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Accepted Manuscript

A new titanosaur (Dinosauria, Sauropoda) from the Upper Cretaceous of Lo Hueco (Cuenca, Spain)

V. Díez Díaz, P. Mocho, A. Páramo, F. Escaso, F. Marcos-Fernández, J.L. Sanz, F. Ortega

PII:

S0195-6671(16)30151-3

Reference:

YCRES 3431

To appear in:

Cretaceous Research

Received Date: 6 May 2016

Revised Date:

Accepted Date: 2 August 2016

24 July 2016

2016 Revised Date: Accepted Date: 2 August 2016 24 July 2016 Please cite this article as:

Please cite this article as: Díez Díaz, V., Mocho, P., Páramo, A., Escaso, F., Marcos-Fernández, 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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ACCEPTED MANUSCRIPT

A new titanosaur (Dinosauria, Sauropoda) from the Upper Cretaceous of

Lo Hueco (Cuenca, Spain)

V. Díez Díaz a , P. Mocho b , A. Páramo b, c , F. Escaso

Sanz c , F. Ortega b*

b , F. Marcos-Fernández

d, b

, J.L.

a Museum für Naturkunde - Leibniz Institute for Evolution and Biodiversity Science.

Invalidenstrasse 43, 10115 Berlin, Germany.

b Grupo de Biología Evolutiva. Facultad de Ciencias. UNED. Senda del Rey 9 28040.

Madrid, Spain.

c Unidad de Paleontología, Departamento de Biología, Universidad Autónoma de

Madrid, Calle Darwin 2, 20049 Madrid, Spain.

d Facultad de Bellas Artes, Universidad Complutense de Madrid. Calle Pintor el Greco

2, 28040 Madrid.

* Corresponding author

E-mail address: fortega@ccia.uned.es

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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

new titanosaurian sauropod is established, Lohuecotitan pandafilandi. This titanosaur is

diagnosed by eight autapomorphic features: dorsally and ventrally widened or

bifurcated posterior centrodiapophyseal lamina in anterior and middle dorsal vertebrae;

short postspinal lamina with a transversely expanded distal end represented by smooth

scars in the dorsal vertebrae; anteriormost caudals with the medial

spinoprezygapophyseal and medial spinopostzygapophyseal laminae ventrally

connected with the prespinal and postspinal laminae, respectively; anterior caudal

neural spines with a dorsal projection of the prespinal and postspinal laminae; anterior

caudal neural spines bears a “greek-cross”-like cross-section; middle caudal centra

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

and the lateral trochanter of the fibula. The herein performed phylogenetic analysis

considered L. pandafilandi as a member of Lithostrotia more derived than

Malawisaurus. The known palaeodiversity of the Late Cretaceous Ibero-Armorican

titanosaurs is increasing, and furher analyses focused on this group will be necesary to

better understand the evolutionary history of European titanosaurs and to clarify their

relationships within Titanosauria.

Keywords: Lohuecotitan pandafilandi gen. et sp. nov., Titanosauria, Lithostrotia, Spain,

Late Cretaceous

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1. Introduction

Titanosaurs had a widespread distribution during the Late Cretaceous, being

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

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-

Maastrichtian of the Ibero-Armorican Island (South France and Iberian Peninsula):

Lirainosaurus astibiae Sanz et al., 1999, Ampelosaurus atacis Le Loeuff, 1995, and

Atsinganosaurus velauciensis Garcia et al., 2010. Nevertheless, a higher diversity of

European titanosaurs has been hypothesized based on the disparity in the morphologies

of teeth (Díez Díaz et al. 2012, 2013a, 2014) and appendicular remains (Vila et al.,

2012; Páramo et al., 2014, 2015a, b).

