anatomical structures we use Romerian terms (Wilson, 2006) for the
structures (e.g. centrum) and their orientation (e.g. anterior). The nomenclature
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used for the caudal vertebrae is the one proposed by Dez Daz et al. (2013b). This study
applies the landmark-based terminology of Wilson (1999, 2012) and Wilson et al.
(2011) for vertebral laminae and fossae. For the haemal arches, it used the nomenclature
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Espraza, France.
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ilped, iliac peduncle; lat. spol, lateral spinopostzygapophyseal lamina; lat. sprl, lateral
spinoprezygapophyseal lamina; lb, lateral bulge; lt, lateral trochanter; med. sprl, medial
spinoprezygapophyseal lamina; pa, parapophysis; pcdl, posterior centrodiapophyseal
lamina; pl, pleurocoel; pcpl, posterior centroparapophyseal lamina; po,
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posterior process of the ulna; pped, pubic peduncle; prsl, prespinal lamina; raa, radial
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3. Systematic palaeontology
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Etymology. Lohuecotitan from Lo Hueco (the type locality) and titan (the giants of the
Greek mythology).
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Fig. 2-5
Holotype. A partial skeleton disarticulated, but whose remains had low dispersion in the
field. The complete set is referred as HUE-EC-01 and is composed of: three cervical
vertebrae and neural arch fragments (HUE-03128, 03076, 03077, 03136); six dorsal
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03134/17, 03134/19); left ulna (HUE-03044); left and right ischia (HUE-03099,
03134/15); left pubis (HUE-03086); right femur (HUE-03108); right fibula (HUE-
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03087); right tibia (HUE-03082), and several other indeterminate remais (HUE-3043,
3046, 3067, 3083, 3084).
Etymology. The specific name pandafilandi refers to Pandafilando de la fosca vista, one
of the characters in the novel The Ingenious Gentleman Don Quixote of La Mancha
(El ingenioso hidalgo don Quijote de la Mancha) written by Miguel de Cervantes and
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published in the early seventeenth century. Pandafilando is, in the mind of the
protagonist, a giant against who he must fight.
Type locality and horizon. The Lo Hueco fossil site (Fuentes, Cuenca, Castilla-La
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Mancha, central Spain) (Fig. 1), Margas, Arcillas y Yesos de Villalba de la Sierra
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vertebrae with a dorsal projection of the prsl and posl, resulting V-shaped outline in
lateral view*; (5) anterior neural spines on caudal vertebrae bear a greek-cross-like
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cross-section*; (6) middle caudal vertebrae having two rough structures in the dorsal
edge of the posterior articulation, which extends to the dorsal surface of the centrum*;
(7) the articular ends of the rami of the posterior haemal arches are fully divided in two
articular facets*; (8) tuberosity between the anterior and the lateral trochanter of the
fibula (shared with Jainosaurus).
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Description. The holotype of Lohuecotitan pandafilandi is composed of several axial
and appendicular elements found in partial or in full articulation. At least three cervical
centra (one anterior, and fragments of one middle and one posterior cervical) and
several neural arch fragments were identified (Fig. 2a-b). They have camellate internal
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subcircular cross-section. Shallow pneumatic fossae appear in the lateral surfaces of the
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centra, and they become deeper in the middle and posterior vertebrae. The pneumatic
fossae are divided in the middle and posterior cervicals by a subvertical lamina. The
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diapophyseal laminae are well developed. Shallow and poorly defined pneumatic fossae
in the cervical vertebrae have been considered as typical of Somphospondyli (e.g.
Salgado et al., 1997; Wilson, 2002; Wedel, 2003; DEmic, 2012). The anterior and
middle sector of the ventral surface of the centra is slightly transversely concave
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delimited by two ventrolateral ridges, and lacks a sagittal crest. The neural arch is
composed by unbifurcated centroprezygapophyseal (cprl), centropostzygapophyseal
(cpol), spinoprezygapophyseal (sprl) and spinopostzygapophyseal (spol) laminae. No
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prsl is present in the preserved neural spine. No complete neural spines are preserved,
and the morphology of the distal tip is unknown.
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dorsal vertebrae (partially articulated) and two possible posterior dorsal vertebrae (Fig.
2c-d). The preserved neural arches are generally badly preserved. The first four
vertebrae represent the transition between the anterior and middle dorsal vertebrae. The
dorsal centra are opisthocoelous and slightly higher than wide, having a subcircular
cross-section. The pneumatic foramina are small, eyed-shaped and not particularly deep.
Some subvertical and stout laminae are present within the pneumatic foramina. The
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morphology of the ventral surface of the centra is flat, presenting a shallow sagittal crest
in the anteriormost dorsal centra. The transverse processes are laterally projected. The
diapophyses are located posterodorsally to the parapophyses. The transverse process has
dorsally facing flattened area. One of the most important features in the dorsal vertebrae
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There is a main spinodiapophyseal lamina (spdl) and an accessory spdl. The accessory
spdl is located on the anterior side of the spdl and connects the ventral sector of the spdl
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to the dorsal part of the prsl. In posterior view, there is a lateral spol (lat. spol), well
developed on the proximal half of the neural spine, and a short med. spol that extends
from the medial side of the poszygapophysis and connects to the ventral tip of the posl.
