Quetecsaurus - Gonzalez Riga Ortiz David 2014

AMEGHINIANA - 2014 - Tomo 51 (1): 3 – 25 ISSN 0002-7014
A NEW TITANOSAUR (DINOSAURIA, SAUROPODA)

FROM THE UPPER CRETACEOUS (CERRO
LISANDRO FORMATION) OF MENDOZA
PROVINCE, ARGENTINA
BERNARDO J. GONZÁLEZ RIGA1,2 AND LEONARDO ORTIZ DAVID1
1
Laboratorio de Dinosaurios, Instituto de Ciencias Básicas, Universidad Nacional de Cuyo, Avenida Padre Contreras 1300, Parque General San Martín, 5500 Mendoza,
Argentina.
2
Departamento de Paleontología, Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales, Consejo Nacional de Investigaciones Científicas y Técni-
cas, Centro Científico Tecnológico, Avenida Ruiz Leal s/n, Parque General San Martín, 5500 Mendoza, Argentina.
[email protected].; [email protected]
Abstract. Quetecsaurus rusconii gen. et sp. nov. is a new titanosaur (Dinosauria, Sauropoda) from the Neuquén Basin of Mendoza Province,
Argentina. The specimen comes from red mudstones of the Cerro Lisandro Formation (middle-late Turonian, Upper Cretaceous), and is the
first sauropod with well-preserved remains to be discovered in this formation. The holotype includes a postorbital, teeth, the atlas, a poste-
rior cervical vertebra, an incomplete dorsal vertebra, a posterior caudal centrum, dorsal ribs, a coracoid, fragments of a humerus, radius and
ulna, and five metacarpals. It is characterized by the following combination of autapomorphies: (1) intercentrum of atlas with a prominent
anteroventral border and expanded posteroventral processes; (2) posterior cervical neural spines with incipient lateral expansions; and (3)
humerus with strongly sigmoid proximal border, rounded proximomedial border, and angular proximolateral corner. A preliminary cladis-
tic analysis placed Quetecsaurus as a sister taxon of Lognkosauria (Mendozasaurus + Futalognkosaurus). Quetecsaurus shares with the lognkosaurs
the presence of cervical neural spines with ‘lateral laminae’, but relatively reduced in comparison with those taxa. This discovery provides new
information on the diagnosis of Lognkosauria within South American titanosaurs.
Key words . Quetecsaurus. Titanosauria. Cerro Lisandro Formation. Cretaceous. Mendoza. Argentina.
Resumen. UN NUEVO TITANOSAURIO (DINOSAURIA, SAUROPODA) DEL CRETÁCICO SUPERIOR (FORMACIÓN

