Cretaceous Research 154 (2024) 105754

Contents lists available at ScienceDirect

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

A rebbachisaurid-mimicking titanosaur and evidence of a Late

Cretaceous faunal disturbance event in South-West Gondwana
Leonardo S. Filippi a, *, Rube

n D. Jua

rez Valieri b, Pablo A. Gallina c, Ariel H. Me

ndez d,
Federico A. Gianechini e, Alberto C. Garrido f, g
a
Museo Municipal Argentino Urquiza, Chos Malal 1277, 8319, Rinco
n de los Sauces, Neuqu
en, Argentina
b
Secretaría de Cultura de La Provincia de Río Negro, 8332, General Roca, Río Negro, Argentina
c
CONICET, Fundacio
n F

elix de Azara e Universidad Maimo
nides, CCNAA, Hidalgo 775, 1405, Buenos Aires, Argentina
d
CONICET, Instituto Patago
nico de Geología y Paleontología (CCT CONICET-CENPAT), Bv. Brown 2915, 9120, Puerto Madryn, Chubut, Argentina
e
gicas de San Luis (CONICET-UNSL), Ej

CONICET, Instituto Multidisciplinario de Investigaciones Biolo ercito de Los Andes 950, 5700, San Luis, San Luis,
Argentina
f
n Provincial de Minería, Etcheluz y Ej

Museo Provincial de Ciencias Naturales “Prof. Dr. Juan Olsacher”, Direccio ercito Argentino, 8340, Zapala, Neuqu
en,
Argentina
g
n en Geociencias de la Patagonia (CIGPat), Departamento de Geología y Petro
Centro de Investigacio
leo, Facultad de Ingeniería, Universidad Nacional del
Comahue (UNCo), Buenos Aires 1400, 8300, Neuqu
en Capital, Argentina

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

Article history: The evolution of ecosystems during the late Mesozoic on the southern landmasses is complex and still
Received 4 August 2023 poorly known. Starting from a single vicariant LaurasianeGondwanan scenario, the paleobiogeographic
Received in revised form and biostratigraphic models have become more complex, including vicariant, dispersal, and local ex-
17 October 2023
tinctions as major drivers of changes in the Cretaceous ecosystems during the isolation and posterior
Accepted in revised form 20 October 2023
Available online 30 October 2023
fragmentation of Gondwana. However, the direct effects of replacement and the adaptive evolution of
terrestrial vertebrates to fill vacant ecological niches after disruptive ecological events have been poorly
discussed. Here, we provide a preliminary description of a nearly complete new titanosaurian sauropod
Keywords:
Titanosauria
from the Upper Cretaceous of Patagonia, Inawentu oslatus gen. et. sp. nov., that shows remarkable
Adaptative convergence convergent anatomical traits with rebbachisaurid sauropods. A phylogenetic analysis recovers it within a
Upper Cretaceous not previously recovered titanosaurian subclade, named Clade A, which would be endemic to the Upper
Gondwana Cretaceous of South America. The convergent evolution between rebbachisaurids and Clade A members
is interpreted as the result of the same ecological niche exploitation. The biostratigraphic scenario during
the Late Cretaceous of South America leads to interpret rapid speciation of the titanosaurs because of
filling the empty ecological niche left by the extinction of the rebbachisaurids, an idea concordant with a
regional disturbance event of the ecosystems in this continent between 90 and 85 Ma.
© 2023 Elsevier Ltd. All rights reserved.

1. Introduction 2005; Perry et al., 2009; Clauss et al., 2013; Sander, 2013; Bates
et al., 2016). They represented the main mid to large-sized herbivo-
During the late Mesozoic, sauropod dinosaurs constituted the rous component of the fauna in most of these southern landmasses,
predominant herbivorous animals in all the non-polar Gondwanan both in diversity and abundance. In this way, it could be expected that
terrestrial ecosystems. With a general bauplan consisting of a the different sauropod clades radiated to cover most of the niche
quadrupedal and graviportal stance, a proportionally small cranial- partitioning opportunities, with different strategies from selective to
to-body ratio, and elongated cervical and caudal series, the sauro- generalists, and from high to low-level browsers. This ecological
pods evolved a notable variety of adaptations related to gigantism, radiation has been previously proposed for some sauropod clades
locomotion, defense, physiology, and feeding behaviors (Carrano, (Stevens and Parrish, 2005; Whitlock, 2011), and some aspects of this
have been discussed previously for Titanosauria (Martínez et al.,
2016; Poropat et al., 2021). Among sauropod dinosaurs proposed as
* Corresponding author. low browsers, three distinctive anatomical traits appear to be pre-
E-mail address: lsfi[email protected] (L.S. Filippi). sent: a short and sub-horizontal neck perpendicularly positioned

https://doi.org/10.1016/j.cretres.2023.105754
0195-6671/© 2023 Elsevier Ltd. All rights reserved.

rez Valieri, P.A. Gallina et al.
L.S. Filippi, R.D. Jua Cretaceous Research 154 (2024) 105754