The titanosaurian record from the Upper Cretaceous of southern France has been

known since the last years of the nineteenth century (Depéret, 1899, 1900). The

discovery in the last decades of more than 20 sites with sauropod remains has greatly

improved the knowledge of the French titanosaurian sauropod faunas (e.g. Le Loeuff,

1995, 2005; Garcia et al., 2010; and see Díez Díaz, 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-Rhône

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 Díez Díaz, 2013 for a

complete list). The first titanosaurian remains found in the Iberian Peninsula were

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

Peninsula: Barcelona (Sellés et al., 2015), Cuenca (Ortega et al., 2008, 2015),

Guadalajara (Ortega and Pérez-García, 2009), Huesca (Canudo, 2001), Lleida

(Casanovas-Cladellas and Santafé-Llopis, 1993; Casanovas et al., 1987; Casanovas-

Cladellas et al., 1995; Masriera and Ullastre, 1987), Segovia (Pérez-García et al., 2015;

Sanz and Buscalioni, 1987), Soria (Lapparent et al., 1957; Pereda Suberbiola and Ruiz-

Omeñaca, 2001), and Valencia (Company et al., 2009; Company and Pereda-

Suberbiola, 2013).

One of the most outstanding sites is the Laño quarry (upper Campanian-lower

Maastrichtian, northern Spain) that has yielded numerous cranial and postcranial

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

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 (Díez Díaz 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

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

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.

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

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

anatomical connection or with a low dispersion of their skeletal elements. Up to now,

the comparison of these partial skeletons with isolated remains indicates the presence of

at least two different titanosaurian morphotypes based on two types of cranial

morphologies, appendicular bones and teeth (Knoll et al., 2013, 2015; Díez Díaz et al.,

2014; Páramo et al., 2014, 2015a, b; Ortega et al., 2015).

This discovery sheds light on the knowledge of the systematics and diversity of

the titanosaurs in the Ibero-Armorican Island during the Late Cretaceous, and it

corroborates the previous hypothesis of the presence of a higher diversity of titanosaurs

in southwestern Europe at the end of the Cretaceous (see e.g. Díez Díaz et al., 2013a).

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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

evolutionary history of the Ibero-Armorican titanosaurs and the phylogenetic

relationships of these forms within Titanosauria.

2. Material and methods

For the anatomical structures we use “Romerian” terms (Wilson, 2006) for the

structures (e.g. “centrum”) and their orientation (e.g. “anterior”). The nomenclature

used for the caudal vertebrae is the one proposed by Díez Díaz 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

proposed by Otero et al. (2012).

Institutional abbreviations. HUE: Lo Hueco collection, housed at the Museo de

Paleontología de Castilla-La Mancha in Cuenca, Spain; MDE: Musée des Dinosaures,

Espéraza, France.

Anatomical abbreviations. *, autapomorphy; 4 th , fourth trochanter; acdl, anterior

centrodiapophyseal lamina; acet, acetabulum; acpl, anterior centroparapophyseal

lamina; act, alp, anterolateral process of the ulna; amp, anteromedial process of the ulna;

accessory trochanter; at, anterior trochanter; cc, cnemial crest; cd, caudal vertebra; cpol,

centropostzygapophyseal lamina; cprf, centroprezygapophyseal fossa; cprl,

centroprezygapophyseal lamina; dart, doble articulation; di, diapophysis; dbi, dorsal

birfucation of the pcdl; ep, epicondyle; fia, fibular articulation; fic, fibular condyle;

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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,

postzygapophysis; podl, postzygadiapophyseal lamina; posl, postspinal lamina; pp,

posterior process of the ulna; pped, pubic peduncle; prsl, prespinal lamina; raa, radial

articulation; rpr, round process; sdf, spinodiapophyseal lamina; spdl, spinodiapophyseal

lamina; spol, spinopostzygapophyseal lamina; sprf, spinoprezygapophyseal fossa; sprl,

spinoprezygapophyseal lamina; tap, triangular aliform process; tb, tuberosity; tic, tibial

condyle; tprl, intraprezygapophyseal lamina; tpol, intrapostzygapophyseal lamina; ts,

trochanteric shelf; vbi, ventral birfucation of the pcdl.

3. Systematic palaeontology

Dinosauria Owen, 1842

Saurischia Seeley, 1888

Sauropoda Marsh, 1878

Titanosauriformes Salgado et al., 1997

Titanosauria Bonaparte and Coria, 1993

Lithostrotia Upchurch et al., 2004

Lohuecotitan gen. nov.