CERRO LISANDRO) DE LA PROVINCIA DE MENDOZA, ARGENTINA. Quetecsaurus rusconii gen. et sp. nov. es un nuevo tita-
nosaurio (Dinosauria, Sauropoda) hallado en la Cuenca Neuquina, Provincia de Mendoza, Argentina. El espécimen proviene de fango-
litas rojas de la Formación Cerro Lisandro (Turoniano medio-tardío, Cretácico Superior) y es el primer saurópodo con huesos bien
preservados descubierto en esta formación. El holotipo comprende un postorbital, dientes, el atlas, una vértebra cervical posterior, una
vértebra dorsal incompleta, un centro caudal anterior, costillas dorsales, un coracoides, fragmentos de húmero, radio y ulna, y cinco me-
tacarpos. Se caracteriza por la siguiente asociación de autapomorfías: (1) intercentro del atlas con un borde anteroventral prominente y
procesos posteroventrales expandidos; (2) espinas neurales cervicales posteriores con láminas laterales incipientes; y (3) húmero con un
borde proximal fuertemente sigmoidal, borde proximomedial redondeado y esquina proximolateral angular. Un análisis cladístico pre-
liminar ubica a Quetecsaurus como un taxón hermano de Lognkosauria (Mendozasaurus + Futalognkosaurus). Quetecsaurus comparte con
los lognkosaurios la presencia de espinas neurales cervicales con láminas laterales, pero relativamente reducidas en comparación con esos
taxa. Este descubrimiento brinda nueva información sobre la diagnosis de Lognkosauria, dentro de los titanosaurios de América del Sur.
Palabras clave. Quetecsaurus. Titanosauria. Formación Cerro Lisandro. Cretácico. Mendoza.
TITANOSAURS comprise the most diverse and abundant clade Brazil, and one from Chile) (see González Riga, 2010; Man-
of sauropod dinosaurs from the Upper Cretaceous. Their fos- nion and Otero, 2012). Titanosaurs show an amazing diversity
sils have been found on all continents and their evolution and encompassed several mega-herbivorous adaptive types
during the breakup of Gondwana has considerable paleo- during the Late Cretaceous (Curry Rogers, 2005; Wilson,
biogeographic relevance (e.g., Le Loeuff, 1993; Casanovas- 2006). The smallest titanosaurs, such as Saltasaurus loricatus
Cladellas et al., 1993; Canudo et al., 2008). There are Bonaparte and Powell, 1980, and Neuquensaurus australis (Ly-
approximately 60 titanosaur (and somphospondylian ti- dekker, 1893) Powell, 1986, reached only 7 m of total length;
tanosauriforms) species recognized across the world. The fos- by contrast, giant forms such as Argentinosaurus huinculensis
sil record of these sauropods is particularly abundant in South Bonaparte and Coria, 1993, and Puertasaurus reuili Novas,
America (with 32 taxa known from Argentina, eight from Salgado, Calvo, and Agnolin, 2005, exceeded 25 m, and were
AMGHB2-0002-7014/12$00.00+.50 3
AMEGHINIANA - 2014 - Tomo 51 (1): 3 – 25
possibly the heaviest terrestrial vertebrates that ever existed GEOLOGICAL SETTING
(~73 metric tons for Argentinosaurus according to Mazzetta et In Neuquén Province, northern Patagonia, the Cerro
al., 2004). Lisandro Formation has provided a rich fossil record that in-
In South America, the most abundant titanosaur record comes cludes freshwater bivalves represented by the genus Diplodon
from the Neuquén Basin in northwestern Patagonia. Most of (Manceñido and Damborenea, 1984; Garrido, 2000; Leanza
these titanosaurs have been found in Upper Cretaceous strata et al., 2004; Salgado et al., 2009), lungfish remains (Ceratodus
of the Neuquén Group and the Allen Formation (Leanza et sp.), osteichthyan vertebrae assigned to Lepisosteidae (Coria
al., 2004; de la Fuente et al., 2007; González Riga, 2010; et al., 1996; Leanza et al., 2004; Apesteguía et al., 2007; Sal-
Calvo et al., 2011). Although their record is diverse, titanosaur gado et al., 2009), and teeth and bones of crocodiles and tur-
remains are still unknown from some stratigraphic intervals in tles (Garrido, 2000; Salgado et al., 2009). Dinosaur remains
the Neuquén Basin. For example, until this paper, there was no include those of ornithischians (Anabisetia saldiviai Coria and
record of well-preserved titanosaur fossils from the Cerro Calvo, 2002; Iguanodontia indet. Coria et al., 1996) and
Lisandro Formation (middle-late Turonian; Garrido, 2010), theropods (Coria et al., 1996, Coria and Currie, 2006). Sauro-
and only fragmentary titanosaur bones and teeth were known pod fossils are scarce and limited to isolated teeth and frag-
from this unit (Salgado et al., 2009). mentary bones (Salgado et al., 2009).
The aim of this paper is to describe a new genus and species The Cerro Lisandro Formation is widely exposed in
of titanosaur, Quetecsaurus rusconii, from the middle-late Neuquén Province and is composed of fine clastic deposits,
Turonian in Cerro Lisandro Formation of Mendoza Province mainly monotonous banks of red mudstones interbedded with
near the northwestern border of Patagonia. This discovery thin levels of fine, yellow and gray-green sandstones (Garrido,
provides new morphological information relevant to the un- 2010). This facies association is interpreted to represent distal
derstanding of titanosaur evolution during the Cretaceous. mud flats of fluvial systems (Leanza and Hugo, 2001). The
Quetecsaurus is the third sauropod discovered in Mendoza; mudstones indicate a dominance of illite and interstratified il-
the others are Mendozasaurus neguyelap González Riga, 2003 lite-smectite, with secondary involvement of kaolinite and
and Malarguesaurus florenciae González Riga, Previtera and chlorite (Giusiano and Pettinari, 1999; Pettinari et al., 1999).
Pirrone, 2009. These dinosaurs come from middle levels of Quetecsaurus comes from Cañada del Pichanal, in the
the Neuquén Group. Malarguesaurus was found in strata as- southern end of Mendoza Province, near the Río Colorado
signed to Los Bastos Formation (early-middle Coniacian) and very close to Neuquén Province (Fig. 1). In this region, the
whereas Mendozasaurus was discovered in the upper levels of Cerro Lisandro Formation is the oldest unit of the Neuquén
the Sierra Barrosa Formation (middle-late Coniacian). Other Group, and it crops out in low anticlinal and synclinal struc-
titanosaur remains found in Mendoza Province include a dis- tures that are generally North-South oriented (González Riga,
tal caudal series from “Cañadón Amarillo” (Wilson et al., 2002).
1999). Finally, exceptionally preserved articulated specimens At the Quetecsaurus site, the dominant facies are massive
were recently discovered in upper levels of the Plottier For- and laminated red-purple mudstones. The presence of clay
mation (late Coniacian-early Santonian) (González Riga et al., minerals is easily recognizable by flooded soils and sparse veg-
2012, 2013). etation. The facies association includes massive mudstones and
The specimen described herein was collected during the thin, lenticular gray shales and sandstones. Fibrous gypsum
development of the mining project ‘Proyecto Potasio Río beds are also common. The sequence represents a muddy
Colorado’ (Vale S.A.). The fieldwork, led by the first author floodplain where clay sediments accumulated in ephemeral
(B.G.R.), was carried out by a team of paleontologists and bodies of water. In some areas there are reactivation surfaces
technicians in strict compliance with laws pertaining to the (lag deposits) that represent episodes of flooding, drying, and
preservation of the paleontological heritage of Argentina. This reworking. The contact with the Portezuelo Formation is lo-
paper is partly based on the first description of the specimen, cated 2 km east of the fossil site, and is easily identified by the
which was part of the degree thesis of the second author presence of yellow, coarse sandy channel beds.
(L.O.D.). Within the Cerro Lisandro Formation, isolated sand bod-
4
GONZÁLEZ RIGA AND ORTIZ DAVID: NEW TITANOSAUR
Figure 1. 1, Locality map of the Neuquén Basin showing La Cañada del Pichanal, the site where fossils of Quetecsaurus rusconii were recovered.
2, Geological section showing the stratigraphic level of Quetecsaurus rusconii.
ies up to 50 m in width are present, and represent fluvial chan- turing and plastic deformation due to lithostatic pressure. In
nels with low to moderate lateral migration. These sand bod- contrast, the cervical and dorsal vertebrae were less affected,
ies are well exposed near the Pata Mora bridge, on the southern and preserve their original anatomical shape. From a system-
margin of the Río Colorado, 20 km east of Cañada del atic viewpoint, it is important to mention that all the skeletal
Pichanal. elements were associated and belong to the same specimen.
The Quetecsaurus specimen consists of cranial remains, the No duplicate bones were found (Fig. 2).
atlas, a cervical vertebra, a dorsal vertebra, some dorsal ribs, a
caudal centrum, and appendicular bones. From a taphonomic
viewpoint, the skeletal elements were disarticulated, with the
exception of the metacarpals, which lay in a semi-articulated
position. The bones showed no preferential orientation due to
hydraulic processes, which is consistent with the presence of
mudstone facies that were deposited in the floodplain. The
preservation of small, fragile elements such as teeth and skull
bones confirms that the dispersal and transport of the bones
Figure 2. Preserved skeletal elements of Quetecsaurus rusconii,
was minimal. After burial, the metacarpals underwent frac- UNCUYO-LD-300.
5
Institutional abbreviations. UNCUYO-LD, Universidad gular coracoid with an anterodorsal angle around 90°; (12)
Nacional de Cuyo, Instituto de Ciencias Básicas, Laboratorio hemal arches with open proximal articular facets; and (13)
de Dinosaurios, Mendoza, Argentina. metacarpals without distal articular facets.
Anatomical nomenclature. For the axial skeleton, we use the Quetecsaurus rusconii sp. nov.
terminology proposed by Wilson (1999) and Wilson et al. Figures 2–15
(2011). We also incorporate selected terms proposed by Sal- Derivation of name. Rusconii, in honor of Carlos Rusconi
gado and Powell (2010). Appendicular nomenclature follows (1898-1969), a naturalist who worked extensively in Mendoza
several recent papers (e.g., Salgado et al., 1997; González Riga, Province and was Director of the Museum of Natural Sciences
2005; Curry Rogers, 2009; Otero, 2010). “Juan Cornelio Moyano”.
Diagnosis. As for the genus.
SYSTEMATIC PALEONTOLOGY
Holotype. UNCUYO-LD-300, represented by the following
DINOSAURIA Owen, 1842
associated bones belonging to a single individual: postorbital,
SAURISCHIA Seeley, 1887
two teeth, atlas, one posterior cervical vertebra, one incom-
SAUROPODA Marsh, 1878
plete anterior dorsal vertebra, one anterior caudal centrum,
NEOSAUROPODA Bonaparte, 1986
eight dorsal ribs, a coracoid, an incomplete humerus, distal
MACRONARIA Wilson and Sereno, 1998
fragments of a radius and an ulna, and five metacarpals.
TITANOSAURIFORMES Salgado, Coria, and Calvo, 1997
Horizon and locality. Uppermost levels of the Cerro Lisandro
SOMPHOSPONDYLI Wilson and Sereno, 1998
Formation (late Cenomanian-early Turonian after Hugo and
TITANOSAURIA Bonaparte and Coria, 1993
Leanza [2001], middle Turonian after Legarreta and Gulisano
LITHOSTROTIA Wilson and Upchurch, 2003
[1989]; middle-late Turonian after Garrido [2010]), Neuquén
Genus Quetecsaurus gen. nov. Group, Cañada del Pichanal section, Malargüe Department,
Type species. Quetecsaurus rusconii sp. nov. Mendoza Province, Argentina.
Derivation of name. From Quetec, fire (Milcayac, the lan-
guage used by the people who inhabited the region of Men- Description and comparisons
doza); saurus (Greek), lizard. Postorbital. The only preserved skull bone is a right postor-
Diagnosis. Titanosaurian sauropod characterized by the fol- bital (UNCUYO-LD-300.1; Fig. 3). The bone is not well pre-
lowing combination of autapomorphies: (1) intercentrum of served and has a narrow anterior process and a wide and
atlas with prominent anteroventral border and expanded quadrangular posterior section. Between them, the anterior
posteroventral processes; (2) posterior cervical neural spines and posterior portions form an angle of ~90º. As in most