concerning the ground; a broad symphyseal surface of the snout with Throughout the basin, the Bajo de la Carpa Formation exhibits a
teeth restricted to the anterior margin; and slender teeth. Diplodo- multiplicity of paleoenvironments developed under a domain of
coid sauropods can be considered as the classic low-level browsers, aeolian and fluvial depositional systems (Garrido, 2010). In the La
with diplodocids and dicraeosaurids as an abundant component of Invernada-Cerro Overo area, the development of an anastomosed
large herbivorous fauna during the Late Jurassic across North fluvial system characterized by the presence of muddy floodplain
America, Europe, and Gondwana, and also during the earliest Early and dominant sandy load paleochannels deposits has been estab-
Cretaceous in western Gondwana (Gallina et al., 2014; McPhee et al., lished for this unit (Cruzado-Caballero et al., 2018).
2016; Gallina et al., 2019). Nevertheless, the archetype of a ground- The recovered skeletal remains were found articulated in a
level browser sauropod is represented by the rebbachisaurids, horizon of massive, reddish-colored, edaphized mudstones which
which survived (and partially replaced) the other families of dip- are covered by a thin sandy layer (30 cm thick) linked to river
lodocoids, becoming an important component of the Cretaceous overbank flooding deposits. The degree of articulation observed in
sauropod faunas in South America, Africa, and Europe until these skeletal remains suggests that the carcass lies closer to the
CenomanianeTuronian times (Lavocat, 1954; Calvo & Salgado, 1995; place of death of the animal. The moderate to poor degree of
Bonaparte, 1996; Sereno et al., 1999; Carvalho et al., 2003; Torcida preservation of the bones is attributed to the action of the edaphic
Ferna
ndez-Baldor, F. et al., 2011; Mannion et al., 2011; Fanti et al., processes that affected these deposits during the development of
2013). The highly derived design present in rebbachisaurids that the floodplain, while the missing skeletal pieces have disappeared
tended towards low or ground-level browsing adaptations is indic- due to the erosion that affected the outcrops during the last times
ative of feeding specialization, and probably, based on snout shape of of the Quaternary.
Nigersaurus, generalist sauropods (Sereno et al., 2007). The dip-
lodocoid extinction, represented by the Rebbachisauridae, may have 4. Systematic paleontology
facilitated the expansion and diversification of derived titano-
saurians (Salgado, 2003). After the extinction of rebbachisaurids Sauropoda Marsh, 1878
(Turonian) the diversity of titanosaurs shows a marked increase in its Titanosauria Bonaparte and Coria, 1993
diversity (Mannion et al., 2013). Here we provide a preliminary Eutitanosauria sensu Carballido, Otero, Mannion, Salgado and
description of a largely complete new titanosaurian sauropod spec-
rez-Moreno, 2022
Pe
imen from the La Invernada area (Rinco
n de los Sauces, Neuque
n,
Inawentu oslatus gen. et. sp. nov.
Patagonia Argentina) (Fig. 1), which shows remarkable convergent
traits of its skull anatomy with rebbachisaurid sauropods. Exploring Etymology. Inawentu, from the Mapundung language ‘inawentu’,
the phylogenetic relationships of this titanosaur using cladistic an- meaning imitator or mimic; oslatus, from the Latin ‘os’, mouth, and
alyses, the new taxon is recovered within a clade of derived titano- ‘latus’, broad.
saurians, which include already known taxa (e.g., rinconsaurians), Holotype. MAU-Pv-LI-595 (Figs. 2e4). Partially complete and artic-
some of them sharing these low-browse adaptations. ulated specimen, composed of an almost complete skull, axial
sequence from the atlas to the last sacral element, and both ilia.
2. Materials and methods Diagnosis. Inawentu oslatus gen. et. sp. nov. is a titanosaurian
sauropod characterized by a series of autapomorphic characters
2.1. Anatomical abbreviations (marked with *) and a unique combination of characters: (1)*
posterior region of skull rotated posteroventrally so that it is on
an, angular; aof, antorbital fenestra; art, articular; bptp, the same level as the snout; (2)* foramen magnum located
basipterygoid process; bt, basal tuber; d, dentary; en, external subparallel to the tooth row; (3) spatulate snout, generated by
naris; f, frontal; fm, foramen magnum; j, jugal; la, lacrimal; lf, lateral expansion of the maxillae and dentaries; (4)* ridge at the
lacrimal fossa; ltf, laterotemporal fenestra; m, maxilla; na, nasal; contact between the ascending process and the body of the
orb, orbit; p, parietal; pmx, premaxilla; po, postorbital; pop, par- premaxilla, which generates a prominence in lateral view; (5)
aoccipital process; prf, prefrontal; pt, pterygoid; q, quadrate; qj, jugal with a highly elongated maxillary process and a wide and
quadratojugal; sa, surangular; sq; squamosal; so, supraoccipital; ventrally extended quadratojugal process, (6)* a double lateral
spl, splenial; stf, supratemporal fenestra. temporal fenestra formed by an anterior subcircular and a pos-
terior enlarged and laterally compressed opening; (7) quadran-
2.2. Institutional abbreviations gular jaw with highly developed lateral expansion; (8) dentition
restricted to the most anterior border of the maxillae and den-
CM e Carnegie Museum of Natural History, Pittsburgh, Penn- taries; (9) anteroposteriorly short jaw that represents half of the
sylvania, USA; MAU-Pv e Museo Municipal Argentino Urquiza, total skull length; (10)* jaw highest point, twice the height of the
Rinco
n de los Sauces, Neuque
n, Argentina; MCT e Museu de dentary, located in the articular sector between the coronoid
^ncias da Terra, Departamento Nacional de Produça
Cie ~o Mineral, Rio process and surangular; (11) short and slender neck, integrated
de Janeiro, Brazil. by 12 cervical vertebrae; and (12) anteroposteriorly long and
dorsoventrally low ilium.
3. Geological Setting Locality and Horizon. The area of La Invernada is located southwest
of Rinco
n de los Sauces city, Neuque
n province, Patagonia,
The specimen described here (MAU-Pv-LI-595) has been Argentina. Bajo de la Carpa Formation (Santonian, Upper Creta-
recovered in floodplain from fluvial system deposits belonging to ceous), Río Colorado Subgroup, Neuque
n Group of the Neuque
n
the Bajo de la Carpa Formation (Santonian) (Fig. 1), an integrant Basin (Fig. 1).
unit of the Río Colorado Subgroup, Neuque
n Group; from the Upper
Cretaceous of the Neuque
n Basin (Cazau & Uliana, 1973; Legarreta
& Gulisano, 1989; Garrido, 2010). The fossiliferous bearing level is 5. Description
located 11.1 m below the contact with the overlying Anacleto For-
mation, over a total thickness measured for the Bajo de la Carpa The skull of Inawentu (Fig. 3) resembles the elongated cranial
Formation at the study site of 129.6 m. morphology present in some titanosaurs such as Rapetosaurus
2
 L.S. Filippi, R.D. Jua

rez Valieri, P.A. Gallina et al.
3

Cretaceous Research 154 (2024) 105754

Fig. 1. Map showing the location of the La Invernada fossil site. The red star indicates the provenance of the holotype of Inawentu oslatus gen. et sp. nov. (MAU-Pv-LI-595). Location map based on satellite image acquired from Google
Earth (9 Jun 2021; Data SIO, NOAA, US Navy, NGA, GEBCO; Image Landsat/Copernicus).
 L.S. Filippi, R.D. Jua

rez Valieri, P.A. Gallina et al.
4

Cretaceous Research 154 (2024) 105754

Fig. 2. Articulated holotype of Inawentu oslatus gen. et sp. nov. (MAU-Pv-LI-595) in left lateral view. (A), skull; (B), atlas and axis; (C), cervical vertebrae from C3 to C10; (D), posterior cervical vertebrae C11eC12; (E), dorsal vertebrae
from D1 to part of D9; (F), posterior dorsal vertebrae from part of D9 and D10, complete sacrum (S1eS6), and left ilium. Scale bars for AeF: 10 cm. Scale bar for the silhouette: 1 m.

rez Valieri, P.A. Gallina et al.
L.S. Filippi, R.D. Jua Cretaceous Research 154 (2024) 105754

Fig. 3. Skull of Inawentu oslatus gen. et sp. nov. (MAU-Pv-LI-595) in left lateral (A), dorsal (B), and ventral (C) view. Diagonal lines represent zones covered with sediment and gray
zones are reconstructed parts. Scale bar: 10 cm.