Type and only included species. Lohuecotitan pandafilandi sp. nov.

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Etymology. Lohuecotitan from Lo Hueco (the type locality) and titan (the giants of the

Greek mythology).

Type locality and horizon. As for type and only species.

Diagnosis. See diagnosis for type and only species below.

Lohuecotitan pandafilandi gen. et sp. nov.

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

vertebrae (HUE-03088, 03134/7, 03135); dorsal ribs (HUE-03045, 03081, 10017);

sacral vertebrae fragments (HUE-03134); 20 caudal vertebrae (HUE-03020-29, 03052,

03101, 03114, 03134/1-6); haemal arches (HUE-03032-34, 03040, 03041, 03054,

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-

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

Mancha, central Spain) (Fig. 1), Margas, Arcillas y Yesos de Villalba de la Sierra

Formation, upper Campanian-lower Maastrichtian.

Diagnosis. Member of Titanosauria, having the following autapomorphies (marked with

an asterisk), as well as an unique combination of characters not seen in other

titanosaurs: (1) posterior centrodiapophyseal lamina (pcdl) dorsally and ventrally

widened (sometimes bifurcated) simultaneous in anterior and middle dorsal vertebrae

(shared with Saltasaurus); (2) short postspinal lamina (posl) with a transversely

expanded distal end represented by smooth scars in the dorsal vertebrae*; (3)

anteriormost caudal vertebrae with the medial spinoprezygapophyseal lamina (med.

sprl) and medial spinopostzygapophyseal lamina (med. spol) ventrally connected with

the prespinal lamina (prsl) and posl, respectively*; (4) anterior neural spines on caudal

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

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

tissue, as occur in titanosauriforms (Upchurch, 1998; Wilson, 2002; Wedel, 2003;

Upchurch et al., 2004; D’Emic, 2012). The centra are opisthocoelous and bear a

subcircular cross-section. Shallow pneumatic fossae appear in the lateral surfaces of the

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

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; D’Emic, 2012). The anterior and

middle sector of the ventral surface of the centra is slightly transversely concave

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

prsl is present in the preserved neural spine. No complete neural spines are preserved,

and the morphology of the distal tip is unknown.

Lohuecotitan pandafilandi type specimen preserves four anterior to middle

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

of Lohuecotitan is the presence of a dorsally and ventrally widened (sometimes

bifurcated) pcdl, that is considered as a local autapomorphy of this taxon, as it is also

shared by some of the dorsal vertebrae of Saltasaurus (Zurriaguz and Powell, 2015).

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

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

titanosaurs Lirainosaurus (Díez Díaz et al., 2013b), Opisthocoelicaudia (Borsuk-

Bialynicka, 1977) and Elaltitan (Mannion and Otero, 2012), but also in other non-

derived titanosaurs (Wilson, 1999). The neural spine is posteriorly deflected (more than

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

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

titanosauriforms, including Lirainosaurus (Díez Díaz et al., 2013b) and Rapetosaurus

(Curry Rogers, 2009). The middle and distal part of the preserved dorsal ribs are

transversally compressed and have a plank-like morphology, features also common in

titanosauriforms (Wilson and Sereno, 1998).

Only some fragments of the sacral vertebrae are present. There are, at least, five

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.

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;

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

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; D’Emic, 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

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

posl, respectively. This condition is considered a diagnostic feature for Lohuecotitan

pandafilandi. Another peculiar feature is the presence of a dorsal projection of the prsl

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

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

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

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

in the posterior haemal arches is not known in other European titanosaurs and is here

considered a diagnostic feature of Lohuecotitan pandafilandi, although this condition is

also present in the South American Aeolosaurini (e.g. Casal et al., 2007; Santucci and

Arruda-Campos, 2011) and some euhelopodids (D’Emic, 2012). The haemal canal is ca.

50% of the total length of the most anterior haemal arch, as occurs in most

titanosauriforms (e.g. Wilson, 2002; Upchurch et al., 2004; D’Emic, 2012; Mannion et

al., 2013; Mocho et al., 2014).