with incipient lateral laminae; and (3) humerus with strongly neosauropods this bone has a posterior process and a jugal
sigmoid proximal border, rounded proximomedial corner, process that is dorsoventrally flattened. The morphology of
and angular proximolateral corner. Quetecsaurus is also diag- the postorbital is similar, in general terms, to that of Nemeg-
nosed by the following unique combination of characters: (1) tosaurus mongoliensis Nowinski, 1971, from Maastrichtian
opisthocoelous cervical centra; (2) deep spinoprezygapophy- strata of Mongolia, and to that of Tapuiasaurus macedoi Zaher,
seal and spinopostzygapophyseal fossae on posterior cervical Pol, Carvalho, Nascimento, Riccomini, Larson, Juarez-Valieri,
vertebrae; (3) accessory posterior centrodiapophyseal lamina Pires-Domingues, da Silva, and Campos, 2011, from the Early
on posterior cervical vertebrae; (4) posterior cervical centra rel- Cretaceous of Brazil. However, in contrast to these two taxa,
atively elongate (total length/posterior cotyle height greater the posterior process is more rounded (Fig. 3).
than 2.5); (5) cervical vertebra height/centrum length less than Teeth. Two teeth have been preserved, one of them is almost
1.5; (6) prespinal lamina extends to the base of the neural complete (UNCUYO-LD-300.2; Fig 4). The best-preserved
spine on anterior dorsal vertebrae; (7) opisthocoelous ante- tooth has an elliptical section, as in most derived titanosaurs
rior dorsal centra; (8) procoelous anterior caudal centra; (9) in which the teeth have been described as “pencil-chisel like”
neural arch located anteriorly on anterior caudal centra; (10) (e.g., Calvo, 1994; Canudo, 2002; Apesteguía, 2004; Garcia
infraglenoid lip very developed on the coracoid; (11) quadran- and Cerda, 2010). In the apex, the crown is labiolingually
6
compressed, but this morphology is subcircular toward the golia, and from that of Kotasaurus yamanpalliensis Yadagiri,
roof (Fig. 4). It has longitudinal, denticulate carinae that are 1988 (Yadagiri, 2001) from the Early Jurassic of India. In these
visible to the naked eye. The carinae extend from the apex to sauropods, the intercentrum is relatively wide and the an-
the middle portion of the tooth on their mesial and distal teroventral border is not prominent. In Quetecsaurus, the ar-
edges, separating the labial and lingual sides, as in Tapuiasaurus ticulation with the occipital condyle is wider than high. The
(Zaher et al., 2011). The enamel has an ornamentation simi- anterior face of the centrum is concave, and the canal with the
lar to that seen in Petrobrasaurus Filippi, Canudo, Salgado, articulation with the odontoid process is open and has a sub-
Garrido, García, Cerda and Otero, 2011; Tapuiasaurus; and circular contour. The posterior face of the intercentrum is flat
Rapetosaurus Curry Rogers and Foster, 2001. It lacks the and has a quadrangular contour, somewhat similar to that of
“bisel” surface, interpreted as a feature of a replacement tooth. Erketu. Moreover, there are two well-developed posteroventral
Atlas. The atlas is incomplete but shows important characters processes that are absent in Rapetosaurus and Futalognkosaurus
(UNCUYO-LD-300.3; Fig. 5). The intercentrum is robust and very reduced in Erketu. The ventral face of the inter-
and has a wide ventral face with a prominent anteroventral centrum is relatively wide and anteroposteriorly concave, and
border. In contrast, the atlas of Rapetosaurus has a short and it is well demarcated from the lateral faces by ridges connected
poorly developed anteroventral border. In Futalognkosaurus to the posteroventral processes. The anteroventral border is
dukei Calvo, Porfiri, González Riga and Kellner, 2007, the in- ventrally oriented and extends anteriorly 1 cm. The total dis-
tercentrum is thin with a short anteroventral border. The atlas tance between the posterior processes is 4.7 cm. Other meas-
of Quetecsaurus is also different from that of Erketu ellisoni urements are given in Tab. 1.
Ksepka and Norell, 2006, from the Early Cretaceous of Mon- Cervical vertebra. A partially complete posterior cervical ver-
tebra is preserved (UNCUYO-LD-300.4; Fig. 6). The cen-
trum is dorsoventrally compressed and opisthocoelous, as in
other titanosaurs (e.g., Futalognkosaurus, Ligabuesaurus lean-
zai Bonaparte, González Riga and Apesteguía, 2006, Men-
dozasaurus, and Pitekunsaurus macayai Filippi and Garrido,
2008). The ventral face of the centrum is anteroposteriorly
concave. The postzygapophyseal processes are relatively short
and have wide, ventrolaterally directed articular facets. The
prezygapophyseal processes are relatively long and surpass the
Figure 3. Quetecsaurus rusconii gen. et sp. nov., UNCUYO-LD-300.1,
right postorbital. 1, lateral view; 2, medial view; 3, dorsal view; 4, ante- anterior face of the centrum. The postzygapophyseal facets are
rior view. Scale bar = 5 cm. connected by the intrapostzygapophyseal lamina. As seen in
posterior view, there is a deep spinopostzygapophyseal fossa
that is delimited by the spinopostzygapophyseal laminae and
the intrapostzygapophyseal lamina. The prezygapophyses ex-
hibit robust, semicircular articular facets. In anterior view, a
relatively wide intraprezygapophyseal lamina connects the
prezygapophyses, forming a roof over the neural canal, as also
occurs in Ligabuesaurus and titanosaurs. Together with the
spinoprezygapophyseal laminae, the intraprezygapophyseal
lamina circumscribes the deep spinoprezygapophyseal fossa.
The spinoprezygapophyseal lamina is strongly curved an-
teriorly and reaches the tip of the neural spine. This lamina is
similar to that of Futalognkosaurus. In contrast, in Mendoza-
saurus and Puertasaurus the spinoprezygapophyseal laminae
tooth. 1–1´, distal view; 2, mesial view. 3, section. Abbreviations: ca:
crown apex; lc: longitudinal carinae; rt: root. Scale bar= 1 cm. are well separated and only reach the middle part of the neu-
7
ral spine. In other titanosaurs such as Saltasaurinae (Powell, prezygapophyseal fossa. The neural spine is posteriorly ori-
1986) and Rinconsaurini (sensu Calvo et al., 2007c) this char- ented and placed on the posterior portion of the neural arch,
acter is present but is not associated with a very deep spino- as in Futalognkosaurus. Its morphology differs from that of
Figure 5. Quetecsaurus rusconii gen. et sp. nov., UNCUYO-LD-300.3, atlas. 1–1´, anterior view; 2–2´, left lateral view; 3–3´, dorsal view; 4–4´, pos-
terior view; 5–5´, right lateral view; 6–6´, ventral view. Abbreviations: na: neurapophysis; awb: wide anteroventral border; pvp: posteroventral
process. Scale bar= 1 cm.
8
Figure 6. Quetecsaurus rusconii gen. et sp. nov., UNCUYO-LD-300.4, middle-posterior cervical vertebra. 1–1´, right lateral view; 2–2´, posterior
view. Abbreviations: nc: neural canal; dp: diapophysis; sdf: spinodiapophyseal fossa; spof: spinopostzygapophyseal fosa; pcdl: posterior cen-
trodiapophyseal lamina; tpol: intrapostzygapophyseal lamina; spol: spinopostzygapophyseal lamina; sprl: spinoprezygapophyseal lamina; ll:
lateral laminae; podl: postzygodiapophyseal lamina ; pp: parapophysis; poz: postzygapophysis; prz: prezygapophysis. Scale bar= 10 cm.
9
Isisaurus colberti Jain and Bandyopadhyay, 1997 (Wilson and odiapophyseal laminae links a reduced neural spine with al-
Upchurch, 2003) from the Maastrichtian of India, Puer- most horizontally oriented diapophyses. The horizontal ori-
tasaurus, and Mendozasaurus, where the neural spines are lat- entation of the diapophysis and the morphology of the neural
erally expanded, reaching or surpassing the width of the spine indicate that this vertebra is an anterior dorsal, proba-
vertebral centrum. The neural spine expands distally through bly the third.
the development of lateral laminae (i.e., ‘lateral expansions’), Caudal vertebra. An anterior caudal centrum is preserved
a feature also present in Mendozasaurus (González Riga, 2005) (UNCUYO-LD-300.6; Fig. 8). It is short as in other ti-
and Futalognkosaurus (Calvo et al., 2007b), although this con- tanosaurs (length: width ratio of 0.86). Similar proportions
dition appears to be less developed than in these taxa. Based on are present in, for example, anterior caudal vertebrae of Men-
comparisons with the articulated cervical vertebrae of Futa- dozasaurus, Uberabatitan ribeiroi Salgado and Souza Carvalho,
lognkosaurus, the cervical vertebra of Quetecsaurus is prelimi- 2008, Adamantisaurus mezzalirai Santucci and Bertini, 2006,
nary assigned to an 11th place on the series. In the posterior Aeolosaurus maximus Santucci and Arruda-Campos, 2011,
cervical of Quetecsaurus, the neural spine is ~13 cm wide, Chubutisaurus insignis del Corro, 1975, and Lirainosaurus
which is much less than the width of the centrum (Tab. 1). In astibiae Sanz, Powell, Le Loeuff, Martínez and Pereda-Suber-
Mendozasaurus, by contrast, the lateral laminae are so promi- biola, 1999. The centrum is procoelous and its anterior cotyle
nent that the neural spine is wider than the centrum; this char- has a slight lateral border. The cotyle is anteriorly titled, such
acter is less pronounced in Futalognkosaurus, where the spine that its dorsal border is placed anteriorly with respect to the
is equal in width or narrower than the centrum. Moreover, in ventral border. The posterior condyle is prominent and dor-
Ligabuesaurus, the neural spines are expanded, but these ex- sally displaced, as in Mendozasaurus. The transverse processes
pansions are constructed from lateral spinoprezygapophyseal are not preserved but their bases indicate that they were pos-
laminae, not from lateral laminae that originate in the spin- terolaterally directed. The centrum has strongly concave lateral
odiapophyseal (=supradiapophyseal) fossa (González Riga, and ventral faces. The neural arch is placed on the anterior
2005). In Quetecsaurus, the spinodiapophyseal fossa is well de- part of the centrum, as in caudals of other titanosauriforms.
fined, but it is shallower than in Mendozasuarus. It is bounded
by spinopostzygapophyseal and spinoprezygapophyseal lami-
nae. Pleurocoels are absent, but the centrum has a large lon-
gitudinal cavity ventral to the posterior centrodiapophyseal
lamina. Dorsal to this lamina there are two small cavities that
are delimited by an accessory posterior centrodiapophyseal
lamina and the postzygodiapophyseal lamina. The para-
pophyses are robust and anteroventrally directed, and the di-
apophyses are short and posteroventrally oriented.
Dorsal vertebra. A fragmentary anterior dorsal vertebra is
preserved (UNCUYO-LD-300.5; Fig. 7). It retains a portion
of the neural arch and spine, and part of the posterior cotyle
and anterior condyle, but lacks most of the centrum, prezy-
gapophyses, and diapophyses. It has an opisthocoelous cen-
trum with a circular anterior and posterior contour. The
neural spine is single, short, and vertically directed, as in
Pitekunsaurus, Mendozasaurus, and Drusilasaura deseadensis
Navarrete, Casal and Martínez, 2011. It exhibits a prespinal Figure 7. Quetecsaurus rusconii gen. et sp. nov., UNCUYO-LD-300.5, an-
terior dorsal vertebrae. 1–1´, anterior view; 2–2´, posterior view. Abbre-
lamina that reaches the spinodiapophyseal laminae. The right viations: nc: neural canal; vc: vertebral centrum; dp: diapophysis; ns:
diapophysis is partially preserved, and its morphology demon- neural spine; spdl: spinodiapophyseal lamina; prsl: prespinal lamina;
spol: spinopostzygapophyseal lamina; poz: postzygapophysis. Scale
strates that the vertebra is wider than high (Tab. 1). The spin- bar= 10 cm.
10
Dorsal ribs. Eight dorsal ribs are preserved, some of them served ribs, there are pneumatic cavities between the tubercu-
complete (UNCUYO-LD-300.7-14; Fig. 9). In the better pre- lum and capitulum, as has been described for Titanosauri-
Figure 8. Quetecsaurus rusconii gen. et sp. nov., UNCUYO-LD-300.6, anterior caudal vertebra. 1–1´, right lateral view; 2–2´, anterior view; 3–3´,
dorsal view; 4–4´, left lateral view; 5–5´, posterior view; 6–6´, ventral view. Scale bar= 5 cm.
11
TABLE 1 - Measurements of the vertebrae of Quetecsaurus rusconii gen. et sp. nov. (LD-Pv 1)
Relationship Ratio total