The presence of a short med. spol connected to the base of the posl occurs in the derived
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titanosaurs Lirainosaurus (Dez Daz et al., 2013b), Opisthocoelicaudia (BorsukBialynicka, 1977) and Elaltitan (Mannion and Otero, 2012), but also in other nonderived titanosaurs (Wilson, 1999). The neural spine is posteriorly deflected (more than
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50 degrees from the vertical plane). It bears a lateral triangular process with an
appreciable dorsoventral development. The posl is well developed and its ventral
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section is transversely narrow. The posl becomes smoother and laterally expanded in the
dorsal end, not reaching the dorsal border of the neural spine. Smooth scars represent
the distal end of the posl. A dorsoventrally short posl with a smooth distal end is a
diagnostic feature of Lohuecotitan.
The last dorsal vertebra is poorly preserved, in association with the sacrum, and
presents an important taphonomical deformation. The centrum is opisthocoelous and
pleurocoels are present. The ventral surface is convex. The cprl are stout and simple,
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and a deep centroprezygapophyseal fossa (cprf) is present between the cprl and the
intraprezygapophyseal lamina (tprl). The ventral tip of the pcdl is forked in this
posterior dorsal vertebra.
Dorsal ribs bear a medial pneumatic fossa in the proximal end as in other
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Only some fragments of the sacral vertebrae are present. There are, at least, five
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fused sacral neural spines bearing a dorsal ligament. Six sacral ribs are attached to the
medial side of the ilium. The sacral vertebrae also present a camellate bone tissue.
Lohuecotitan preserves an almost complete anterior to middle caudal series of 20
caudal vertebrae (the first nine anterior caudal vertebrae were found articulated) (Fig.
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3). The caudal neural arches are anteriorly displaced as in Titanosauriformes (e.g.
Salgado et al., 1997; Wilson, 2002; Upchurch et al., 2004). The hyposphenic ridge is
absent in the caudal vertebrae, as in most somphospondylians (Upchurch et al., 2004;
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Carballido et al., 2011). All of them are procoelous, including the first caudal vertebra,
showing a large posterior condyle (see measurements in Supplementary Material I). The
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posterior convexity in the anterior and middle caudal vertebrae bears a slight dorsal
displacement. Along the tail, the procoelous condition becomes progressively less
pronounced. The presence of procoelous anterior and middle caudal vertebrae is shared
by the members of Lithostrotia (e.g. Salgado et al., 1997; Upchurch, 1998; Upchurch et
al., 2004; DEmic, 2012). The caudal ribs are posterolaterally projected and ventrally
deflected in anterior view. The anterior caudal vertebrae lack pronounced diapophyseal
laminae or tuberosity in the caudal rib. The anterior caudal centra are higher and wider
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than long, and have a sub-circular outline in anterior/posterior view. The ventral
surfaces of the centrum are flat, and some of them bear a slightly concavity in their
posterior third. The neural arch in the anterior caudal vertebrae presents a complex and
autapomorphic lamination. The anterior caudal neural spines bear robust sagittal prsl
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and posl. The lateral sprl (lat. sprl) are short (they do not reach the midlength of the
lateral aspect of the spine) and the lat. spol extends along the neural spine height. The
med. sprl and med. spol are present and they are ventrally connected with the prsl and
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pandafilandi. Another peculiar feature is the presence of a dorsal projection of the prsl
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and posl, resulting in a V-shaped outline in lateral view. The neural spines bear a greek
cross-like cross-section. This morphology on the anteriormost caudal neural spines is
also tentatively considered as diagnostic of Lohuecotitan pandafilandi. The anterior
neural spines are subvertical bearing a slight posterior deflection and from the ninth
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caudal they become vertical. The prezygapophyses are ventrally and dorsally expanded
on the most anterior caudal vertebrae, a feature present in aeolosaurines such as
Aeolosaurus, Overosaurus and Panamericansaurus (Calvo and Porfiri, 2010; Santucci
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and Arruda-Campos, 2011; Coria et al., 2013). In the middle caudals, the width of the
prezygapohyses reaches 40-to-50% of the total length of the centrum. The seventh and
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the eighth caudal vertebrae are pathologically fused to each other and to the haemal arch
(Fig. 3d). In the middle caudals, the dorsal edge of the posterior articular facet bears two
rough structures which extend to the dorsal surface of the centrum, considered here as a
putative autapomorphy of Lohuecotitan pandafilandi.
All the anterior haemal arches are open Y-shaped, whereas the posterior ones are
open V-shaped (Fig. 4). The haemal arches are also dorsally open as in most
macronarians (Wilson, 2002; Curry Rogers, 2005; Mannion and Calvo, 2011; Otero et
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al., 2012). The two surfaces for articulations with the centra in each haemal arch are
subdivided into an anterior and a posterior articular facet that are separated by a
conspicuous transverse groove. These articulations are only completely separated in
middle and distal haemal arches. The presence of two clearly separated articular facets
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in the posterior haemal arches is not known in other European titanosaurs and is here
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Arruda-Campos, 2011) and some euhelopodids (DEmic, 2012). The haemal canal is ca.