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

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.

The proximal and distal ends are not fully preserved. The anterior process seems to be

much larger than the lateral one.

The ilium (Fig. 5a) is internally pneumatized (camellate bone tissue), and the

preacetabular process is perpendicular to the sagittal plane and is subhorizontal. It

reaches its greatest height above the pubic peduncle. The ventral margin of the

preacetabular process is slightly concave. Two ischia are preserved, with the right one

almost complete (Fig. 5e). The ischia are shorter than the pubes and the ischiatic blade

is transversely flat, resulting in a plate-like morphology. The ischiatic blade lacks an

emargination on its anterior margin. A lateral tuberosity is present near the posterior

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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

common in titanosaurs (Wilson, 2002; Upchurch et al., 2004; González 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

ridge (and associated groove) are present in the lateral surface.

The shaft of the femur (Fig. 5d, g) is straight and anteroposteriorly compressed

with an elliptical cross-section (mediolateral/anteroposterior width of the femoral

diaphysis = 1.6). On the posterior surface a wide trochanteric shelf extends from the

greater trochanter. The proximolateral margin of the femur is deflected medially and

bears a well-developed lateral bulge. The fourth trochanter is placed in the posterior

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 D-

shaped profile, and the cnemial crest is anterolaterally directed, specially its most distal

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

than anteroposteriorly, as typical of titanosaurs (Salgado et al., 1997; Upchurch et al.,

2004). The shaft of the fibula (Fig. 5h, j) is slightly sigmoidal in lateral view, but not as

pronounced as in other titanosaurs such as Opisthocoelicaudia (Borsuk-Bialynicka,

1977) and Saltasaurus (Powell, 1992). The tibial articular facet is well marked and

occupies ¼ of the fibular total length. An anterior crest is present in the proximal end

and is medially directed. The lateral trochanter is located above the mid-length of the

fibula and is formed by two subparallel ridges delimiting a shallow oval fossa. Between

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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).

4. Phylogenetic analysis

To establish the phylogenetic relationships of Lohuecotitan pandafilandi a

phylogenetic analysis was performed using the data matrix proposed by Carballido and

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

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

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

considered ordered.

The result of this analysis yielded 20 MPTs of 1004 steps with a consistency

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 40º) 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).

The present topology also recovered as possible autapomorphies the following

characters: i) cervical pleurocoels divided by a septum (character #115); ii) dorsal

neural spines with subparallel margins (character #138); iii) in anterior view, neural

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

middle and posterior dorsal centrum (character #162); vi) absence of spdl and spol

contact in middle dorsals (character #169); vii) femoral transverse diameter is 125-

150% of the anteroposterior diameter (character #301).

Previous phylogenetic analyses show that relationships of the Ibero-Armorican

titanosaurs remain uncertain (e.g. Curry Rogers, 2005; Mannion and Upchurch, 2011;

Díez Díaz, 2013; Gorscak and O’Connor, 2016). Future analyses focused on European

titanosaurian taxa will be important to better understand the evolutionary history of this

group in the European territory and to clarify the relationships of these taxa with non-

European titanosaurs. The results obtained here support the presence of a new

lithostrotian titanosaur in the Ibero-Armorican Island during the Late Cretaceous.

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,

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

Lohuecotitan and Atsinganosaurus. Atsinganosaurus has a prominent anterodorsal-

posteroventral lamina on the spinodiapophyseal fossa (sdf), dividing it into two

subfossae, feature that it is not present in Lohuecotitan

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; Apesteguía,

2005; pers. obs., PM). A hyposphenic crest is common in basal titanosaurs such as

Andesaurus, Argentinosaurus or Epachthosaurus (e.g. Apesteguía, 2005), and contrary

to other European titanosaurs, such as Lirainosaurus where this crest is absent.

Nevertheless, the presence of a hyposphene-hypantrum complex is not possible to test

in Lohuecotitan due the poor preservation of the dorsal vertebrae. Lohuecotitan presents

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 (Díez Díaz 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

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

spine. Lohuecotitan shares with Lirainosaurus the presence of deep fossae below the

prezygapophyses (cprf, observed in anterior view) in the proximal anterior caudal

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.