Ratio length /
Length overall height / height / height
Skeletal element Total height Center height Center Width height of the
of center length of the of center
center
center
Atlas 3 2,3 5 — — 0,937
Posterior cervical vertebra 49 40 14 23 1,225 1,642 2,857
Anterior dorsal vertebra 39 — 20 24* — 1,95 —
Anterior caudal vertebra — 16 14 13 — — 1,143
Measurements in cm

dorsal rib. Abbreviations: ca: capitulum; tu: tuberculum. Scale bar=
10 cm.
formes (Wilson and Sereno, 1998). The tuberculum is longer

and more robust than the capitulum. All ribs have elliptical
sections.
Coracoid. A nearly complete right coracoid is preserved (UN-
CUYO-LD-300.15; Figs. 10–11). The lateral surface is convex
and the medial surface is concave. On the lateral surface there
is a slight ridge that extends posteroventrally, ventral to the
coracoid foramen. The bone is very robust in its posteroven-
tral portion; that is, in the area of contact with the scapula,
the glenoid fossa, and the infraglenoid lip. In contrast, the an- Figure 10. Quetecsaurus rusconii gen. et sp. nov., UNCUYO-LD-300.15,
terodorsal portion is thinner (Tab. 2). The coracoid shows a right coracoid. 1–1´, medial view; 2–2´, lateral view. Abbreviations:
igl: infraglenoid lip; gf: glenoid fossa; scs: scapulocoracoid suture; cf:
very well developed infraglenoid lip that forms an angle of coracoid foramen. Scale bar= 10 cm.
approximately 90° with the anterior border. The anterior
border is straight and intersects the dorsal border with an angle closed and placed posteromedially, very close to the posterior
of ~90°. Because of this, the coracoid exhibits a quadrangu- border.
lar contour, as in Neuquensaurus (Otero, 2010), Saltasaurus Humerus. The proximal end of the left humerus has been re-
(Powell, 1992, 2003), and Rinconsaurus caudamirus Calvo covered (UNCUYO-LD-300.16; Fig. 12). The humeral head
and González Riga, 2003. In contrast, other titanosaurs, such is robust and the dorsal border of the bone has a strongly sig-
as Opisthocoelicaudia skarzynskii Borsuk-Bialyncka, 1977, moidal outline. This morphology is different from that of
Rapetosaurus and Isisaurus, have a more rounded or subrec- other titanosaurs, and is herein considered an autapomorphy
tangular outline. The contact with the scapula is almost of Quetecsaurus (Fig. 12). The titanosaurs Saltasaurus and Par-
straight and it is relatively extended in relation with the an- alititan Smith, Lamanna, Lacovara, Dodson, Smith, Poole,
teroposterior length of the bone. The coracoid foramen is Giegengack, and Attia 2001, from the Late Cretaceous of
12
Egypt have sigmoidal proximal humeri, but this condition is the sauropod humerus based on the shape of its proximolateral
relatively less pronounced than that of Quetecsaurus. More- corner, differentiating between rounded and square mor-
over, the humerus of Quetecsaurus has a broad and rounded phologies (Wilson, 2002; character 159). Later, in a character
proximomedial border and an angular proximolateral corner, matrix, González Riga (2003; character 27) described the
the proximal and lateral margins of which form an obtuse morphology of the proximal ends of sauropod humeri as
angle (Fig. 12). Wilson (2002) described the proximal end of strongly rounded, straight or slightly rounded, or sigmoidal
(Fig. 16). In fact, in basal titanosauriforms such as Ligabue-
saurus, the proximal border is slightly rounded. Conversely, in
most derived titanosaurs, the proximal end is straight (e.g.,
Rapetosaurus, Mendozasaurus). In Quetecsaurus, a broad and
deep cavity extends in the anterior margin, below the humeral
head; this is different from the conditions in Neuquensaurus
and Rapetosaurus. The deltopectoral crest is relatively reduced
and positioned medially. The robustness of the humeral head
is correlated with the broad glenoid fossa of the coracoid.
Radius and ulna. Only the distal ends of a radius and an ulna
have been preserved (UNCUYO-LD-300.17-18; Fig. 13). The
radius has an anteroposteriorly expanded distal end and dis-
plays a slight depression corresponding to the ulnar articular
surface. The distal end of the ulna has a triangular outline,
with a typical radial articular surface. The distal articular sur-
face has noticeable rugosities and tuberosities which are inter-
preted as sites of ligament insertion.
Metacarpals. The five left metacarpals have been preserved
right coracoid, glenoid view. Abbreviations: igl: infraglenoid lip; gf:
glenoid fossa; scs: scapulocoracoid suture; cf: coracoid foramen. (UNCUYO-LD-300.19-23; Figs. 14–15; Tab. 3). Metacarpals
Scale bar= 10 cm.
Figure 12. Quetecsaurus rusconii gen. et sp. nov., UNCUYO-LD-300.16, left humerus in anterior view. Abbreviations: hh: humeral head; dpc: del-
topectoral crest; erpb: expanded and rounded proximomedial corner; apc: angular proximolateral corner. Scale bar= 10 cm.
13
II, III, and IV were articulated whereas metacarpals I and V cal positions. The complete metacarpal structure is known in
were displaced a few centimeters from their original anatomi- some titanosaurs such as Rapetosaurus, Opisthocoelicaudia, and
Epachthosaurus sciuttoi Powell, 1990.
The proximal and distal ends are well preserved, whereas
the shafts are broken at their mid sections and exhibit numer-
ous diagenetic fractures. When articulated, the metacarpals are
oriented almost vertically and form a semilunate arch with a
flat proximal surface (Fig. 15). Due to breakage of the shafts,
it is not possible to precisely measure the length of most
metacarpals, but the length of metacarpal III can be estimated
at about 36 cm.
Metacarpal I has a dorsomedially elongate proximal end.
The diaphysis is flattened on its medial and lateral faces; a
different condition is observed in Epachthosaurus, where
metacarpal I is the most robust of the series. The lateral surface
of the proximal shaft has a broad concavity for articulation
with metacarpal II. The distal end has a subtriangular outline,
being rounded at its medial border. Metacarpal II has the most
robust proximal end of the series, and has a subcircular outline
as in Petrobrasaurus. The proximal end is more robust than the
distal end. The shaft has a triangular cross section and its me-
dial side has a slightly concave surface for articulation with
metacarpal III. The distal end has a subquadrangular outline
and is slightly concave. Metacarpal III has the most slender
Figure 13. Quetecsaurus rusconii gen. et sp. nov., UNCUYO-LD-300.17, shaft of the series; it has a circular cross section. The proximal
UNCUYO-LD-300.18, left radius and ulna. 1–1´, distal shaft of radius in
posterior view; 2–2´, distal shaft of ulna in anterior view; 3, distal shaft
end has a triangular outline; the distal end is dorsoventrally
of radius in ventral view; 4, distal shaft of ulna in ventral view. Abbre- compressed and shows a subquadrangular outline. The surface
viations: uas: ulnar articular surface; ras: radial articular surface. Scale
bar= 5 cm. of the distal end is not perpendicular to the shaft but instead
TABLE 2 - Measures of various skeletal elements of Quetecsaurus rusconii gen. et sp. nov. (LD-Pv 1)
Width in previous Width at posterior

Length Height
portion portion
Postorbital 11,8 4,3 1,5 3,3
Width of the
middle section
Tooth 4 0,5
Dorsal rib 132 6

Postero-dorsal Anteroventral Length of the Infraglenoid
Width
border length border length glenoid fossa lip length
Coracoid 32 25 34 16 13
Width of the diaphysis at
Proximal border width
proximal portion
Humerus 37 15,5
Measurements in cm
14
is oriented toward the ventral side. Metacarpal IV is proxi-

mally incomplete. The distal end is subquadrangular and
slightly concave lateromedially. On the ventral side, the distal
end has a prominent process. The diaphysis is expanded dor-
somedially and has a concave articular surface for metacarpal
III. Metacarpal V has a dorsoventrally elongate proximal end.
The distal end is subquadrangular and slightly concave. This
element has been distorted by taphonomic processes.
Although their preservation is poor, the distal ends of the
metacarpals show important characters. The articular surfaces
for the phalanges are not well defined, as in most titanosaurs.
However, in metacarpal I and IV there is a prominent, ven-
trally-directed process. Moreover, metacarpals II and V have a
lateromedially concave surface. The described structure is sim-
ilar to those of other titanosaurs, following the detailed study
of Apesteguia (2005).
PHYLOGENETIC ANALYSIS AND DISCUSSION

Quetecsaurus rusconii displays a unique combination of
characters within Titanosauria, which justifies its diagnosis as
a new taxon. Three characters are here considered autapo-
morphies: (1) intercentrum of atlas with a prominent an-
teroventral border and expanded posteroventral processes; (2)
posterior cervical neural spines with an incipient lateral lam-
ina; and (3) humerus with a strongly sigmoidal proximal
border, a rounded proximomedial border, and an angular
proximolateral corner.
A morphological comparison of the proximal ends of ti-
tanosaur humeri revealed interesting information (Fig. 16).
For example, Rapetosaurus and Mendozasaurus have a straight
proximal end, with a proximolateral corner that forms an angle
of ~90°. In contrast, Quetecsaurus is the only titanosaur that
has a strongly sigmoidal proximal end, with an expanded and
rounded proximomedial border and an angular proximolat-
eral corner that describes an obtuse angle (Fig. 16). Only
Saltasaurus and Paralititan show a partly similar morphology,
but with a less sigmoidal outline; moreover, the humeri of

UNCUYO-LD-300.20, UNCUYO-LD-300.21 UNCUYO-LD-300.22, UNCUYO-
LD-300.23, left metacarpals.1–4, metacarpal I; 1, medial view; 2, lateral
view; 3, proximal view; 4, distal view. 5–8, metacarpal II; 5, dorsal view;
6, ventral view; 7, proximal view; 8, distal view. 9–12, metacarpal III; 9,
dorsal view; 10, ventral view; 11, proximal view; 12, distal view. 13–16,
metacarpal IV; 13, medial view; 14, lateral view; 15, proximal view; 16,
distal view. 17–20, metacarpal V; 17, medial view; 18, lateral view; 19,
proximal view; 20, distal view. Scale bar= 5 cm.
15
these two taxa do not have a rounded and expanded proxi- through a cladistic analysis based on 82 characters scored
molateral border (Fig. 16). for 22 taxa (see Tab. 4 and Appendix). Camarasaurus gran-
The phylogenetic relationships of Quetecsaurus were tested dis (Cope, 1877) was considered as the outgroup of 21 ti-
Figure 15. Quetecsaurus rusconii gen. et sp. nov., UNCUYO-LD-300.19, UNCUYO-LD-300.20, UNCUYO-LD-300.21, UNCUYO-LD-300.22, UNCUYO-LD-
300.23, left metacarpus. 1–1´, proximal view; 2–2´, distal view. Abbreviations: mcI: metacarpal I; mcII: metacarpal II; mcIII: metacarpal III; mcIV:
metacarpal IV; mcV: metacarpal V. Scale bar= 5 cm.
TABLE 3 - Measures of the metacarpals of Quetecsaurus rusconii gen. et sp. nov. (LD-Pv 1)
Proximal epiphysis Distal epiphysis