50% of the total length of the most anterior haemal arch, as occurs in most
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titanosauriforms (e.g. Wilson, 2002; Upchurch et al., 2004; DEmic, 2012; Mannion et
The ulna of Lohuecotitan (Fig. 5b-c) seems to be a robust element (the proximal
and distal end are not preserved) as is also the case in the ulnae of Ampelosaurus. The
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proximal end of the ulna is triradiate, as in most sauropods (e.g. Wilson, 2002;
Upchurch et al., 2004). The anteromedial process is longer than the anterolateral
process. The posterior process is posteriorly developed and transversely constricted.
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The proximal and distal ends are not fully preserved. The anterior process seems to be
much larger than the lateral one.
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The ilium (Fig. 5a) is internally pneumatized (camellate bone tissue), and the
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edge of the ischium and raised from the surface without an associated groove. The
anteroposterior width of the pubic peduncle corresponds to the 50% of the total length
of the ischium. The distal end of the ischium is coplanar. The ischium is shorter than
pubis, distal end with a plate-like morphology and the absence of an emargination are
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common in titanosaurs (Wilson, 2002; Upchurch et al., 2004; Gonzlez Riga et al.,
2009). The preserved left pubis are not complete (Fig. 5f). The main axis of the pubis is
straight, except from the proximal posterior edge, which is laterally twisted. No weak
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The shaft of the femur (Fig. 5d, g) is straight and anteroposteriorly compressed
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surface of the shaft near the medial edge, and its distal tip reach the midheight of the
femur. The distal condyles are medially beveled. The proximal end of the tibia has a Dshaped profile, and the cnemial crest is anterolaterally directed, specially its most distal
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point, when the flat subtriangular surface of the distal end is treated as the anterior
surface (Fig. 5i, k). The distal articular surface is more or less oval, longer transversely
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the crest-like anterior trochanter and the lateral trochanter there is an additional anterior
trochanter, considered as an autapomorphy of Lohuecotitan pandafilandi and shared
with Jainosaurus (Wilson et al., 2011) and Laplatasaurus (Gallina and Otero, 2015).
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4. Phylogenetic analysis
phylogenetic analysis was performed using the data matrix proposed by Carballido and
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Sander (2014). The data matrix was analyzed using TNT 1.1 (Goloboff, Farris and
Nixon, 2003) to find the most parsimonious trees (MPTs). We used a heuristic tree
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search performing 1000 replications of Wagner trees (using random addition sequences)
followed by tree bisection reconnection (TBR) as swapping algorithm, saving 100 trees
per replicate. To test the support of the phylogenetic hypotheses, Bremer support and
bootstrap (absolute frequencies based on 10000 replicates) values were performed in
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TNT 1.1. (Goloboff, Farris and Nixon, 2003). As in the original analysis (Carballido
and Sander, 2014), the multistate characters 12, 58, 95, 96, 102, 106, 108, 115, 116,
119, 120, 154, 164, 213, 216, 232, 233, 234, 235, 256, 267, 298, 299 and 301 were
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considered ordered.
The result of this analysis yielded 20 MPTs of 1004 steps with a consistency
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index (CI) of 0.405 and a retention index (RI) of 0.735 (Fig. 6). The general topology
obtained is similar to that proposed by Carballido and Sander (2014). Lohuecotitan
pandafilandi is placed within Lithostrotia, and this position is supported by the presence
of anterior and middle dorsal vertebrae with a strongly posteroventraly oriented (more
than 40) zygapophyseal articulation (character #149). Lohuecotitan is considered a
more derived lithostrotian than Malawisaurus, and is recovered in a polytomy with
Opisthocoelicaudia, Alamosaurus, Trigonosaurus, Nemegtosaurus, Saltasaurinae and
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Rapetosaurus + (Isisaurus + Tapauisaurus). The incorporation of Lohuecotitan within
this clade is supported by disappearance of the caudal ribs by caudal 10 (character
#189), and slightly directed posteriorly neural spines in middle caudal vertebrae
(character #213).
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neural spines with subparallel margins (character #138); iii) in anterior view, neural
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canal of the middle and posterior dorsal vertebrae is enclosed in a deep fossa (character
#155); iv) divided cpol in middle and posterior dorsals (character #158); v) subcircular
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middle and posterior dorsal centrum (character #162); vi) absence of spdl and spol
contact in middle dorsals (character #169); vii) femoral transverse diameter is 125150% of the anteroposterior diameter (character #301).
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titanosaurs remain uncertain (e.g. Curry Rogers, 2005; Mannion and Upchurch, 2011;
Dez Daz, 2013; Gorscak and OConnor, 2016). Future analyses focused on European
titanosaurian taxa will be important to better understand the evolutionary history of this
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group in the European territory and to clarify the relationships of these taxa with nonEuropean titanosaurs. The results obtained here support the presence of a new
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5. Discussion
In the last twenty years, five titanosaur taxa were established for the European
Campanian-Maastrichtian: Magyarosaurus, Lirainosaurus, Atsinganosaurus,
Ampelosaurus and Paludititan (Nopcsa, 1915; Huene, 1932; Le Loeuff, 1995; Sanz et
al., 1999; Csiki et al., 2010; Garcia et al., 2010).