Dorsoventrally expanded prezygapophyses that characterize Lohuecotitan are absent in

Lirainosaurus, Atsinganosaurus, Magyarosaurus and Ampelosaurus.

From the European record, only Ampelosaurus and Paludititan preserve

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

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)

differs from that of Lirainosaurus and Atsinganosaurus by having a more rounded

outline of the proximal articulation, and not showing a prominent anteromedial ridge

close to the distal extremity delimiting two highly concave surfaces, features diagnostic

of these last taxa (Díez Díaz et al., 2013c). The fibula of Lohuecotitan has a shaft

slightly more sigmoidal than that of Lirainosaurus and Ampelosaurus. However,

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

morphological variability in the appendicular material is observed in Bellevue site. In

Lo Hueco this variability might represent at least two different taxa (Ortega et al.,

2015), and future analysis in the material of the Ampelosaurus type locality might

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.

Other important fossil sites from the Upper Cretaceous of the Iberian Peninsula

are located in Laño (Treviño 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 Laño site, only one titanosaurian taxon is identified (Díez Díaz et al., 2015).

From Chera, two morphotypes were identified, one of them assigned to Lirainosaurus

(Company et al., 2009; Díez Díaz 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

the tibiae is clearly different, although both present a rounded profile of the proximal

articulation, in contrast to Lirainosaurus.

The discovery of Lohuecotitan pandafilandi provides relevant information for

understanding the titanosaurian evolutionary history and diversity during the Late

Cretaceous in Europe. After almost 20 years since the publication of Lirainosaurus

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.

It is important to note that in the Lo Hueco site two different morphotypes of

titanosaurian teeth and braincases have been found (Díez Díaz et al., 2014; Knoll et al.,

2013, 2015; Páramo 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,

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

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 Fox-

Amphoux-Métisson (Díez Díaz et al., 2012) and Massecaps, where a new titanosaur has

been found and its description is in progress (Díez Díaz 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.

6. Conclusion

A new titanosaur, Lohuecotitan pandafilandi is described from the Upper

Cretaceous of the Ibero-Armorican Island. This new taxon came from the Lo Hueco

fossil site (upper Campanian-lower Maastrichtian). This taxon is diagnosed by eight

autapomorphic features highlighting the presence of short posl with a transversely

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.

Lohuecotitan is placed within Lithostrotia, and this position is supported by the

presence of anterior and middle dorsal vertebrae with strongly and posteroventrally

oriented (more than 40º) zygapophyseal articulation. Lohuecotitan is considered a more

derived lithostrotian than Malawisaurus and is recovered in a polytomy with

Opisthocoelicaudia, Alamosaurus, Trigonosaurus, Saltasaurinae and Nemegtosauridae.

The phylogenetic relationships of the Ibero-Armorican titanosaurs remain uncertain.

Future analyses focused on European titanosaur taxa will be important to better

understand the evolutionary history of this group in the European territory and to clarify

the relationships of these taxa with non-European titanosaurs.

Previous works confirm a high titanosaurian diversity in the Ibero-Armorican

Island summarizing the recent discoveries from Spain and France: from one known

titanosaurian taxon in Spain (Lirainosaurus) and two in France (Ampelosaurus and

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Atsinganosaurus) to, at least, six or seven taxa (Díez Díaz 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.