Width of the middle
Estimated total length
section
Length Width Length Width
Metacarpal I 28.5 7.6 12.8 4.4 11.3 6.4
Metacarpal II 31.2 7 9.8 9.3 10.4 8.6
Metacarpal III 29.2 4.5 9.2 8.7 10.2 7.3
Metacarpal IV 30.3 7.5 8.9 5.6 8.1 10.1
Metacarpal V 25.1 6.5 10.4 5.3 8.7 7.3
Measurements in cm
16
tanosauriform taxa, which include two basal forms (Bra- tree search. The strict consensus cladogram (Fig. 18) shows
chiosaurus brancai Janensch, 1950, and Chubutisaurus insignis some differences with recent studies (e.g., González Riga et al.,
Del Corro, 1975) and a diverse sample of 19 titanosaurs (see 2009; Gallina and Apesteguía, 2011), particularly in the poly-
Tab. 4). We used a list of characters derived from previous tomy formed by Drusilasaura Navarrete, Casal and Martinez,
analyses (e.g. Wilson, 2002; Gonzalez Riga, 2003; Curry 2011 and the clade (Bonitasaura + (Quetecsaurus + (Mendoza-
Rogers, 2005; Calvo et al., 2007a; González Riga et al., 2009; saurus + Futalognkosaurus))) (Fig. 18). Most titanosaurs are
Gallina and Apesteguía, 2011; Zaher et al., 2011) after a revi- represented by very incomplete skeletons: some have caudal
sion of each of them, we redefined two characters (58 and 82), vertebrae, others dorsal vertebrae, or cranial remains without
and added three new characters referred to the cervical verte- postcranial bones. This results in the weak support of some
brae (24, 26 and 27) (see Tab. 4 and Appendix). terminal taxa and nodes (see González Riga, 2010). In this
The data matrix was analyzed using the program NONA, case Drusilasaura is represented by an incomplete skeleton:
version 2.0 (Goloboff, 1993), with multistate characters con- dorsal and caudal vertebrae, one sacral vertebra, a scapula, and
sidered unordered, and the character polarity was determined rib fragments (Navarrete et al., 2011).
by comparison with the outgroup. Two most parsimonious Titanosauria, defined as ‘Andesaurus delgadoi (Calvo and
trees (157 steps, C.I.: 0.64; R.I.: 0.71) were obtained after the Bonaparte, 1991), Saltasaurus loricatus (Bonaparte and Pow-
TABLE 4 - Matrix of characters used for the phylogenetic analysis
Taxon 1-10 11-20 21-30 31-40 41-50 51-60 61-70 71-80 81-82
Camarasaurus 0011000001 1001000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 00
Brachiosaurus 0001000011 0000000010 1010100001 0010100010 0000000000 0000001000 0000000000 000101110? 00
Chubutisaurus ?????????? ?????????? ?????????? ????????2? 0001?0??00 100???1??0 ???01????? 0?????111? ??
Andesaurus ?????????? ?????????? ?????????? ????1??011 1011?0?000 2000001100 00?01????1 110???01?? ??
Malawisaurus 0??????11? ?0?????000 201??00001 00??11?011 1111??1000 32000?1110 0???1111?1 ?1??????1? ?1
Ligabuesaurus ?????????? ?????????? 101??00001 010?111011 1111?????? ?????????? ??10?????? 0?????111? ??
Mendozasaurus ?????????? ?????????? ?????1100? 122?120?2? ?111?11000 3210011110 00101111?? 1?????0111 ?1
Futalognkosaurus ?????????? ?????????? ???0211001 12221?0?2? ?111111?0? 3????11??0 ?????????1 ?12111???? ??
Quetecsaurus ???000???? ?????????? 2211?10001 010?1?0?2? ?1?1????00 3?????1??0 ??1?2?111? 1????????1 ??
Epachthosaurus ?????????? ?????????? ?????????? ????101121 101111?000 3320001110 0010111??1 1??1110111 10
Rapetosaurus 111?001110 1111111110 2210200001 00?1121121 11111?0000 33200110?0 001011100? 11011101?1 ?1
Drusilasaura ?????????? ?????????? ?????????? ????100?2? ?111?11?00 33?0011?10 001??????? ?????????? ?0
Rinconsaurus ?????????? ?????????? 211??00001 011?101121 11111???00 3320011010 1110111011 11111101?? ??
Muyelensaurus ?1????1??1 1???1?111? 211??0000? 011?121121 11111?0?00 3320011010 11101110?1 111111011? ??
Bonitasaura ?11???111? 1?1??????1 221??00?01 011?120121 1111?11000 3320011010 01?01?1??1 112???011? ??
Gondwanatitan ?????????? ?????????? ?????????? ???1?11211 111111???1 3320102020 01?0?????? ?0111???1? ??
Aeolosaurus ?????????? ?????????? 221??????? ????1???2? ?????30?01 3320112020 01?01?10?1 101????11? ?1
Opisthocoelicaudia ?????????? ?????????? ?????????? ???1001121 0111121100 0000021000 0011111011 1111110111 10
Alamosaurus ?????????? ?????????? 22???00001 001?1??120 0111131100 3320021000 0010111111 112111011? ??
Neuquensaurus ?????????? ?????????? ?????00101 012?101121 1111131?10 3321021000 001011101? 112111?11? ?2
Saltasaurus ?11?????11 1???01011? 221??00111 012?111121 1111110?10 3321021001 0011211011 112111011? ?2
Rocasaurus ?????????? ?????????? ?????0???? 01??101121 1111????10 33210?1001 00???????1 ?12111?1?? ??
17
18
Figure 17. Posterior cervical vertebrae of Argentinean titanosauriforms. 1, Bonitasaura in anterior view; 2, Bonitasaura in lateral view; 3, Futa-
lognkosaurus in anterior view; 4, Futalognkosaurus in lateral view; 5, Mendozasaurus in anterior view; 6, Mendozasaurus in anterior view; 7, Quetec-
saurus in posterior view; 8, Quetecsaurus in lateral view (reversed); 9, Puertasaurus in anterior view; 10, Puertasaurus in lateral view; 11, Ligabuesaurus
in anterior view; 12, Ligabuesaurus in lateral view. Not to scale. Abbreviations: msprl: medial spinoprezygapophyseal lamina; sprl: spinoprezy-
gapophyseal lamina; spol: spinopostzygapophyseal lamina; spdl: spinodiapophyseal lamina; le: lateral expansion. Modified from Gallina, 2011.
ell, 1980), their most recent common ancestor and all de- tree. Obviously, a more detailed analysis including all new taxa
scendants’ (Salgado et al., 1997, modified by Wilson and recently described from Brazil and other countries is necessary
Upchurch, 2003) is supported by two unambiguous synapo- for clarifying the phylogenetic relationships of these clades, a
morphies: absence of well-developed distal phalangeal articu- goal beyond the scope of this contribution.
lar facets on metacarpals (character 71, state1), and humerus/ Our analysis recovers Quetecsaurus rusconii as the sister
femoral ratio less than 0.9 (character 77, state 1). Within Ti- taxon of Lognkosauria (Mendozasaurus + Futalognkosaurus).
tanosauria, the node-based Lithostrotia (defined as ‘the most Lognkosauria was defined by Calvo et al. (2007a) as ‘the most
recent common ancestor of Malawisaurus and Saltasaurus, and recent common ancestor of Mendozasaurus neguyelap (Gon-
all the descendants of that ancestor’ sensu Upchurch et al., zález Riga, 2003) and Futalognkosaurus dukei (Calvo et al.,
2004), is supported by three unambiguous synapomorphies: 2007a), and all of its descendants’. In the original paper, five
articular face shape on anterior caudal centra strongly pro- synapomorphies supported this clade: (1) presence of laterally
coelous with prominent condyles (character 51, state 3), ar- expanded posterior cervical neural spines that are wider than
ticular face shape on middle caudal centra strongly procoelous their respective centra; (2) posterior cervical vertebrae 1.5
with prominent condyles (character 52, state 2), and length times taller than long; (3) deep and extended spinodiapophy-
proportions of prezygapophyses between 40 to 50% with re- seal fossa in posterior cervical vertebrae; (4) ratio of antero-
spect to the centrum length in middle caudal vertebrae (char- posterior length/ height of posterior face in posterior cervical
acter 59, state 1). centra less than 1.5; and (5) anteriormost caudal vertebrae
Within Lithostrotia we recognize five different clades: with transversely expanded neural spines.
Longkosauria, Rinconsauria, Opisthocoelicaudinae, Aeolo- In our analysis, Lognkosauria is supported by three un-
saurini and Saltasaurinae (Fig. 18), all supported by unambi- ambiguous synapomorphies: (1) laterodorsally expanded cer-
guous synapomorphies. Boostrap and Jackknife values up to vical neural spines originated by lateral laminae that reach or
50% are indicated in selected nodes of the strict consensus surpass the wide of the centra (character 27, state1); (2) pos-
Figure 16. Humeri of sauropods (only the anterior view of the proximal end is represented). 1, Quetecsaurus; 2, Saltasaurus; 3, Paralititan; 4, Chubuti-
saurus; 5, Diamantinasaurus; 6, Opisthocoelicaudia; 7, Narambuenatitan; 8, Epachthosaurus; 9, Neuquensaurus; 10, Mendozasaurus; 11, Magyarosaurus;
12, Rapetosaurus; 13, Atacamatitan; 14, Petrobrasaurus; 15, Bonitasaura; 16, Ligabuesaurus; 17, Panamericansaurus; 18, Tornieria; 19, Kotasaurus; 20,
Borealosaurus. (Images reversed for Opisthocoelicaudia, Mendozasaurus, Paralititan, Epachthosaurus, Magyarosaurus, Petrobrasaurus, Kotasaurus,
Tornieria, and Borealosaurus.) Not to scale.
19
Figure 18. Cladogram showing the phylogenetic relationships of Quetecsaurus rusconii based on strict consensus of two most parsimonious trees
(157 steps, C.I.: 0.64, R.I.: 0.71). The robustness of nodes is indicated in brackets (bootstrap and jackknife values, respectively).
terior cervical vertebrae 1.5 times taller than long (character the available information and the results of this analysis, Drusi-
31, state 1); and (3) deep and extended spinodiapophyseal lasaura should not be included within Lognkosauria.
fossa in posterior cervical vertebrae (character 32, state 2). A revision of the characters referred to the cervical neural
The synapomorphy defined by Calvo et al., (2007a) as ‘an- spines is important. González Riga (2010, p. 129) recognized
teriormost caudal vertebrae with transversely expanded neu- two distinct structural patterns among the huge cervical
ral spines’ (character 56, state 1, in this paper) is not recognized neural spines of titanosars: the Mendozasaurus-type and the
here as diagnostic of Longkosauria diagnosis since it has a Ligabuesaurus-type. In the first one (e.g., Mendozasaurus, Fu-
broader distribution (e.g., Rapetosaurus, Muyelensaurus, Rin- talognkosaurus) a very wide neural spine is formed by distinct
consaurus, Bonitasaura, Aeolosaurus, Drusilasaura). In particu- structures named ‘lateral expansions’ (lateral laminae in this
lar, Drusilasaura (Navarrete et al., 2011: fig. 9.A) shares with paper), which are not related to the spinopostzygapophyseal
Futalognkosaurus (Calvo et al., 2007b: figs. 16–17) a robust or the spinoprezygapophyseal laminae (González Riga, 2005,
neural spine in the most anterior caudals with well-developed figs. 2–3; Calvo et al., 2007b, fig. 11). Thus, synapomorpy 1
lateral expansion. This condition is absent in Mendozasaurus, of Lognkosauria has been here redefined as ‘laterodorsally ex-
which has a transversely expanded neural spine (González Riga, panded cervical neural spines originated by lateral laminae that
2003: figs. 4.D–E) but that is nonetheless relatively reduced in reach or surpass the wide of the centra’. It is relevant to note
comparison with these two taxa. Unfortunately, cervical ver- that these same lateral laminae are also present in Quetec-
tebrae of Drusilasaura have not been preserved, which are char- saurus, but are less strongly developed. In the second type, the
acteristic of lognkosaurs and would allow testing its affinities huge neural spines of Ligabuesaurus are formed by splayed lat-