(Curry Rogers & Forster, 2004), Tapuiasaurus (Zaher et al., 2011; 2005). Additionally, the skull has highly derived characters,
Wilson et al., 2016), and diplodocoids such as Nigersaurus including a posteriorly located skull roof, perpendicular to the
(Sereno et al., 2007), Diplodocus (skull CM 11161, McIntosh & tooth row (Fig. 3A), which is a feature more pronounced than in
Berman, 1975; Tschopp et al., 2015), and Bajadasaurus (Gallina Rapetosaurus, Tapuiasaurus, Nemegtosaurus, and Sarmientosaurus.
et al., 2019), although differing from those “boot-shaped” types Also, the squamosal and quadratojugal are located in a position
that are characteristic of brachiosaurids, such as Giraffatitan almost parallel to the longitudinal axis of the skull, accompanied
(Janensch, 1936) and Abydosaurus (Chure et al., 2010), some early by a strong reduction of the laterotemporal fenestra and an
branching titanosaurs, such as Sarmientosaurus (Martínez et al., anteroposterior shortening of the jaw, due to the reduction of the
2016) and other titanosaurs, such as Nemegtosaurus (Wilson, angular and surangular (Fig. 3A).
5

rez Valieri, P.A. Gallina et al.
L.S. Filippi, R.D. Jua Cretaceous Research 154 (2024) 105754

6

rez Valieri, P.A. Gallina et al.
L.S. Filippi, R.D. Jua Cretaceous Research 154 (2024) 105754

The jaw is quadrangular, a shared feature with other titanosaurs, The complete neck of Inawentu is composed of 12 vertebrae,
such as Antarctosaurus (Huene, 1929), Bonitasaura (Apesteguía, thus being the titanosaur with the least number of cervical verte-
2004; Gallina & Apesteguía, 2011), Brasilotitan (Machado et al., brae known so far. This contrasts with the 17 and 14 cervical
2013), and Baalsaurus (Calvo & Gonz
alez Riga, 2019) (Fig. 4). vertebrae observed in Rapetosaurus and Futalognkosaurus (Curry
However, the latter taxa are represented by fragmentary specimens Rogers, 2009; Calvo et al., 2007a), respectively. The remaining
that prevent knowing the degree of their skull modifications titanosaurian specimen known with a complete cervical sequence
(except for Antarctosaurus which has some parts of the skull). The is an unnamed taxon from Brazil (MCT 1487-R) and contains 13
laterally expanded maxilla of Inawentu (Fig. 3B) is similar to that of cervical vertebrae. The low number of cervical vertebrae and the
Narambuenatitan (Filippi et al., 2011), which allows us to infer that elongation index (EI ¼ 4.3 in C6) in Inawentu, results in a propor-
the latter taxon would have had a similar quadrangular jaw. The tionally shorter neck than that of other titanosaurs such as Futa-
new taxon shares with Rapetosaurus an anteroposteriorly elongated lognkosaurus and Rapetosaurus. The cervical ribs were preserved in
antorbital fenestra, although in Inawentu it is more posterodorsally almost complete articulation. These are relatively short, and their
located and close to the maxilla-lacrimal contact (Fig. 3AeB). posterior ends reach the posterior edge of the centrum of the
Although the occipital condyle is not preserved, the position of the subsequent vertebra (Fig. 5B), a condition that is observed in the
foramen magnum with respect to the skull roof indicates a highly anterior cervical vertebrae. Thus, the length of the cervical ribs is
derived ventral deflection of the snout, as present in the rebac- lesser than that observed in titanosauriforms, in which these ele-
chisaurid Nigersaurus (Sereno et al., 2007). The isolated braincases ments extend up to three vertebrae posteriorly (Cerda, 2009). This
of Antarctosaurus, Saltasaurus (Powell, 1992, 2003), and Bonatitan feature is reminiscent of the anteroposteriorly short cervical ribs of
(Martinelli & Forasiepi, 2004; Salgado et al., 2014) could indicate a most diplodocimorphs. The cervicodorsal vertebrae have highly
similar skull configuration. The quadratojugal shows two ventral modified centra and neural arches and are interpreted as a pivotal
hypertrophied structures in its mandibular ramus, homologous to point enabling multidirectional movement for the entire neck.
those present in Tapuiasaurus and Nemegtosaurus, but notably more Except for the atlas-axis (Fig. 5A), C11 and C12 (Fig. 5C), the
developed. Besides, the anterior portion of the quadratojugal is other cervical vertebrae are morphologically similar (Fig. 5B),
dorsoventrally deep, extending dorsally to the contact with the showing opisthocoelous centra, a slightly anteroposteriorly
jugal (Fig. 3A). The jugal is tetraradiated as in most titanosaurs, concave ventral surface, with slightly excavated lateral surface. The
although the maxillary process is highly elongated and the quad- cervical vertebrae lack the two different PODL segments present in
ratojugal process is wide and ventrally extended, a condition not Bonitasaura (Gallina & Apesteguía, 2015; Fig. 3C). The C11 and C12
present in other related taxa (e.g., Nemegtosaurus, Rapetosaurus). A are different morphologically from the remaining cervical verte-
divided laterotemporal fenestra (LTF) is present as in Tapuiasaurus, brae and they also differ from each other (Fig. 5C). The C11 has an
although Inawentu shows an anterior subcircular opening and a anteroposteriorly elongated centrum with an anteroposteriorly
posterior, laterally compressed, and larger fenestra. In Tapuiasaurus, concave ventral surface like the preceding cervical vertebrae,
the LTF is 8-shaped, divided by a second postorbital squamosal although it is transversely wider. The neural spine of C11 is very
contact, into a smaller posterodorsal opening and a larger, elon- prominent, located at the level of the middle of the vertebral
gated anteroventral opening (Wilson et al., 2016). Inawentu present centrum, tilted slightly posteriorly, and the neural spine has lateral
a LTF divided by a “bone bridge” formed by the contact of the jugal- expansions that resemble those present in Futalognkosaurus,
quadratojugal and jugal-squamosal (Fig. 3A). Mendozasaurus (Gonza
lez Riga, 2005), and Bonitasaura. Overall, the
The dentition, is restricted to the most anterior sector of the C12 is similar to the C11, except that in C12 the centrum is shorter
snout, even more than in the titanosaurids Nemegtosaurus and anteroposteriorly, the neural arch is higher, with pre and post-
Tapuiasaurus, but not as anterior as in the rebbachisaurid Lav- zygapophyses at the same level, and the neural spine is slenderer
ocatisaurus (Canudo et al., 2018). The snout is markedly trans- and anteroposteriorly more compressed. The C12 lacks a ventral
versely wide, developing a spatulate distal morphology keel or a ventral pneumatic fossa, such as that present in the last
(Fig. 3AeB). It is wider than the transverse width of the skull at cervical vertebra of Overosaurus (Coria et al., 2013).
the orbit level, as in Antarctosaurus and rebbachisaurid sauro- As in other titanosaurs, the dorsal series of Inawentu is
pods. The dentaries have a quadrangular L-shaped morphology composed of ten opisthocoelous vertebrae with acuminated lateral
(Figs. 3C and 4AeB), as in some titanosaurs, such as Antarcto- pleurocoels. The first dorsal vertebra has the most ante-
saurus, Bonitasaura, Brasilotitan, and Baalsaurus (Fig. 4CeF), and roposteriorly elongated centrum, with elongated and laterally
in the rebbachisaurids Lavocatisaurus and Nigersaurus. Both Ina- projected transverse processes. As in Bonitasaura, the neural spine
wentu and Antarctosaurus and to a lesser degree, Brasilotitan of the first dorsal vertebra is dorsoventrally tall, unlike the verti-
developed a markedly distinctive laterodistal deflection of the cally shortened neural spine of Overosaurus. The inclination of the
tooth row just in the joining zone of the symphyseal portion and neural spine serially varies in Inawentu, it is sub-vertically oriented
the posterior rami of the dentary. This morphology is homolo- in the anteriormost dorsal vertebrae, then progressively projects
gous to that present in the rebbachisaurid Nigersaurus. Inawentu backward in the middle dorsals, and it returns to a vertical orien-
presents a square jaw with highly developed lateral expansion tation in the posteriormost elements. Although this variation in
(Fig. 4AeB) which clearly differentiates it from the slight neural spine angle is observed in some titanosaurs, it is more
expansion observed in Antarctosaurus (Fig. 4D). In Inawentu the similar to that of Bonitasaura (Gallina, 2011), Trigonosaurus
symphysial sector is relatively robust and wide, similar to Baal- (Campos et al., 2005), and Overosaurus, than in Rapetosaurus,
saurus and different from Antarctosaurus and Brasilotitan, which Opisthocoelicaudia (Borsuk-Bialynicka, 1977), and the saltasaurines.
have an anteroposteriorly thinner symphysis. On the other hand, The sacrum consists of six vertebrae and has a similar
Inawentu shares vertical walls with Antarctosaurus, Brasilotitan morphology to that of other titanosaur sauropods, with the
and Baalsaurus, a condition that differentiates it from Bonita- exception of Neuquensaurus (Salgado et al., 2005) which has seven
saura, which presents a curved symphysial sector. sacral vertebrae. The last sacral centrum has a convex posterior