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Several features allow us to compare and discuss the systematic affinities of
Lohuecotitan with the remaining European forms. Cervical vertebrae are relatively
unknown for the European titanosaurs, and only isolated and badly preserved specimens
were recovered. Lohuecotitan is characterized by the presence of long prezygapophyses,
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single sprl, spol and cpol, and the absence of a prsl. These laminae are also present in
Atsinganosaurus, except prsl. The diapophyseal laminae are well developed in
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Dorsal vertebrae are relatively diagnostic within sauropods (Wilson, 1999, 2012;
Wilson et al., 2011), being rare in the Upper Cretaceous of Europe. One of the more
remarkable features in some of the European taxa, i.e. Ampelosaurus and
Magyarosaurus, is the presence of a hyposphenic crest (Le Loeuff, 2005; Apestegua,
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2005; pers. obs., PM). A hyposphenic crest is common in basal titanosaurs such as
Andesaurus, Argentinosaurus or Epachthosaurus (e.g. Apestegua, 2005), and contrary
to other European titanosaurs, such as Lirainosaurus where this crest is absent.
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an accessory spdl, absent in Lirainosaurus and Paludititan. The transverse processes are
transversely projected as occurs in Atsinganosaurus, Magyarosaurus and Ampelosaurus
(in some vertebrae the transverse process are laterodorsally projected probably due
deformation), differing from the condition present in Lirainosaurus (Dez Daz et al.,
2013b). Lohuecotitan can be distinguished from the remaining European titanosaur taxa
by the presence of a well-developed triangular aliform process in the middle dorsals. In
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the Ampelosaurus holotype (MDE C3-247), the triangular aliform process is absent and
the neural spines are transversely narrow, differing from that of Lohuecotitan.
The proximal anterior caudals of Lohuecotitan mainly differ from those of
Ampelosaurus, Magyarosaurus and Paludititan in the complexity and development of
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the neural arch structures, bearing the following autapomorphic features: i) med. sprl
and med. spol ventrally connected with the prsl and posl, respectively; ii) dorsal
projection of the prsl and posl, and iii) greek-cross-like cross section of the neural
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spine. Lohuecotitan shares with Lirainosaurus the presence of deep fossae below the
prezygapophyses (cprf, observed in anterior view) in the proximal anterior caudal
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vertebrae, although in Lohuecotitan these are smaller, deeper, and more anteriorly
placed. In addition, and distinct to Lirainosaurus, the posl is only slightly posteriorly
expanded in Lohuecotitan. In Lohuecotitan the caudal neural spines are longer than that
of the other Ibero-Armorican titanosaurs, and dorsally directed as in the Romanian taxa.
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ischiatic remains. The general morphology of the ischium of Ampelosaurus (e.g. MDE
C3-594) presents the common features found in titanosaurs (i.e. plate-like shaft and a
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dorsoventrally long pubic articulation). The iliac peduncle of the ischium in Paludititan
is longer than the ones of Lohuecotitan and Ampelosaurus. Lohuecotitan does not
present a posterolateral buttress in the iliac peduncle of the ischium, as in most
titanosaurs, and differing from that of Ampelosaurus and Paludititan (Csiki et al.,
2010).
Some differences are observed in the hindlimb elements. The femoral shaft in
Lohuecotitan is only slightly anteroposteriorly compressed, differing from
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Lirainosaurus, some specimens referred to Magyarosaurus, and some specimens from
the Bellevue locality referred to Ampelosaurus. Lohuecotitan shares with Lirainosaurus
the presence of mediolaterally compressed tibial diaphysis, contrary to that of the
French taxa. However, the tibiae of Lohuecotitan and Ampelosaurus (e.g. MDE C3-138)
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outline of the proximal articulation, and not showing a prominent anteromedial ridge
close to the distal extremity delimiting two highly concave surfaces, features diagnostic
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of these last taxa (Dez Daz et al., 2013c). The fibula of Lohuecotitan has a shaft
slightly more sigmoidal than that of Lirainosaurus and Ampelosaurus. However,
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Lohuecotitan shares with the fibulae of Ampelosaurus, the expanded proximal and distal
ends, and the presence of a sigmoidal ridge associated with the lateral trochanter (Le
Loeuff, 1992, 2005) similar to the lower ridge placed near the posterior edge of the
lateral trochanter in Lohuecotitan.. As occurs in the Lo Hueco fossil site, a large
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reveal the presence of more than one taxa. The study of this variability will be important
in further phylogenetic analyses on the Ibero-Armorican titanosaurs.
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Other important fossil sites from the Upper Cretaceous of the Iberian Peninsula
are located in Lao (Trevio County, Sanz et al., 1999; Pereda-Suberbiola et al., 2015),
Chera (Valencia, Company, 2010) and South Pyrenees area (e.g. Vila et al., 2012).
From the Lao site, only one titanosaurian taxon is identified (Dez Daz et al., 2015).
From Chera, two morphotypes were identified, one of them assigned to Lirainosaurus
(Company et al., 2009; Dez Daz et al., 2015). Relatively to the second morphotype,
Lohuecotitan corresponds to a different form based on the morphology of the middle
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caudal vertebrae, femora and tibiae. The second morphotype of Chera has caudal
vertebrae with a subquadrangular outline in posterior view and a higher developed
articulation for the haemal arches. In addition, the femora from this second morphotype
of Chera are more robust than the one of Lohuecotitan, and the general morphology of
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the tibiae is clearly different, although both present a rounded profile of the proximal
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understanding the titanosaurian evolutionary history and diversity during the Late
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astibiae, it is possible to establish a new titanosaurian taxon, and this confirms that the
Iberian Peninsula was inhabited by at least two different titanosaurian species between
the Campanian and Maastrichtian.