with this clade, as noted by Navarrete et al. (2011). Based on eral spinoprezygapophyseal laminae (Bonaparte et al., 2006;
20
González Riga, 2010). Bonitasaura Apesteguía, 2004, is an- nosaurs, Indiana University Press, Indianopolis, p. 321–345.
Apesteguía, S., Agnolín F.L., and K. Claeson. 2007. Review of Cretaceous
other titanosaur that also has a huge cervical neural spine, dipnoans from Argentina (Sarcopterygii, Dipnoi) with description of
which was described by Gallina (2011) as a ‘rhomboid-shaped new species. Revista del Museo Argentino de Ciencias Naturales, Nueva
Serie 9: 27–40.
neural spine forming a simple expansion of the distal spine Bonaparte, J.F. 1986. History of the terrestrial Cretaceous vertebrates of
without a clear contribution of anterior or posterior laminae’. Gondwana. Actas 4th Congreso Argentino de Paleontología y Bioestratigrafía
(Mendoza), Resúmenes 2: 63–95.
Therefore, the broad neural spine of Bonitasaura lacks the lat- Bonaparte, J.F. and Powell, J.E. 1980. A continental assemblage of tetrapods
from the Upper Cretaceous beds of El Brete, northwestern Argentina
eral laminae that form the lateral border of the neural spines (Sauropoda, Coelurosauria, Carnosauria, Aves). Mémoires de la Société
in lognkosaurs. Geólogique de France 139: 19–28.
Bonaparte, J.F. and Coria, R.A. 1993. Un nuevo y gigantesco saurópodo ti-
Other lognkosaurian characters are not present in Quetec- tanosaurio de la Formación Río Limay (Albiano-Cenomaniano) de la
saurus. For example, with regard to synapomorphy 2, the ratio provincia del Neuquén, Argentina. Ameghiniana 30: 271–282.
Bonaparte, J.F., González Riga, B.J. and Apesteguía, S. 2006. Ligabuesaurus
of the height of the cervical vertebra to the length of its cen- leanzai nov. gen. et sp., a new titanosaur from the Aptian of Patagonia,
trum is less than 1.5 (see Tab. 1). The synapomorphy 3, the Argentina. Cretaceous Research 27: 364–376.
Borsuk-Bialynicka, M. 1977. A new camarasaurid sauropod Opisthocoelicau-
deep spinodiapophyseal fossa that is characteristic of Men- dia skarzynskii, gen. n. sp. n. from the Upper Cretaceous of Mongolia.
Palaeontologica Polonica 37: 45–64.
dozasaurus (González Riga, 2005: fig. 2) and Futalognkosaurus
Calvo, J.O. 1994. Jaw mechanics in sauropod dinosaurs. GAIA 10: 183–193.
(Calvo et al., 2007b: fig. 11), does not have the same devel- Calvo, J.O. and Bonaparte, J.F. 1991. Andesaurus delgadoi n. g. n. sp. (Sauris-
chia, Sauropoda) dinosaurio Titanosauridae de la Formación Río Limay
opment in Quetecsaurus, where this fossa is smaller and shal- (Albiano-Cenomaniano), Neuquén, Argentina. Ameghiniana 28: 303–310.
lower. In this context, Quetecsaurus does not appear to be a Calvo, J.O. and Salgado, L. 1995. Rebbachisaurus tessonei sp. nov., a new
Sauropoda from the Albain-Cenomanian of Argentina; New evidence on
member of Lognkosauria, but is instead recovered as a close the origin of the Diplodocidae. GAIA 11: 13–33.
phylogenetic relative of the lognkosaurs Mendozasaurus and Calvo, J.O. and González Riga, B.J. 2003. Rinconsaurus caudamirus gen. et sp.
nov., a new titanosaurid (Dinosauria, Sauropoda) from the Late Creta-
Futalognkosaurus. ceous of Patagonia, Argentina. Revista Geológica de Chile 30: 333–353.
Calvo, J.O., Porfiri, J.D., González Riga, B.J. and Kellner, A.W.A. 2007a. A
new Cretaceous terrestrial ecosystem from Gondwana with the descrip-
ACKNOWLEDGMENTS tion of a new sauropod dinosaur. Anais da Academia Brasileira de Cien-
We thank M. Tovar, Director of the Instituto de Ciencias Basicas, and A. So- cias 79: 529–541.
moza, Rector of the Universidad Nacional de Cuyo, for their permanent sup- Calvo, J.O., Porfiri, J.D., González Riga, B.J. and Kellner, A.W.A. 2007b.
port in our researchs in the Laboratorio de Dinosaurios. We are grateful to the Anatomy of Futalognkosaurus dukei Calvo, Porfiri, Gonzalez Riga and
Dirección de Patrimonio del Gobierno de Mendoza for its management in Kellner, 2007 (Dinosauria, Titanosauridae) from the Neuquén Group
the preservation of the dinosaur fossils (Directora Prof. R. Agüero, Vice-Di- (Late Cretaceous), Patagonia, Argentina. Arquivos do Museu Nacional 65:
rector Arq. E. Priori, Lic. E. Albarrán). We thanks the Editor, D. Pol, for its 511–526.
accurate and valuable revision, and two anonymous reviewers that greatly im- Calvo, J.O., González Riga, B.J. and Porfiri, J.D. 2007c. A new titanosaur
proved the early version of the paper. We also wish to thank to M. Lamanna sauropod from the Late Cretaceous of Neuquén, Patagonia, Argentina.
who improved the English of the manuscript and provided important scien- Arquivos do Museu Nacional 65: 485–504.
tific comments. We are grateful to participants in paleontological team that Calvo, J.O., Porfiri, J.D., Pol, D., González Riga, B.J., de la Fuente, M. and
worked in the field (J. Porfiri, G. Retamal, D. Eseisa, D. Rosales, C. Fuentes, Rougier, G.W. 2011. Vertebrados continentales Mesozoicos. 18° Con-
G. Garat, M. Milani, and J. Giacoboni) and F. Ortiz for the preparation of the greso Geológico Argentino (Neuquén), Relatorio: 539–556.
fossils in laboratory. We thank the company Vale S.A. of the “Proyecto Pota- Canudo, J.I. 2002. Una mirada de dentista: los dientes de los dinosaurios sau-
sio Río Colorado” and their environmental consultant, Knight Piésold Ar- rópodos. Asociación Paleontológica Aragonesa 32: 12–24.
gentina, for their willingness to comply with the procedures established by Canudo, J.I., Royo-Torres, R. and Cuenca-Bescós, G. 2008. A new sauropod:
law for the collection and preservation of the fossils discovered during con- Tastavinsaurus sanzi gen. et. sp nov. from the Early Cretaceous (Aptian)
struction of the mine (Ch. Monárdez, J. Leis, C. Delgado, A. Demonte, and of Spain. Vertebrate Paleontology 28: 712–731.
their team). The collaboration of the following students in the laboratory work Casanovas-Cladellas, M.L., Santafé-Llopis, J.V. and Santisteban-Bove, C. 1993.
was also important: G. Sanchez Tiviroli, P. Gutiérrez, R. Sánchez Domina, L. First dinosaur teeth from the Lower Cretaceous of Benicatazara (Aras de
Pinto, L. Resa, J. Menéndez, F. Jofré, L. Martínez, V. Penas, M. Pascual, R. Alpuente, Valencia). Revue de Palèobiologie, Vol. spec. (7), 37–44.
López, D. Solíz, M. Fredes, and F. Santos. This research was supported by Cope, E.D. 1877. On a gigantic saurian from the Dakota Epoch of Colorado.
R. Villalba and S. Londero from the Instituto Argentino de Nivología, Gla- Paleontological Bulletin 25: 5–10.
ciología y Ciencias Ambientales, and funded by grants from the Consejo Coria, R.A., Cladera, G. and Salgado, L. 1996. Sobre una nueva localidad fo-
Nacional de Investigaciones Científicas y Técnicas (CONICET 713/09), the silífera de la Formación Río Limay? Cretácico Superior temprano, en la lo-
Agencia Nacional de Promoción Científica y Tecnológica (FONCYT PICT- calidad de Cerro Bayo Mesa, Provincia de Neuquén. Ameghiniana 33: 463.
2011-2591), and the Universidad Nacional de Cuyo (06/M044) to B. Gon- Coria, R.A. and Calvo, J.O. 2002. A new iguanodontian ornithopod from
zález Riga. Neuquén Basin, Patagonia, Argentina. Journal Vertebrate Paleontology 22:
503–509.
Coria, R.A. and Currie, P.J. 2006. A new carcharodontosaurid (Dinosauria,
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Ameghiniana 34: 3–32. APPENDIX 1
Salgado, L. and Calvo, J.O. 1997. Evolution of titanosaurid sauropods. II:
The cranial evidence. Ameghiniana 34: 33–48. List of characters used for the phylogenetic analysis
Salgado, L. and Azpilicueta, C. 2000. Un nuevo saltasaurino (Sauropoda, Ti- 1. Short, deep snout: present (0); absent (1) (modified from Upchurch 1998
tanosauridae) de la provincia de Río Negro (Formación Allen, Cretácico
by Curry Rogers, 2005).
Superior), Patagonia, Argentina. Ameghiniana 37: 259–264.
Salgado L. and de Souza Carvalho, I. 2008. Uberabatitan ribeiroi, a new ti- 2. Frontal contribution to supratemporal fossa: present (0); absent (1) (Wil-
tanosaur from the Marilia Formation (Bauru Group, Upper Cretaceous), son and Sereno, 1998).
Minas Gerais, Brazil. Palaeontology 51: 881–901. 3. Frontal, dorsal texture: smooth (0); rugose (1) (Gallina and Apesteguía,
Salgado, L., Canudo, J.I., Garrido, A.C., Ruiz-Omeñaca, J.I., García, R.A., de 2011).
la Fuente, M.S., Barco, J.L. and Bollati, R. 2009. Upper Cretaceous 4. Postorbital, ventral process shape: transversely narrow (0); broader trans-
vertebrates from El Anfiteatro area, Río Negro, Patagonia, Argentina.
versely than anteroposteriorly (1) (Zaher et al., 2011).
Cretaceous Research 30: 767–784.
Salgado, L. and Powell, J.E. 2010. Reassessment of the vertebral laminae in 5. Postorbital, posterior process: present (0); absent (1) (Zaher et al., 2011).
some South American titanosaurian sauropods. Journal of Vertebrate Pa- 6. Postorbital, posterior margin articulating with the squamosal: with ta-
leontology 30: 1760–1772. pering posterior process (0); with a deep posterior process (1) (Zaher et
Santucci, R.M. and Bertini, R.J. 2006. A new titanosaur from western Sao al., 2011).
Paulo State, Upper Cretaceous Bauru Group, south-east Brazil. Palaeon- 7. Parietal occipital process, dorsoventral height: deep, nearly twice the di-
tology 49: 59–66. ameter of the foramen magnum (0); short, less than the diameter of the
Santucci, R.M. and Arruda-Campos, A.C. 2011. A new sauropod (Macrona-
foramen magnum (1) (Wilson, 2002).
ria, Titanosauria) from the Adamantina Formation, Bauru Group, Upper
Cretaceous of Brazil and the phylogenetic relationships of Aeolosaurini. 8. Parietal, elongate lateral process: absent (0); present (1) (Curry Rogers,
Zootaxa 3085: 1–33. 2005).
Sanz, J.L., Powell, J.E., Le Loeuff, J.L., Martínez, R. and Pereda-Suberbiola, 9. Parietal, cranial inclination with wide posterodorsal exposure of crest: ab-
X. 1999. Sauropod remains from the Upper Cretaceous of Laño (North- sent (0); present (1) (Salgado and Calvo, 1997).
central Spain). Titanosaur phylogenetic relationships. Estudios del Museo 10. Parietal, contribution to post-temporal fenestra: present (0); absent (1)
de Ciencias Naturales de Alava 14: 235–255. (Wilson, 2002).
Seeley, H.G. 1887. On the classification of the fossil animals commonly called
11. Parietal, distance separating supratemporal fenestrae: less (0); or twice (1)
Dinosauria. Proceedings of the Royal Society of London 43: 221–228.
Smith, J.B., Lamanna, M.C., Lacovara, K.J., Dodson, P., Smith, J.R., Poole, the transverse diameter of the supratemporal fenestra (Wilson, 2002).
J.C., Giegengack, R. and Attia, Y. 2001. A giant sauropod dinosaur from 12. Ascending process of premaxilla: directed dorsally (0); directed pos-
an Upper Cretaceous mangrove deposit in Egypt. Science 292: 1704–1706. terodorsally (1) (Gauthier 1986).
Upchurch, P. 1998. The phylogenetic relationships of sauropod dinosaurs. 13. External nares, configuration of lateral margin: lacrimal excluded, maxilla-