Fig. 4. Compared jaws of basally branching titanosaurs of Clade A, in dorsal view. Inawentu oslatus gen. et sp. nov. (MAU-Pv-LI-595) (A and B); Brasilotitan (C); Antarctosaurus
(D); Bonitasaura (E); and Baalsaurus (F). The black arrow indicates the portion of the dentary displaced from its original position. The images are not in scale.

7

rez Valieri, P.A. Gallina et al.
L.S. Filippi, R.D. Jua Cretaceous Research 154 (2024) 105754

8

rez Valieri, P.A. Gallina et al.
L.S. Filippi, R.D. Jua Cretaceous Research 154 (2024) 105754

surface, and consequently, the first caudal centrum (not preserved) European taxa. On the other hand, in Mannion et al. (2019b),
must have been procoelous. The first sacral vertebra has a delicate Antarctosaurus is recovered in a clade with the Rinconsauria.
rib, similar to the last dorsal ribs, which rests on the anterior edge of Although defined in different ways (node based vs. stem-based),
the ilium as in Overosaurus, Trigonosaurus, and Futalognkosaurus. Lognkosauria clusters the same taxa as Colossosauria in this phy-
Although they are incomplete, the sacral ribs of S6 possibly did not logeny, similarly to the results obtained by Pe
rez Moreno et al.
come into direct contact with the ilia. Instead, the S6 sacral ribs (2022). Besides, the new topology recovered Colossosauria
appear to be articulated both with the fifth sacral ribs and to the (Lognkosauria þ Rinconsauria) outside of Lithostrotia such as the
ilium, as in Overosaurus, Trigonosaurus, and Malawisaurus (Gomani, analyses by Pe
rez Moreno et al. (2022), which differs from previous
2005). The pubic and ischial pedicel of the ilium are not preserved. analyses (Gonza
lez Riga et al., 2019; Hechenleitner et al., 2020;
The iliac blade is well expanded anteroposteriorly as in other tita- Gallina et al., 2021). In our analysis, the Clade A is composed of a
nosaurs, but the lateral projection of the preacetabular process is subclade integrated by two branches, one containing Bonitasaura as
less prominent than in Saltasaurus and Neuquensaurus, which have the sister taxon of Inawentu plus Antarctosaurus. The second in-
laterally wider hips. cludes Narambuenatitan and a series of derived taxa, as the other
squared-jaw Brasilotitan and several forms which have an unknown
snout shape, such as Uberabatitan (Salgado & De Souza Carvalho,
6. Phylogenetic analysis 2008), Muyelensaurus (Calvo et al., 2007b), Rinconsaurus (Calvo &
Gonza
lez Riga, 2003), Overosaurus, Trigonosaurus, and Arrudatitan
To elucidate the phylogenetic placement of Inawentu, and other (Silva Junior et al., 2021) (Fig. 6). Inawentu includes some
related forms among the titanosaurian sauropods, we generated a anatomical traits shared with both Antarctosaurus and Bonitasaura.
new dataset primarily derived from that of Gallina et al. (2021). This Antarctosaurus and Inawentu overlap part of the skull, and Bonita-
dataset (see Supp. Data) was modified increasing in several cranial saura and Inawentu overlap in the skull, cervical and dorsal verte-
characters and taxa-preserving cranial elements to highlight brae. Nevertheless, based on the few comparable materials
morphological differences among titanosauriforms and titano- between these three taxa, such as the dentary, a more similar
saurians in particular. The resultant dataset is composed of 421 morphology is observed between Inawentu and Antarctosaurus
characters and 98 operational taxonomic units. The analysis was than Bonitasaura. An example of this is the development of the
carried out using the software T.N.T. 1.5 (Goloboff and Catalano, anterolateral corner of the dentary (character 421). Another taxon
2016). Characters were ordered as in the original analysis. The known by very fragmentary cranial material, Baalsaurus, was also
chosen parameters included the tree bisection and reconnection included in a posteriori analysis since it presents a derived squared
algorithm (TBR), with 1000 replications of Wagner trees and 10 dentary and represents the oldest known evidence of this
trees saved per replication. This procedure retrieved 890 most morphology in the fossil record, as it is found in Coniacian strata
parsimonious trees (MPTs) of 1521 steps (CI: 0.34; RI: 0.71), found (Calvo & Gonza
lez Riga, 2019). Baalsaurus is recovered within Clade
in 89 of the replicates. The strict consensus shows Inawentu as a A in the strict consensus analysis, although the precise placement
sister taxon to Antarctosaurus within an unresolved group, which of this taxon within the clade remains uncertain. Its inclusion
includes titanosaurs forms from the TuronianeMaastrichtian time generates a clade that is only supported by a single character
lapse (Supplementary information Fig. S2). In a second analysis (character 99).
using the TNT command “Pruned trees”, seven unstable taxa are
eliminated allow improving relationships within unresolved group,
positioning to Uberabatitan as a sister taxon to Brasilotitan 7. Discussion
(Supplementary information Fig. S3). Finally, a majority rule
consensus was made (þ50) (Supplementary information (Fig. S4)), The Clade A consisting in a group of deeply nested titanosaurs
where Inawentu is recovered as a deeply nested titanosaur in close that were restricted to the last stages of the Late Cretaceous in
association with several Late Cretaceous taxa from South America South America. The discovery of new materials and different data
that share a derived squared mandibular shape, such as Antarcto- sets that provide new morphological information will allow us to
saurus and Bonitasaura (Fig. 4). These three taxa form the sister provide better support in future phylogenies that confirm the
group of a clade that includes other squared-snout titanosaurs. i.e., presence of this clade of square-jawed titanosaurs.
Narambuenatitan and Brasilotitan, and Rinconsauria plus Aeolo- Inawentu and probably other members of this clade, possess
saurini (Fig. 6). The latter two clades, however, do not preserve apparent trophic adaptations seen in the preceding rebbachisaurid
cranial elements (except Muyelensaurus) and are characterized by sauropods, such as a broad snout and a relatively short neck. In this
mid-to-small-sized and slender bodies. The new topology recov- context, the shortened cervical series of Inawentu could be in
ered a clade of square-jawed titanosaurian taxa, here named Clade concordance with low-browsing feeding behavior. This would have
A, supported by three synapomorphies (Ch. 99-2, tooth rows, shape paleoecological implications, such as a faunistic turnover in post-
of anterior portions: rectangular, tooth-bearing portion of jaw Turonian times in the Gondwanan continental ecosystems and
perpendicular to jaw rami; Ch. 100-3, tooth rows, length: restricted low-browsing dietary habits in two different lineages of sauropod
anterior to subnarial foramen; and Ch. 415-0, ilium, postacetabular dinosaurs.
posteroventral edge: open concave). Consequently, these results Among deeply nested sauropods, i.e., the Neosauropoda, two
show a novel distant relationship between Lognkosauria and Rin- principal lineages diverged near the Lower to Middle Jurassic: the
consauria in a database that is mainly focused on South American macronarians and the diplodocoids (Xu et al., 2018). These are
taxa (contrasting with the results of Carballido et al., 2017; abundant components in the Late Jurassic faunas in North America,
Gonzalez Riga et al., 2019). A similar topology is only recovered in Europe, Africa, and South America (Marsh, 1884; Rauhut et al.,
Gorscak & O'Connor (2016) and subsequent phylogenetic results 2015; Mannion et al., 2017, 2019a). These Late Jurassic sauropod
obtained from that dataset, which also includes several African and faunas are indicative of the existence of niche partition preferences