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titanosaurian teeth and braincases have been found (Dez Daz et al., 2014; Knoll et al.,
2013, 2015; Pramo et al., 2014, 2015a, b; Ortega et al., 2015), but these remains have
been found isolated, and no direct correlation with the bones of Lohuecotitan is, so far,
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possible. One of the braincases has several similarities with Ampelosaurus, however, no
other remains found in Lo Hueco seem to have clear affinities with the French taxon. In
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the case of the teeth morphotypes, one of them shows the same features as the other
French taxon Atsinganosaurus, but (as in the case of the previous braincase) no
postcranial remains present any similarities with Atsinganosaurus. However, a tooth
morphotype from Lo Hueco is similar to one found in the French sites of FoxAmphoux-Mtisson (Dez Daz et al., 2012) and Massecaps, where a new titanosaur has
been found and its description is in progress (Dez Daz et al., 2013a). If the
titanosaurian remains found in Massecaps show similarities with the ones of
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Lohuecotitan we could finally correlate one of the tooth morphotypes found in Lo
Hueco with Lohuecotitan, and to consider the presence of this taxon in several other
sites of southwestern Europe.
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6. Conclusion
Cretaceous of the Ibero-Armorican Island. This new taxon came from the Lo Hueco
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expanded distal end represented by smooth scars in the dorsal vertebrae; anteriormost
caudals with the med. sprl and med. spol ventrally connected with the prsl and posl,
respectively; anterior neural spines with prsl and posl dorsally projected; double
articular facets of the rami of the distal haemal arches.
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Atsinganosaurus) to, at least, six or seven taxa (Dez Daz et al., 2015). The diagnosis of
Lohuecotitan is relevant to begin to update the titanosaurian diversity of Europe, and
more specifically of the Iberian Peninsula.
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7. Acknowledgements
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(Museo de Ciencias Naturales de Alava/Arabako Natur Zientzien Museoa, VitoriaGasteiz, Spain), B. Madarieta (Museo Vasco de Historia de la Medicina y de las
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software. The comments made by L. Salgado, P. Mannion, and the editor improved an
early version of this paper.
8. References
RI
PT
Antunes, M. T., & Mateus, O. (2003). Dinosaurs of Portugal. Comptes Rendus Palevol,
2, 77-95.
SC
M
AN
U
TE
D
Barroso-Barcenilla, F., Cambra-Moo, O., Escaso, F., Ortega, F., Pascual, A., PrezGarca, A., Rodrguez-Lzaro, J., Sanz, J. L., Segura, M., & Torices, A. (2009).
New and exceptional discovery in the Upper Cretaceous of the Iberian Peninsula:
EP
AC
C
ACCEPTED MANUSCRIPT
Calvo, J. O., & Porfiri, J. D. (2010). Panamericansaurus schroederi gen. nov. sp. nov.
Un nuevo Sauropoda (Titanosauridae-Aeolosaurini) de la Provincia del Neuqun,
Cretcico Superior de Patagonia, Argentina. Brazilian Geographical Journal:
Geosciences and Humanities research medium, 1, 100-115.
RI
PT
SC
M
AN
U
Albarracn, 1.
Carballido, J.L., Rauhut, O. W. M., Pol, D., & Salgado, L. (2011). Osteology and
phylogenetic relationships of Tehuelchesaurus benitezii (Dinosauria, Sauropoda)
from the Upper Jurassic of Patagonia. Zoological Journal of the Linnean Society,
TE
D
EP
AC
C
doi:10.1080/14772019.2013.764935
Casal, G., Martnez, R., Luna, M., Sciutto, J. C., & Lamanna, M. (2007). Aeolosaurus
colhuehuapensis sp. nov. (Sauropoda, Titanosauria) de la formacin Bajo Barreal,
Cretcico Superior de Argentina. Revista Brasileira de Paleontologia, 10(1), 5362.
ACCEPTED MANUSCRIPT
Casanovas, M. L., Santaf, J. V., Sanz, J. L., & Buscalioni, A. D. (1987). Arcosaurios
(Crocodilia, Dinosauria) del Cretcico Superior e la Conca de Tremp (Lleida,
Espaa). Estudios Geolgicos, Vol. Extr. Galve-Tremp, 95-110.
Casanovas-Cladellas, M. L., & Santaf-Llopis, J. V. (1993). Presencia de
RI
PT
SC
M
AN
U
5, 277-283.
Cerda, I. A., Carabajal, A. P., Salgado, L., Coria, R. A., Reguero, M. A., Tambussi, C.
P., & Moly, J. J. (2012). The first record of a sauropod dinosaur from Antarctica.
Die Naturwissenschaften, 99, 83-87. doi:10.1007/s00114-011-0869-x
TE
D
EP
Company, J., Pereda Suberbiola, X., & Ruiz-Omeaca, J.I. (2009). Nuevos restos
fsiles del dinosaurio Lirainosaurus (Sauropoda, Titanosauria) en el Cretcico
AC
C
Coria, R. A., Filippi, L. S., Chiappe, L. M., Garca, R., & Arcucci, A. B. (2013).