7. Acknowledgements

This research was supported by the projects CGL2012-35199 and CGL2015-68363-P of

the Spanish Ministerio de Economía y Competitividad; and the project

SBPLY/15/180601/000045 of the Junta de Comunidades de Castilla-La Mancha. The

holotype HUE-EC1 described in this paper was collected under permission obtained

from the Dirección General de Patrimonio y Museos of the Junta de Comunidades de

Castilla-La Mancha (ref. 04-0392-P11). This research was also supported by

SFRH/BD/68450/2010 PhD scholarship (PM), financed by the “Fundação para a

Ciência e Tecnologia” (Portugal); a Synthesys Project

(http://synthesys3.myspecies.info/) which is financed by the European Community

Research Infrastructure Action under the FP7 (FR-TAF-5072) (PM); and the Spanish

Government Grant (ref. BES-2013-065509) (AP). We thank J. C. Corral and J. Alonso

(Museo de Ciencias Naturales de Alava/Arabako Natur Zientzien Museoa, Vitoria-

Gasteiz, Spain), B. Madarieta (Museo Vasco de Historia de la Medicina y de las

Ciencias of Leioa, Spain), J. Company (Universidad Politécnica de Valencia, Spain), J.

Le Loeuff (Musée des Dinosaures d’Espéraza, France), G. Garcia and X. Valentin

(IPHEP, Université de Poitiers, France), P. Barrett and S. Chapman (Natural History

Museum, U.K.), R. Allain (Muséum national d'Histoire naturelle, France), D. Schwarz

and O. Hampe (HNM, Germany). Special thanks to C. de Miguel Chaves (UNED), I.

Nárvaez (UNED) and A. Elvira (UNED) for help in the preparation of the specimens

and suggestions. The Willi Hennig Society sponsors the use of the TNT cladistics

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software. The comments made by L. Salgado, P. Mannion, and the editor improved an

early version of this paper.

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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

specimen of Lohuecotitan pandafilandi (HUE-EC-01) in the field (d); general

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

location of the holotype in the field (c) in the stratigraphic column (b); and excavation

map (e).

Fig. 2. Presacral vertebrae of Lohuecotitan pandafilandi type specimen (HUE-EC-01).

Anterior cervical vertebra (HUE-03128) in left (a) and anterior (b) views; anterior-

middle dorsal vertebrae (HUE-03135) in right view (c); middle dorsal vertebra (HUE-

03135) in right view (d). Scale bar: 5 cm.

Fig. 3. Caudal vertebrae and haemal arches of Lohuecotitan pandafilandi type specimen

(HUE-EC-01). Anterior caudal vertebrae: 1

st

and 2 nd caudal vertebra (HUE-03134/6) in

right (a), anterior (b) and posterior (c) view; 7 th and 8 th caudal vertebra (HUE-03134/1)

showing a pathological central fusion in right view (c); 9 th caudal vertebra (HUE-

03101) in left view (e). Middle caudal vertebrae: HUE-03028 in left view (f); HUE-

03028 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

(k) views. Scale bar: 10 cm.

Fig. 6. Lohuecotitan pandafilandi phylogeny. Strict consensus of 20 MPTs of 1004

steps with a consistency index (CI) of 0.405 and a retention index (RI) of 0.735

obtained from Carballido and Sander (2014) data matrix. Numbers without brackets

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: 1 st and 2 nd caudal vertebra (HUE-03134/6) in

right (a), anterior (b) and posterior (c) view; 7

th

and 8

th

caudal vertebra (HUE-03134/1)

showing a pathological central fusion in right view (d); 9

th

caudal vertebra (HUE-

03101) in left view (e). Middle caudal vertebrae: HUE-03028 in left view (f); HUE-

03028 in anterior (g), posterior (h) and right (i) views. Scale bar: 5 cm.

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Fig. 2. Presacral vertebrae of Lohuecotitan pandafilandi type specimen (HUE-EC-01).

Anterior cervical vertebra (HUE-03128) in left (a) and anterior (b) views; anterior-

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.

Fig. 3. Caudal vertebrae and haemal arches of Lohuecotitan pandafilandi type specimen

(HUE-EC-01). Anterior caudal vertebrae: 1

st

and 2

nd

caudal vertebra (HUE-03134/6) in

right (a), anterior (b) and posterior (c) view; 7

th

and 8

th

caudal vertebra (HUE-03134/1)

showing a pathological central fusion in right view (d); 9

th

caudal vertebra (HUE-

03101) in left view (e). Middle caudal vertebrae: HUE-03028 in left view (f); HUE-

03028 in anterior (g), posterior (h) and right (i) views. Scale bar: 5 cm.

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