Fig. 5. Cervical vertebrae of Inawentu oslatus gen. et sp. nov. (A), Atlas (C1) and Axis (C2); (B), Cervical C4 and C5; (C), Cervical C11 and C12. Anatomical abbreviations: At, Atlas; Ax,
Axis; ce, centrum, cr, cervical rib; ic, intercentrum; ns, neural spine; pa, parapophysis; podl; postzygodiapophyseal lamina; pos, postzygapophysis; pre, prezigapophysis; tp,
transverse process. Scale bar: 10 cm.

9

rez Valieri, P.A. Gallina et al.
L.S. Filippi, R.D. Jua Cretaceous Research 154 (2024) 105754

10

rez Valieri, P.A. Gallina et al.
L.S. Filippi, R.D. Jua Cretaceous Research 154 (2024) 105754

Fig. 7. Life reconstruction of Inawentu oslatus gen. et sp. nov. (MAU-Pv-LI-595). Illustration by Gabriel Lío.

between the low-browser feeder diplodocoids and the higher- macronarians and basal titanosauriforms (Janensch, 1936; Madsen
browser basal macronarians and basal titanosauriforms et al., 1995; Marpmann et al., 2015; Chure et al., 2010; Moore et al.,
(Whitlock, 2011). The cranial morphology of the basal 2018; D'Emic & Carrano, 2020) is plesiomorphic and more

Fig. 6. Phylogenetic relationships of Inawentu oslatus gen. et sp. nov. (MAU-Pv-LI-595). Time-calibrated simplified majority rule (þ50%) consensus tree. L, Lower; U, Upper; Alb,
Albian; Apt, Aptian; Bar, Barremian; Ber, Berriasian; Cam, Campanian; Cen, Cenomanian; Con, Coniacian; Hau, Hauterivian; Maa, Maastrichtian; San, Santonian; Tur, Turonian;
Vlg, Valanginian. The titanosaur's jaw images (Rapetosaurus is reversed) are not in scale.

11

rez Valieri, P.A. Gallina et al.
L.S. Filippi, R.D. Jua Cretaceous Research 154 (2024) 105754