Overosaurus paradasorum; gen. et sp. nov., a new sauropod dinosaur
(Titanosauria: Lithostrotia) from the Late Cretaceous of Neuqun, Patagonia,
Argentina. Zootaxa 3683, 357. doi:10.11646/zootaxa.3683.4.2
ACCEPTED MANUSCRIPT
Csiki, Z., Codrea, V., Jipa-Murzea, C., & Godefroit, P. (2010). A partial titanosaur
(Sauropoda, Dinosauria) skeleton from the Maastrichtian of Nla-Vad, Haeg
Basin. Neues Jahrbuch fr Geologie und Palaontologie - Abhandlungen, 258,
297-324.
RI
PT
SC
M
AN
U
TE
D
EP
AC
C
ACCEPTED MANUSCRIPT
Fox-Amphoux-Mtisson (Var, SE France). Proceedings of the Geologists
Association, 123, 626-637.
Dez Daz, V., Ortega, F., & Sanz, J. L. (2014). Titanosaurian teeth from the Late
Cretaceous of Lo Hueco (Cuenca, Spain). Cretaceous Research, 51, 285-291.
RI
PT
Dez Daz, V., Pereda Suberbiola, X., & Company, J. (2015). Puesta al da de la
SC
Dez Daz, V., Pereda Suberbiola, X., & Sanz, J. L. 2013b. The axial skeleton of the
titanosaur Lirainosaurus astibiae (Dinosauria: Sauropoda) from the latest
M
AN
U
Dez Daz, V., Pereda Suberbiola, X., & Sanz, J. L. (2013c). Appendicular skeleton and
dermal armour of the Late Cretaceous titanosaur Lirainosaurus astibiae
(Dinosauria: Sauropoda) from Spain. Palaeontologia Electronica, 16(2), 19A, 18p.
TE
D
Dez Daz, V., Tortosa, T., & Le Loeuff, J. (2013a). Sauropod diversity in the Latest
Cretaceous of south-western Europe: The lessons of odontology. Annales de
Palontologie, 99(2): 119-129.
EP
Garca, G., Amico, S., Fournier, F., Thouand, E., & Valentin, X. (2010). A new
titanosaur genus (Dinosauria, Sauropoda) from the Late Cretaceous of southern
AC
C
Goloboff, P., Farris, J., & Nixon, K. (2008). TNT, a free program for phylogenetic
analysis. Cladistics, 24, 774-786.
Gonzlez Riga, B. J., Previtera, E., & Pirrone, C. A. (2009). Malarguesaurus florenciae
gen. et sp. nov., a new titanosauriform (Dinosauria, Sauropoda) from the Upper
ACCEPTED MANUSCRIPT
Cretaceous of Mendoza, Argentina. Cretaceous Research, 30, 135-148.
doi:10.1016/j.cretres.2008.06.006
Gorscak, E., & OConnor, P. M. (2016). Time-calibrated models support congruency
between Cretaceous continental rifting and titanosaurian evolutionary history.
RI
PT
SC
M
AN
U
Knoll, F., Ridgely, R. C., Ortega, F., Sanz, J. L., & Witmer, L. M. (2013). Neurocranial
osteology and neuroanatomy of a Late Cretaceous titanosaurian sauropod from
Spain (Ampelosaurus sp.). PLoS ONE, 8, e54991.
doi:10.1371/journal.pone.0054991
TE
D
Knoll, F., Witmer, L. M., Ridgely, R. C., Ortega, F., & Sanz, J. L. (2015). A new
titanosaurian braincase from the Cretaceous Lo Hueco locality in Spain sheds
light on neuroanatomical evolution within Titanosauria. PLOS ONE, 10,
EP
e0138233. doi:10.1371/journal.pone.0138233
Lapparent, A. F. de, & Aguirre, E. (1956). Algunos yacimientos de Dinosaurios en el
AC
C
ACCEPTED MANUSCRIPT
Le Loeuff, J. (1992). Les vrtbrs continentaux du Crtac suprieur dEurope:
Palocologie, Biostratigraphie et Paleobiogographie. Unpublished Ph.D. Thesis
(273 pp). Mmoires des Sciences de la Terre, Universit Pierre and Marie Curie,
Paris VI, France, 92-93.
RI
PT
SC
693-699.
M
AN
U
Mannion, P. D., & Calvo, J.O. (2011). Anatomy of the basal titanosaur (Dinosauria,
TE
D
EP
155-181. doi:10.1111/j.1096-3642.2011.00699.x
Mannion, P. D., & Upchurch, P. (2011). A re-evaluation of the mid Cretaceous
AC
C
sauropod hiatus and the impact of uneven sampling of the fossil record on
patterns of regional dinosaur extinction. Palaeogeography, Palaeoclimatology,
Mannion, P. D., & Otero, A. (2012). A reappraisal of the Late Cretaceous Argentinean
sauropod dinosaur Argyrosaurus superbus, with a description of a new titanosaur
genus. Journal of Vertebrate Paleontology, 32, 614-638.
doi:10.1080/02724634.2012.660898
ACCEPTED MANUSCRIPT
Mannion, P. D., 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 titanosauriforms. Zoological Journal of the
Linnean Society, 168, 98-206. doi:10.1111/zoj.12029
RI
PT
Masriera, A., & Ullastre, J. (1988). Nuevos datos sobre las capas maestrichtienses con
SC
M
AN
U
TE
D
EP
Ortega, F., & Prez-Garca, A. (2009). cf. Lirainosaurus sp. (Dinosauria: Titanosauria)
en el Cretcico Superior de Sacedn (Guadalajara). Geogaceta, 46, 87-90.