Zoological Journal of the Linnean Society 124: 43–103. nasal contact (0); lacrimal participates, separates maxilla and nasal (1)
Upchurch, P., Barrett, P.M. and Dodson, P. 2004. Sauropoda. In: D.B.
(Gallina and Apesteguía, 2011).
Weishampel, P. Dodson and H. Osmólska (Eds.), The Dinosauria (second
edition). University of California Press, Berkeley, p. 259–322. 14. Preantorbital fenestra: absent (0); present (1) (Wilson and Sereno, 1998).
Wilson, J.A. 1999. A nomenclature for vertebral laminae in sauropods and other 15. Supraoccipital, height: twice (0); or subequal (1) than the height of the
saurischian dinosaurs. Journal of Vertebrate Paleontology 19: 639–653. foramen magnum (Wilson, 2002).
Wilson, J.A. 2002. Sauropod dinosaur phylogeny: critique and cladistic analy- 16. Paraoccipital process, ventral non-articular process: absent (0); present (1)
sis. Zoological Journal of the Linnean Society 136: 217–276. (Wilson, 2002).
Wilson, J.A. 2006. An overview of titanosaur evolution and phylogeny. In: 17. Longitudinal groove on the supraoccipital: absent (0); present (1) (Curry
Colectivo Arqueológico-Paleontológico de Salas (Eds.), Actas de las 3as
Rogers, 2005).
Jornadas sobre Dinosaurios y su Entorno, Burgos, p. 169–190.
Wilson, J.A. and Sereno, P. 1998. Early evolution and higher level phylogeny 18. Basipterygoid processes, angle of divergence: approximately 45º (0); less
of sauropod dinosaurs. Journal of Vertebrate Paleontology 18: 1–68. than 30º (1) (Wilson, 1998).
Wilson, J.A., Martínez, R.N. and Alcober, O. 1999. Distal tail segment of a 19. Basal tubera, anteroposterior depth: approximately half the dorsoventral
titanosaur (Dinosauria-Sauropoda) from the Upper Cretaceous of Men- height (0); sheetlike, 20% the dorsoventral height (1) (Wilson, 1998).
doza, Argentina. Journal of Vertebrate Paleontology 19: 591–594. 20. Mandible shape: U shape (0); L shape (1) (Gallina and Apesteguía, 2011).
Wilson, J.A. and Upchurch, P. 2003. A revision of Titanosaurus Lydekker (Di- 21. Tooth shape: spoon-like (0); compressed cone chisel-like (1); pencil chisel-
nosauria - Sauropoda), the first dinosaur genus with a ‘Gondwanan’ dis-
like (2) (modified from Calvo, 1994 by Calvo and González Riga, 2003).
tribution. Journal of Systematic Palaeontology 1: 125–160.
Wilson, J.A., D’Emic, M.D., Ikejiri, T., Moacdieh, E.M. and Whitlock, 22. Tooth crowns, cross-sectional shape at mid-crown: D-shaped (0); sub-
J.A. 2011. A nomenclature for vertebral fossae in sauropods and other cilindrical with crest (1); cylindrical (2) (modified from Wilson and
saurischian dinosaurs. PLoS ONE 6: e17114. Sereno, 1998 by Calvo et al., 2007b).
23
23. Wear facets of teeth sharply inclined: absent (0); present (1) (Salgado and coelous-distoplatyan (1); slightly procoelous (2); strongly procoelous with
Calvo, 1997). prominent condyles (3) (modified from González Riga, 2003 by Gon-
24. Atlas, anteroventral border of the intercentrum: reduced (0); extended (1) zalez Riga et al. 2009).
(this paper). 52. Articular face shape on middle caudal centra: non-procoelous (0); pro-
25. Cervical vertebrae, number: 12 (0); 13 (1); 14 or more (2) (Upchurch, coelous-distoplatyan (1); slightly procoelous (2); strongly procoelous with
1998). prominent condyles (3) (modified from González Riga, 2003 by Gonza-
26. Lateral laminae on posterior cervical neural spines: absent (0); present (1) lez Riga et al. 2009).
(this paper). 53. Articular face shape on posterior caudal centra: non-procoelous (0);
27. Laterodorsally expanded cervical neural spines originated by lateral lami- slightly procoelous with reduced condyles (1); strongly procoelous with
nae, that reach or surpass the wide of the centra: absent (0); present (1) prominent condyles (2) (Gonzalez Riga et al. 2009).
(this paper). 54. Anterodorsal border of neural spine in middle caudal vertebrae located
28. Cervical prezygapophyses, relative length: articular facets that surpass (0); posteriorly with respect to anterior border of the postzygapophyses: ab-
or not surpass (1) the centra (Salgado et al., 1997). sent (0); present (1) (Salgado et al., 1997).
29. Neural spines in cervical vertebrae: tall (0); small (1) (Calvo et al., 2007b). 55. Anteriorly directed anterior caudal neural spine: absent (0); present (1)
30. Anterior cervical neural spines: bifid (0); single (1) (Upchurch, 1998). (Calvo et al., 2007c).
31. Posterior cervical vertebrae, proportions: ratio total height-centrum length; 56. Shape of the section of neural spines in most anterior caudal vertebrae
less (0); or more (1) than 1.5 (modified from Calvo and Salgado, 1995 by in dorsal view: axially elongated (0); transversely elongated (1); quad-
González Riga, 2005). rangular (2) (Calvo et al., 2007c).
32. Spinodiapophyseal fossa in posterior cervical vertebrae: absent (0); shal- 57. Neural arch in anterior caudal vertebrae: placed in the middle of the cen-
low or reduced (1); deep and extended (2) (González Riga, 2005). trum (0); anteriorly (1); on the anterior border (2) (modified from Sal-
33. Posterior cervical centra, proportions: ratio anteroposterior length/ height gado et al., 1997 by Calvo et al., 2007b).
of posterior face: >3 (0); 2,5 to 1,5 (1); less than 1,5 (2) (modified from 58. Neural spine in middle caudal vertebrae, shape: tall and short anteropos-
Wilson, 2002 by Calvo et al., 2007b). teriorly (0); low, anteroposteriorly elongated, laminated, with a sub-
34. Dorsal vertebrae, number: 12 or more (0); 11(1), 10 or fewer (2) (Wilson quadrangular contour (1) (modified from González Riga, 2003 by this
and Sereno, 1998). paper).
35. Anterior dorsal neural spines, shape: bifid (0); single (1) (McIntosh, 1990). 59. Length proportions of prezygapophyses with respect to the centrum
36. Anterior dorsal vertebrae, infrapostzygapophyseal fossa: absent (0); pres- length in middle caudal vertebrae: shorter than 50% (0); between 40 to
ent not divided (1); present divided in two subtriangular fossa (2) (Gal- 50% (1); longer than 50% (2) (modified from Gonzalez Riga 2003 by
lina and Apesteguía, 2011). Calvo et al., 2007b).
37. Anterior dorsal neural spines inclined posteriorly more than 20º from ver- 60. Ventral depression divided by a longitudinal septum in anterior and
tical: absent (0); present (1) (modified from Wilson and Sereno (1998) middle caudal vertebrae: absent (0); present (1) (Salgado and Azpilicueta
by González Riga, 2003). 2000).
38. Posterior dorsal neural spines, dorsal development: more (0); or less (1) 61. Postzygapophyseal process in middle caudal vertebra: absent (0); present
than 20% the total height of the vertebra (modified from Sanz et al. (1) (Calvo et al., 2007b).
1999 by González Riga, 2003). 62. Well-developed interprezigapophyseal lamina in middle caudal vertebrae:
39. Prespinal lamina in dorsal vertebrae: absent (0); present in the distal end absent (0); present (1) (Calvo et al, 2007b).
of neural spine (1); well developed up to the base of the neural spine (2) 63. Scapular glenoid orientation: relatively flat (0); strongly beveled medially
(modified from Salgado et al., 1997 by Bonaparte et al., 2006). (1) (Wilson and Sereno, 1998).
40. Centroparapophyseal lamina in posterior dorsal vertebrae: absent (0); pres- 64. Humerus, breadth of proximal end: less (0); equals or more than 50% (1)
ent (1) (Bonaparte and Coria, 1993). the humeral length (González Riga, 2003).
41. Ventrally widened or slightly forked centrodiapophyseal laminae in pos- 65. Humerus, type of proximal border: strongly curved (0); straight or slightly
terior dorsal vertebrae: absent (0); present (1) (Salgado et al., 1997). curved (1); sigmoidal (2) (modified from Upchurch, 1998 by González
42. Hyposphene-hypantrum articulation in dorsal vertebrae: present (0); ab- Riga, 2003).
sent (1) (Salgado et al., 1997). 66. Ulnar olecranon process, development: rudimentary, levels with prox-
43. Pleurocoels in dorsal vertebrae, shape: circular or elliptical (0); caudally imal articulation (0); prominent, projecting above proximal articulation
acuminate (1) (Salgado et al., 1997). (1) (Wilson and Sereno, 1998).
44. Camellate or somphospondylous types of internal structures of presacral 67. Sternal plates, shape: suboval (0); semulinar (1) (Salgado et al., 1997).
vertebrae: absent (0); present (1) (Wilson and Sereno, 1998). 68. Semilular sternal plate with straight caudal border: absent (0); present (1)
45. Sacral vertebrae, number: five (0); six of more (1) (McIntosh, 1990). (González Riga, 2003).
46. First caudal vertebrae, type: platycoelous (0); procoelous (1); opisthocoe- 69. Coracoid, shape: suboval (0); quadrangular (1) (Salgado et al., 1997).
lous (2); biconvex (3) (Salgado et al., 1997). 70. Pubis, length with respect to ischium length: shorter or equal (0); longer
47. Wide and deep interzigapophyseal cavity in proximal caudal vertebrae: (1) (Salgado et al., 1997).
absent (0); present (1) (Calvo et al., 2007b). 71. Metacarpals, distal phalangeal articular facets well developed: present (0);
48. Caudal transverse processes: disappear by caudal 15 (0); disappear by cau- absent (1) (Salgado et al., 1997).
dal 10 (1) (Wilson, 2002). 72. Ischium, posterior process twice or more the length of pubis articulation:
49. Anterior and middle caudal centra, proportions: as high as wide (0); de- present (0); absent (1) (modified from Salgado et al., 1997 by Calvo and
pressed, wider than high (1) (Salgado et al., 1997). González Riga, 2003).
50. Mid caudal centra with the anterior face strongly inclined anteriorly: ab- 73. Ischium, iliac pedicel: short and poorly developed (0); slender and well de-
sent (0); present (1) (Franco-Rosas et al., 2004). veloped (1); wide and well developed (2) (Calvo and González Riga,
51. Articular face shape on anterior caudal centra: non-procoelous (0); pro- 2003).
24
74. Shape of preacetabular lobe of ilium: moderately expanded (0); broadly ex- 81. Calcaneum: present (0); ossified calcaneum absent (1) (McIntosh, 1990).
panded and dorsally directed (1) (Salgado et al., 1997). 82. Osteoderm: absent (0); present, ellipsoid (1); present, keeled (2) (modi-
75. Orientation of preacetabular lobe of ilium: nearly vertical (0); nearly hor- fied from Curry Rogers, 2005 by this paper).
izontal and laterally projected (1) (Salgado et al., 1997).
76. Relative orientation of the pubic peduncle of ilium: angled (0); perpen-
dicular with respect to the sacral axis (1) (Salgado et al., 1997a).
77. Humerus/ femoral ratio of 0.90 o more: absent (0); present (1) (McIntosh,
1990).
78. Lateral bulge of femur, below the major trochanter: absent (0); present
(1) (McIntosh, 1990).
doi: 10.5710/AMEGH.26.12.1013.1889
79. Distal end of tibia broader transversely than anteroposteriorly: absent (0);
present (1) (Salgado et al., 1997).
80. Metatarsal I, length: shortest metatarsal (0); metatarsal V shorter than Recibido: 23 de octubre de 2013
metatarsal I (1) (Curry Rogers, 2005). Aceptado: 26 de diciembre de 2013
25