reminiscent to that of other clades of non-neosauropod eusauro- Description of cranial material Diamantinasaurus matildae
pods (Ouyang & Ye, 2002; Royo-Torres & Upchurch, 2012; Poropat published after the conclusion of this study
& Kear, 2013). In contrast, diplodocids were highly modified in their
cranial morphology, as evidenced in the relative position of the Posterior to the finalization of this manuscript has been pub-
occipital condyle with respect to the tooth row, the position of the lished the description of cranial material of the basal titanosaur
nares, the slender tooth morphology, and the teeth placement in Diamantinasaurus matildae (Poropat et al., 2023). It displays a suit of
the front of the snout (Holland, 1906; Berman & McIntosh, 1978; anatomical characters that are congruent with those of basal tita-
Whitlock et al., 2010; Tschopp & Mateus, 2017; Woodruff et al., nosaurs as Sarmientosaurus and Choconsaurus, but contrast with
2018). the derived morphology present in Inawentu and their close rela-
During the Early Cretaceous, the diplodocoids appear to have tives as Antarctosaurus, Muyelensaurus or Bonitasaura. Although
been globally restricted to Europe and Gondwana (Africa and South Diamantinasaurus is not included in the present phylogenetic
America). At this time, the three diplodocoid families of the group, analysis, the proper analysis of these authors is not in conflict with
i.e., the diplodocids, the dicraeosaurids, and the rebbachisaurids our results, displaying Diamantinasaurus as a basal member of the
were present in at least the first stages of the Early Cretaceous group closely related with another ‘Middle’ Cretaceous taxa from
(Calvo & Salgado, 1995; Haluza et al., 2012; Ibiricu et al., 2012, 2013; Australia and southern South America and recovering an unnamed
McPhee et al., 2016; Bellardini et al., 2022). After the Barremian, the clade including Rinconsauria and Aelosaurini, including Antarcto-
rebbachisaurids are the only surviving family of low-browsing saurus within this last group. We believe that future works, some of
diplodocoid sauropods, reaching Turonian times (Gallina & these currently under development by some of the current authors,
Apesteguía, 2005; Haluza et al., 2012; Ibiricu et al., 2013; should include more taxa and characters in order to test our results
Bellardini et al., 2022). On the other hand, the titanosauriforms are and those of another authors.
broadly present in the known faunal contents of large landmasses.
During the early Late Cretaceous (CenomanianeTuronian) the Data availability
rebbachisaurids appear to be restricted to Africa and South
America, which possibly corresponds to a relictual presence of Data will be made available on request.
the family in both landmasses since they drifted and reached a
complete separation at the Cenomanian times. This could have
Acknowledgments
favored the appearance of independent and vicariant lineages,
although both, Africa and South America, suffered sporadic ex-
We would like to thank ExxonMobil S.A. Exploration Argentina
changes with other continental masses during the rest of the
for their funding fieldwork in La Invernada e Cerro Overo area
Cretaceous (McLoughlin, 2001; Ezcurra & Agnolín, 2012; Sallam
(2013e2015) and the Municipality of Rinco
n de los Sauces city for
et al., 2018). The last records of Rebbachisauridae in South
supporting the paleontological research projects. Thanks to the ex-
America are dated from the early Turonian, following a regional
technicians of the Museo Municipal “Argentino Urquiza”: A.
extinction (Calvo et al., 2006; Ibiricu et al., 2020). This extinction
Schenkel, C. Fuentes, and S. Palomo (the latter who found the
event appears to have affected multiple components of the
specimen presented here), for their participation during the field-
tetrapod fauna of South America, including not only rebbachi-
work. Our special thanks to H. Zaher and B. Navarro for providing
saurids but also basal lineages of titanosauriform sauropods (e.g.,
Tapuiasaurus photographs. C. Woodruff, V. Diez Diaz, P. Mannion
Andesaurus, Epachthosaurus), multiple lineages of theropods,
and P. Mocho, are acknowledged for comments in early stages of
such as spinosaurids, carcharodontosaurids, bahariasaurids,
the manuscript. The final version of the manuscript was subjected
elaphrosaurines, and non-furileusaurian abelisaurids; and the
to a language review by E. Galanda. Finally, we would like to thank
uruguaysuchid and candidodontid crocodylomorphs (Apesteguía,
the editor E. Koutsoukos and the reviewers J.L. Carballido and M.E.
2002; Coria & Salgado, 2005; Novas, 2009; Novas et al., 2005,
Hechenleitner for the corrections and useful comments that
2013; Delcourt et al., 2020; Meso et al., 2021).The change be-
improved the quality of this contribution. Financial support was
tween the “mid-Cretaceous” (AlbianeCenomanian) and post-
provided by Foncyt, PICT-2012-I-INVI-00513 (LSF).
Turonian vertebrate assemblage, is identified as a major faunal
turnover (Krause et al., 2020). In turn, multiple adaptive radia-
tions are recorded in the derived titanosaurs, such as the major
References
diversification of large-body sized lognkosaurians, the small-
body sized saltasaurines, and the square-jawed titanosaurs such Apesteguía, S., 2002. Successional structure in continental tetrapod faunas from
as Baalsaurus and Inawentu (Fig. 6). Adaptive radiations also are Argentina along the Cretaceous. In: Boletim do VI Simpo
sio sobre o Cret

aceo do
Brasil. II Simposio sobre el Cret
acico de Ame
rica del Sur (Sao Pedro, Brasil),
observed between the megaraptorids and furileusaurian thero-
Abstracts, pp. 135e141.
pods, and the deeply nested notosuchians, peirosaurids, and Apesteguía, S., 2004. Bonitasaura salgadoi gen. et sp. nov.: A break sauropod from
sebecosuchian crocodylomorphs (Calvo and Gonza
lez Riga, 2019; the Late Cretaceous of Patagonia. Naturwissenchaften 91, 493e497.
Baiano et al., 2022). As previously proposed, the extinction of the Baiano, M.A., Pol, D., Bellardini, F., Windholz, G.J., Cerda, I.A., Garrido, A.C.,
Coria, R.A., 2022. Elemgasem nubilus: a new brachyrostran abelisaurid (Ther-
large herbivorous and carnivorous dinosaurs from South America opoda, Ceratosauria) from the Portezuelo Formation (Upper Cretaceous) of
and Africa appears to be synchronic, but it can be likely an arti- Patagonia, Argentina. Papers in Palaeontology 8, e1462. https://doi.org/10.1002/
fact since in Africa there is a very scarce post-Cenomanian fossil spp2.1462.
Bates, K.T., Mannion, P.D., Falkingham, P.L., Brusatte, S.L., Hutchinson, J.R., Otero, A.,
record. Before Inawentu (Fig. 7) discovery, the extreme adapta- Allen, V., 2016. Temporal and phylogenetic evolution of the sauropod dinosaur
tions in the snout and neck it was just reported for non- body plan. Royal Society Open Science 3 (3), 150636.
titanosaur rebbachisaurid. So, perhaps the empty niche that Bellardini, F., Filippi, L.S., Garrido, A.C., Carballido, J.L., Baiano, M.A., 2022. New
rebbachisaurid remains from the Huincul Formation (Middle
rebbachisaurids left (ground-level browsers with quadrangular CenomanianeEarly Turonian) of the Central Neuque
n Basin, Patagonia,
snout) was rapidly occupied by some lineages of titanosaurs, here Argentina. Publicacio
n Electro
nica de la Asociacio

n Paleontolo
gica Argentina 22
named clade A. Although is evident a faunistic replacement event (2), 1e24.
Berman, D.S., Mcintosh, J.S., 1978. Skull and relationships of the Upper Jurassic
followed by radiation adaptative events, it remains obscure the sauropod Apatosaurus (Reptilia, Saurischia). Bulletin of Carnegie Museum of
causes of the fauna changes. Natural History 8, 1e35.

12

rez Valieri, P.A. Gallina et al.
L.S. Filippi, R.D. Jua Cretaceous Research 154 (2024) 105754