AC
C
Ortega, F., Bardet, N., Barroso-Barcenilla, F., Callapez, P. M., Cambra-Moo, O.,
Daviero- Gmez, V., Dez Daz, V., Domingo, L., Elvira, A., Escaso, F., GarcaOliva, M., Gmez, B., Houssaye, A., Knoll, F., Marcos-Fernndez, F., Martn, M.,
Mocho, P., Narvez, I., Prez-Garca, A., Peyrot, D., Segura, M., Serrano, H.,
Torices, A., Vidal, D., & Sanz, J. L. (2015). The biota of the Upper Cretaceous
site of Lo Hueco (Cuenca, Spain). Journal of Iberian Geology, 41, 83-99.
doi:10.5209/rev_JIGE.2015.v41.n1.48657
ACCEPTED MANUSCRIPT
Ortega, F., Sanz, J. L., Barroso-Barcenilla, F., Cambra-Moo, O., Escaso, F., GarcaOliva, M., & Marcos-Fernndez, F. (2008). El yacimiento de macrovertebrados
fsiles del Cretcico Superior de Lo Hueco (Fuentes, Cuenca). Paleontologica
Nova. Publicaciones del Seminario de Paleontologa de Zaragoza, 8, 119-131.
RI
PT
Ortega, F., Sanz, J. L., Dez Daz, V., & Escaso, F. (2012). Armoured titanosaurs from
the Upper Cretaceous of Lo Hueco (Cuenca, Spain). Fundamental! 20, 173-174.
Otero, A., Gallina, P. A., Canale, J. I., & Haluza, A. (2012). Sauropod haemal arches:
SC
M
AN
U
doi:10.1080/08912963.2011.618269
Owen, R. (1842). Report on British fossil reptiles. Reports of the British Association for
the Advancement of Science, II, 60-204.
Pramo, A., Ortega, F., Escaso, F., Narvez, I., & Sanz, J. L. (2014). Ejemplares
TE
D
Pramo, A., Ortega, F., & Sanz, J. L. (2015a). Two types of appendicular bones of
EP
AC
C
Pramo, A., Ortega, F., & Sanz, J. L. (2015b). Preliminar assessment of the
morphological variability of appendicular bones of titanosaurs (Dinosauria,
Sauropoda) from Lo Hueco (Fuentes, Cuenca). In Reolid, M. (Ed.), Libro de
resmenes de las XXXI Jornadas de Paleontologa de la Sociedad Espaola de
Paleontologa (pp. 225-226).
ACCEPTED MANUSCRIPT
Pereda-Suberbiola, X., & Ruiz-Omeaca, J. I. (2001). Un dinosaurio saurpodo
(Titanosauria) en el Cretcico superior de Cubilla, Soria (Espaa). Geogaceta, 30,
175-178.
Pereda-Suberbiola, X., Corral, J. C., Astibia, H., Badiola, A., Bardet, N., Berreteaga, A.,
RI
PT
Buffetaut, E., Buscalioni, A. D., Cappetta, H., Cavin, L., Dez Daz, V.,
Gheerbrant, E., Murelaga, X., Ortega, F., Prez-Garca, A., Poyato-Ariza, F.,
Rage, J. -C., Sanz, J. L., & Torices, A. (2015). Late Cretaceous continental and
SC
M
AN
U
doi:10.5209/rev_JIGE.2015.v41.n1.48658
Prez-Garca, A., Ortega, F., Bolet, A., Escaso, F., Houssaye, A., Martnez-Salanova, J.,
de Miguel Chaves, C., Mocho, P., Narvez, I., Segura, M., Torices, A., Vidal, D.,
& Sanz, J. L. (2015). A review of the upper Campanian vertebrate site of Armua
TE
D
Salgado, L., Coria, R. A., & Calvo, J.O. (1997). Evolution of titanosaurid sauropods I:
32.
EP
AC
C
Sanz, J. L., & Buscalioni, A.D. (1987). New evidence of armoured titanosaurs in the
Upper Cretaceous of Spain. In Currie, P. M., & Koster, E. H. (Eds.), Fourth
Symposium on Mesozoic Terrestrial Ecosystems, Short Papers (pp. 199-204).
ACCEPTED MANUSCRIPT
Sanz, J. L., Powell, J. E., Le Loeuff, J., Martnez, R., & Pereda-Suberbiola, X. (1999).
Sauropod remains from the Upper Cretaceous of Lao (Northecentral Spain).
Titanosaur phylogenetic relationships. Estudios del Museo de Ciencias Naturales
de lava, 14 (Nmero Especial 1), 235-255.
RI
PT
Sells, A. G., Marmi, J., Llcer, S., & Blanco, A. (2015). The youngest sauropod
doi: 10.1080/08912963.2015.1059834
SC
M
AN
U
Upchurch, P., Barrett, P., & Dodson, P. (2004). Sauropoda. In Weishampel, D. B.,
Dodson, P., & Osmlska, H. (Eds.), The Dinosauria, second edition (pp. 259-
TE
D
Vila, B., Galobart, ., Canudo, J. I., Le Loeuff, J., Dinars-Turell, J., Riera, V., Oms,
O., Tortosa, T., & Gaete, R. (2012). The diversity of sauropod dinosaurs and their
EP
AC
C
Vilas, L., Mas, R., Garca, A., Alonso, A., Melndez, N., & Rincn, R. (1982). Ibrica
Suroccidental. In Garca, A. (Ed.), El Cretcico de Espaa (pp. 457-508). Madrid:
Universidad Complutense de Madrid.