Quetecsaurus - Gonzalez Riga Ortiz David 2014

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AMEGHINIANA - 2014 - Tomo 51 (1): 3 – 25 ISSN 0002-7014

A NEW TITANOSAUR (DINOSAURIA, SAUROPODA)

Resumen. UN NUEVO TITANOSAURIO (DINOSAURIA, SAUROPODA) DEL CRETÁCICO SUPERIOR (FORMACIÓN

Relationship Ratio total

Atlas 3 2,3 5 — — 0,937

Posterior cervical vertebra 49 40 14 23 1,225 1,642 2,857

Anterior dorsal vertebra 39 — 20 24* — 1,95 —

Anterior caudal vertebra — 16 14 13 — — 1,143

Figure 9. Quetecsaurus rusconii gen. et sp. nov., UNCUYO-LD-300.7,

formes (Wilson and Sereno, 1998). The tuberculum is longer

Width in previous Width at posterior

Dorsal rib 132 6

is oriented toward the ventral side. Metacarpal IV is proxi-

PHYLOGENETIC ANALYSIS AND DISCUSSION

Figure 14. Quetecsaurus rusconii gen. et sp. nov., UNCUYO-LD-300.19,

Proximal epiphysis Distal epiphysis

Metacarpal I 28.5 7.6 12.8 4.4 11.3 6.4

Metacarpal II 31.2 7 9.8 9.3 10.4 8.6

Metacarpal III 29.2 4.5 9.2 8.7 10.2 7.3

Metacarpal IV 30.3 7.5 8.9 5.6 8.1 10.1

Metacarpal V 25.1 6.5 10.4 5.3 8.7 7.3

TABLE 4 - Matrix of characters used for the phylogenetic analysis

Camarasaurus 0011000001 1001000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 00

Brachiosaurus 0001000011 0000000010 1010100001 0010100010 0000000000 0000001000 0000000000 000101110? 00

Chubutisaurus ?????????? ?????????? ?????????? ????????2? 0001?0??00 100???1??0 ???01????? 0?????111? ??

Andesaurus ?????????? ?????????? ?????????? ????1??011 1011?0?000 2000001100 00?01????1 110???01?? ??

Malawisaurus 0??????11? ?0?????000 201??00001 00??11?011 1111??1000 32000?1110 0???1111?1 ?1??????1? ?1

Ligabuesaurus ?????????? ?????????? 101??00001 010?111011 1111?????? ?????????? ??10?????? 0?????111? ??

Mendozasaurus ?????????? ?????????? ?????1100? 122?120?2? ?111?11000 3210011110 00101111?? 1?????0111 ?1

Futalognkosaurus ?????????? ?????????? ???0211001 12221?0?2? ?111111?0? 3????11??0 ?????????1 ?12111???? ??

Quetecsaurus ???000???? ?????????? 2211?10001 010?1?0?2? ?1?1????00 3?????1??0 ??1?2?111? 1????????1 ??

Epachthosaurus ?????????? ?????????? ?????????? ????101121 101111?000 3320001110 0010111??1 1??1110111 10

Rapetosaurus 111?001110 1111111110 2210200001 00?1121121 11111?0000 33200110?0 001011100? 11011101?1 ?1

Drusilasaura ?????????? ?????????? ?????????? ????100?2? ?111?11?00 33?0011?10 001??????? ?????????? ?0

Rinconsaurus ?????????? ?????????? 211??00001 011?101121 11111???00 3320011010 1110111011 11111101?? ??

Muyelensaurus ?1????1??1 1???1?111? 211??0000? 011?121121 11111?0?00 3320011010 11101110?1 111111011? ??

Bonitasaura ?11???111? 1?1??????1 221??00?01 011?120121 1111?11000 3320011010 01?01?1??1 112???011? ??

Gondwanatitan ?????????? ?????????? ?????????? ???1?11211 111111???1 3320102020 01?0?????? ?0111???1? ??

Aeolosaurus ?????????? ?????????? 221??????? ????1???2? ?????30?01 3320112020 01?01?10?1 101????11? ?1

Opisthocoelicaudia ?????????? ?????????? ?????????? ???1001121 0111121100 0000021000 0011111011 1111110111 10

Alamosaurus ?????????? ?????????? 22???00001 001?1??120 0111131100 3320021000 0010111111 112111011? ??

Neuquensaurus ?????????? ?????????? ?????00101 012?101121 1111131?10 3321021000 001011101? 112111?11? ?2

Saltasaurus ?11?????11 1???01011? 221??00111 012?111121 1111110?10 3321021001 0011211011 112111011? ?2

Rocasaurus ?????????? ?????????? ?????0???? 01??101121 1111????10 33210?1001 00???????1 ?12111?1?? ??

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