Bonaparte, J.F., 1996. Cretaceous Tetrapods of Argentina. In: Contributions on the Gallina, P.A., 2011. Notes on the axial skeleton of the titanosaur Bonitasaura salgadoi
Southern South America to Vertebrate Paleontology e Münchner Geo- (Dinosauria - Sauropoda). Anais da Academia Brasileira de Cie ^ncias 83 (1),
wissenschaftliche Abhandlungen (A), vol. 30, pp. 73e130. 235e245.
Borsuk-Bialynicka, M., 1977. A new camarasaurid sauropod Ophistocoelicaudia Gallina, P.A., Apesteguía, S., 2005. Cathartesaura anaerobica gen. et sp. nov., a new
skarzynskii gen. n., sp. n. from the Upper Cretaceus of Mongolia. Palaeontolo
gica rebbachisaurid (Dinosauria, Sauropoda) from the Huincul Formation (Upper
Polonica 37, 5e64. Cretaceous), Río Negro, Argentina. Revista del Museo Argentino de Ciencias
Calvo, J.O., Gonz
alez Riga, B., 2003. Rinconsaurus caudamirus gen. et sp. nov., a new Naturales 7, 153e166.
titanosaurid (Dinosauria, Sauro
poda) from the late Cretaceous of Patagonia, Gallina, P.A., Apesteguía, S., 2011. Cranial anatomy and phylogenetic position of the
Argentina. Revista Geologica de Chile 30 (2), 333e353. titanosaurian sauropod Bonitasaura salgadoi. Acta Palaeontologica Polonica 56
Calvo, J.O., Gonza
lez Riga, B.J., 2019. Baalsaurus mansillai gen. et sp. nov. a new (1), 45e60.
titanosaurian sauropod (Late Cretaceous) from Neuque
n, Patagonia, Argentina. Gallina, P.A., Apesteguía, S., 2015. Postcranial anatomy of Bonitasaura salgadoi
1. Anais da Academia Brasileira de Cie ^ncias 91, e20180661. (Sauropoda, Titanosauria) from the Late Cretaceous of Patagonia. Journal of
Calvo, J.O., Salgado, L., 1995. Rebbachisaurus tessonei sp. nov. A new Sauropoda from Vertebrate Paleontology 35, dx. https://doi.org/10.1080/02724634.2014.924957.
the Albian-Cenomanian of Argentina; new evidence on the origin of the Dip- Gallina, P.A., Apesteguía, S., Haluza, A., Canale, J.I., 2014. A diplodocid sauropod
lodocidae. Gaia 11, 13e33. survivor from the early cretaceous of South America. PLoS One 9, e97128.
Calvo, J.O., Gandossi, P., Porfiri, J.D., 2006. Dinosaur faunal replacement during Gallina, P.A., Apesteguía, S., Canale, J.I., Haluza, A., 2019. A new long-spined dinosaur
Cenomanian times in Patagonia, Argentina. In: The 9 International Symposium from Patagonia sheds light on sauropod defense system. Scientific Reports 9,
on Mesozoic Terrestrial Ecosystems, pp. 17e20. 1392.
Calvo, J.O., Porfiri, J.D., Gonz
alez Riga, B.J., Kellner, A.W.A., 2007a. Anatomy of Gallina, P.A., Canale, J.I., Carballido, J.L., 2021. The earliest known titanosaur
Futalognkosaurus dukei Calvo, Porfiri, Gonza
lez Riga & Kellner, 2007 (Dino- sauropod dinosaur. Ameghiniana 58 (1), 35e51.
sauria, Titanosauridae) from the Neuque
n Group (Late Cretaceous), Patagonia, Garrido, A.C., 2010. Estratigrafía del Grupo Neuque
n, Creta

cico Superior de la
Argentina. Arquivos do Museu Nacional 65, 511e526. Cuenca Neuquina (República Argentina): Nueva propuesta de ordenamiento
Calvo, J.O., Gonza
lez Riga, B.J., Porfiri, J.A., 2007b. A new titanosaur sauropod from
fico. Revista del Museo Argentino de Ciencias Naturales Nueva
litoestratigra
the Late Cretaceous of Neuque
n, Patagonia, Argentina. Arquivos do Museu Serie 12 (2), 121e177.
Nacional 65, 485e504. Goloboff, P.A., Catalano, S.A., 2016. TNT version 1.5, including a full implementation
Campos, D.A., Kellner, A.W.A., Bertini, R.J., Santucci, R.M., 2005. On a Titanosaurid of phylogenetic morpho-metrics. Cladistics 32 (3), 221e238. https://doi.org/
(Dinosauria, Sauropoda) vertebral columna from the Bauru group, Late Creta- 10.1111/cla.12160.
ceous of Brazil. Arquivos do Museu Nacional, Rio de Janeiro 63 (3), 565e593. Gonza
lez Riga, B.J., 2005. Nuevos restos fo
siles de Mendozasaurus neguyelap (Sau-
Canudo, J.I., Carballido, J.L., Garrido, A.C., Salgado, L., 2018. A new rebbachisaurid ropoda: Titanosauridae) del Cret
acico Tardío de Mendoza, Argentina. Ame-
sauropod from the Aptian-Albian, Lower Cretaceous Rayoso Formation, Neu- ghiniana 42, 535e538.
que
n Argentina. Acta Palaeontologica Polonica 63, 679e691. Gonza
lez Riga, B.J., Lamanna, M.C., Otero, A., Ortíz David, L.D., Kellner, A.,
Carballido, J.L., Pol, D., Otero, A., Cerda, I.A., Salgado, L., Garrido, A.C., Ramezzani, J., Ibiricu, L.M., 2019. An overview of the appendicular skeletal anatomy of South
Cúneo, N.R., Krause, M.J., 2017. A new giant titanosaur sheds light on body size American titanosaurian sauropods, with definition of a newly recognized clade.
evolution amongst sauropod dinosaurs. Proceedings of the Royal Society B: Anais da Academia Brasileira de Cie ^ncias 91 (2), e20180374. https://doi.org/
Biological Sciences 284, 20171219. 10.1590/0001-3765201920180374.
Carballido, J.L., Otero, A., Mannion, P.D., Salgado, L., Moreno, A.P., 2022. Titanosauria: Gomani, E.M., 2005. Sauropod dinosaurs from the Early Cretaceous of Malawi, Af-
a critical reappraisal of its systematics and the relevance of the South American rica. Palaeontologia Electronica 8, 1e37.
record. In: South American Sauropodomorph Dinosaurs. Springer, Cham, Haluza, A., Canale, J.I., Otero, A., Pe
rez, L.M., Scanferla, C.A., 2012. Changes in
pp. 269e298. vertebral laminae across the cervicodorsal transition of a well-preserved
Carvalho, I.D.S., Avilla, L.D.S., Salgado, L., 2003. Amazonsaurus maranhensis gen. et rebbachisaurid (Dinosauria, Sauropoda) from the Cenomanian of Patagonia,
sp. nov. (Sauropoda, Diplodocoidea) from the Lower Cretaceous (Aptian-Albian) Argentina. Journal of Vertebrate Paleontology 32, 219e224.
of Brazil. Cretaceous Research 24, 697e713. Hechenleitner, E.M., Leuzinger, L., Martinelli, A.G., Rocher, S., Fiorelli, L.E.,
Carrano, M.T., 2005. The evolution of Sauropod Locomotion: morphological di- Taborda, J.R.A., Salgado, L., 2020. Two Late Cretaceous sauropods reveal tita-
versity of a secondarily quadrupedal radiation. The Sauropods: Evolution and nosaurian dispersal across South America. Communications Biology. https://
Paleobiology 229e251. doi.org/10.1038/s42003-020-01338-w.
Cazau, L.B., Uliana, M.A., 1973. El Creta
cico superior continental de la Cuenca Holland, W.J., 1906. The osteology of Diplodocus Marsh. Memoirs of the Carnegie
Neuquina. In: 5 Congreso Geolo
gico Argentino, vol. 3. Actas, pp. 131e163. Museum 2, 225e226.
Cerda, I.A., 2009. Consideraciones sobre la histoge
nesis de las costillas cervicales en Ibiricu, L.M., Casal, G.A., Martínez, R.D., Lamanna, M.C., Luna, M., Salgado, L., 2013.
los dinosaurios sauro
podos. Ameghiniana 46, 193e198. Katepensaurus goicoecheai, gen. et sp. nov., a Late Cretaceous rebbachisaurid
Chure, D.J., Britt, B.B., Whitlock, J.A., Wilson, J.A., 2010. First complete sauropod (Sauropoda, Diplodocoidea) from Central Patagonia, Argentina. Journal of
dinosaur skull from the Cretaceous of the Americas and the evolution of Vertebrate Paleontology 33, 1351e1366.
sauropod dentition. Naturwissenschaften 97, 379e391. Ibiricu, L.M., Casal, G.A., Martínez, R.D., Alvarez, B.N., Poropat, S.F., 2020. New ma-
Clauss, M., Steuer, P., Müller, D.W.H., Codron, D., Hummel, J., 2013. Herbivory and terials and an overview of Cretaceous vertebrates from the Chubut Group of the
body size: allometries of diet quality and gastrointestinal physiology, and im- Golfo San Jorge Basin, central Patagonia, Argentina. Journal of South American
plications for herbivore ecology and dinosaur gigantism. In: 2. Sauropod Earth Sciences 98, 102460.
Gigantism, a Cross-disciplinary Approach e PLoS Collections One, vol. 8, Ibiricu, L.M., Martínez, R.D., Casal, G.A., 2012. The first record of Pterosauria in the
e68714. Bajo Barreal Formation (Upper Cretaceous), central Patagonia, Argentina.
Coria, R.A., Salgado, L., 2005. Mid-Cretaceous turnover of saurischian communities: Ameghiniana 49, 657e661.
evidence from the Neuque
n Basin. In: Veiga, G., Spalletti, L., Howell, J.A., Janensch, W., 1936. Die Scha €del der sauropoden Brachiosaurus, Barosaurus und
Schwartz, E. (Eds.), Neuque
n Basin: A Case Study in Sequence Stratigraphy and Dicraeosaurus aus den Tendaguruschichten Deutsch-Ostafrikas (Schluz). [The
Basin Dynamics, vol. 252. Special Publications of the Geological Society, heads of the sauropods Brachiosaurus, Barosaurus and Dicraeosaurus from the
pp. 317e327. German East African Tendaguru beds (conclusion). Paleontology Supplies 7 2,
Coria, R.A., Filippi, L.S., Chiappe, L.M., García, R.A., Arcucci, A.B., 2013. Overosaurus 249e298.
paradasorum gen. et sp. nov., a new sauropod dinosaur (Titanosauria: Lithos- Krause, J.M., Ramenazi, J., Umazano, A.M., Pol, D., Carballido, J.L., Sterli, J., Puerta, P.,
trotia) from the Late Cretaceous of Neuque
n, Patagonia, Argentina. Zootaxa Rube
n Cúnero, N., Bellosi, E.S., 2020. High-resolution chronostratigraphy of the
3683, 357e376. Cerro Barcino Formation (Patagonia): Paleobiologic implications for themid-
Cruzado-Caballero, P., Filippi, L.S., Me
ndez, A.H., Garrido, A.C., Díaz-Martínez, I., cretaceous dinosaurrich fauna of South America. Gondwana Research 80,
2018. First ornithopod remains from Bajo de la Carpa Formation (Santonian, 33e49.
Upper Cretaceous), northern Patagonia, Argentina. A new view about the Lavocat, R., 1954. Sur les dinosauriens du Continental Intercalaire des Kem-Kem de
biodiversity of Late Cretaceous South American ornithopods. Cretaceous la Daoura. In: 19 International Geological Congress, vol. 1, pp. 65e68.
Research 83, 182e193. Legarreta, L., Gulisano, C.A., 1989. An