Wedel, M. J. (2003). The evolution of vertebral pneumaticity in sauropod dinosaurs.
Journal of Vertebrate Paleontology, 23(2), 344-357.
ACCEPTED MANUSCRIPT
Wilson, J. A. (1999). A nomenclature for vertebral laminae in sauropods and other
saurischian dinosaurs. Journal of Vertebrate Paleontology, 19, 639-653.
Wilson, J. A. (2002). Sauropod dinosaur phylogeny: critique and cladistic analysis.
Zoological Journal of the Linnean Society, 136, 217-276.
RI
PT
Wilson, J. A. (2012). New vertebral laminae and patterns of serial variation in vertebral
SC
M
AN
U
Wilson, J. A., DEmic, M. D., Ikejiri, T., Moacdieh, E. M., & Withlock, J. A. (2011). A
nomenclature for vertebral fossae in sauropods and other saurischian dinosaurs.
PLoS One, 6(2), e17114. http://dx.doi.org/10.1371/journal.pone.0017114.
Wilson, J. A., & Sereno, P. C. (1998). Early evolution and higher-level phylogeny of
TE
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Zurriaguz, V., & Powell, J. (2015). New contributions to the presacral osteology of
Saltasaurus loricatus (Sauropoda, Titanosauria) from the Upper Cretaceous of
FIGURE CAPTIONS
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Fig. 1. Geographic and geologic setting of the palaeontological site of Lo Hueco:
general geographic location (a); detailed geological situation: sections of the Villalba de
la Sierra Formation (b, left), and of the site (b, right); lateral view of the eastern talus of
the outcrop during the excavation phase, with the identified litosomes (c); type
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cartography of one of the excavation areas (G1) at Lo Hueco (e); and map of the
distribution of bones of the holotype specimen in field (f). The red dot marks the
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location of the holotype in the field (c) in the stratigraphic column (b); and excavation
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map (e).
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Fig. 3. Caudal vertebrae and haemal arches of Lohuecotitan pandafilandi type specimen
(HUE-EC-01). Anterior caudal vertebrae: 1st and 2nd caudal vertebra (HUE-03134/6) in
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right (a), anterior (b) and posterior (c) view; 7th and 8th caudal vertebra (HUE-03134/1)
showing a pathological central fusion in right view (c); 9th caudal vertebra (HUE-
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03101) in left view (e). Middle caudal vertebrae: HUE-03028 in left view (f); HUE03028 in anterior (g), posterior (h) and right (i) views. Scale bar: 5 cm.
Fig. 4. Middle haemal arch (HUE-3041) in left and posterior view and a posterior
haemal arch (HUE-3040) in dorsal (f) and left (g) views. Scale bar: 5 cm.
Fig.5. Sacrum and anterior sector of the Lohuecotitan pandafilandi type specimen
(HUE-EC-01) in articulation (a). Appendicular bones of L. pandafilandi type specimen
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(HUE-EC-01): left ulna (HUE-03044) in lateral (b) and proximal (c) views; right femur
(HUE-03108) in distal (d) and posterior (g) views; right ischium (HUE-03099) in lateral
view (e); left pubis (HUE-03086) in anterior view (f); right fibula (HUE-03087) in
proximal (h) and lateral (j) views; right tibia (HUE-03082) in proximal (i) and posterior
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obtained from Carballido and Sander (2014) data matrix. Numbers without brackets
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indicate bootstrap values, and those within brackets indicate the Bremer support.
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Fig. 3. Caudal vertebrae and haemal arches of Lohuecotitan pandafilandi type specimen
(HUE-EC-01). Anterior caudal vertebrae: 1st and 2nd caudal vertebra (HUE-03134/6) in
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right (a), anterior (b) and posterior (c) view; 7th and 8th caudal vertebra (HUE-03134/1)
showing a pathological central fusion in right view (d); 9th caudal vertebra (HUE-
03101) in left view (e). Middle caudal vertebrae: HUE-03028 in left view (f); HUE-
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03028 in anterior (g), posterior (h) and right (i) views. Scale bar: 5 cm.
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middle dorsal vertebrae (HUE-03135) in right view (c); two fragments of a middle
dorsal vertebra (HUE-03135) in right view: neural spine (d) and centrum (e). Scale bar:
5 cm.
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Fig. 3. Caudal vertebrae and haemal arches of Lohuecotitan pandafilandi type specimen
(HUE-EC-01). Anterior caudal vertebrae: 1st and 2nd caudal vertebra (HUE-03134/6) in
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right (a), anterior (b) and posterior (c) view; 7th and 8th caudal vertebra (HUE-03134/1)
showing a pathological central fusion in right view (d); 9th caudal vertebra (HUE03101) in left view (e). Middle caudal vertebrae: HUE-03028 in left view (f); HUE-
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03028 in anterior (g), posterior (h) and right (i) views. Scale bar: 5 cm.
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