fico secuencial de la Cuenca
alisis estratigra
Curry Rogers, K.A., Forster, C.A., 2004. The skull of Rapetosaurus krausei (Sauropoda: Neuquina (Tri
asico superior e Terciario inferior). In: Chebli, G., Spalletti, L.
Titanosauria) from the Late Cretaceous of Madagascar. Journal of Vertebrate 
(Eds.), Cuencas Sedimentarias Argentinas. 10 Congreso Geolo
gico Argentino,
Paleontology 24, 121e144. Serie Correlacio
n Geolo

gica, vol. 6, pp. 221e243. Buenos Aires.
D'emic, M.D., Carrano, M.T., 2020. Redescription of Brachiosaurid Sauropod Dino- Machado, E.B., Avilla, L.S., Nava, W.R., Campos, D.A., Kellner, A.W.A., 2013. A new
saur Material from the Upper Jurassic Morrison Formation, Colorado, USA. The titanosaur sauropod from the Late Cretaceous of Brazil. Zootaxa 3701 (3),
Anatomical Record 303, 732e758. 301e321.
Ezcurra, M.D., Agnolin, F.L., 2012. A new global palaeobiogeographical model for the Madsen, J.H.J., Mcintosh, J.S., Berman, D.S., 1995. Skull and atlas-axis complex of the
late Mesozoic and early Tertiary. 1. Systematic Biology 61, 553e566. Upper Jurassic sauropod Camarasaurus Cope (Reptilia: Saurischia). Bulletin of
Fanti, F., Cau, A., Hassine, M., Contessi, M., 2013. A new sauropod dinosaur from the Carnegie Museum of Natural History 31, 1e115.
Early Cretaceous of Tunisia with extreme avian-like pneumatization. Nature Mannion, P.D., Upchurch, P., Hutt, S., 2011. New rebbachisaurid (Dinosauria: Sau-
Communications 4, 2080. ropoda) material from the Wessex Formation (Barremian, Early Cretaceous), Isle
Filippi, L.S., García, R.A., Garrido, A., 2011. A new titanosaur sauropod dinosaur from of Wight, United Kingdom. Cretaceous Research 32, 774e780.
Upper Cretaceous of North Patagonia, Argentina. Acta Palaeontologica Polonica Mannion, P.D., Upchurch, P., Barnes, R.N., Mateus, O., 2013. Osteology of the Late
56, 505e520. Jurassic Portuguese sauropod dinosaur Lusotitan atalaiensis (Macronaria) and

13

rez Valieri, P.A. Gallina et al.
L.S. Filippi, R.D. Jua Cretaceous Research 154 (2024) 105754

the evolutionary history of basal titanosauriforms. Zoological Journal of the Royo-Torres, R., Upchurch, P., 2012. The cranial anatomy of the sauropod Turiasaurus
Linnean Society 168, 98e206. riodevensis and implications for its phylogenetic relationships. Journal of Sys-
Mannion, P.D., Allain, R., Moine, O., 2017. The earliest known titanosauriform tematic Palaeontology 10, 553e583.
sauropod dinosaur and the evolution of Brachiosauridae. PeerJ 5, e3217. Salgado, L., 2003. Los sauro
podos de Patagonia: sistema
tica, evolucio
n y paleo-
Mannion, P.D., Upchurch, P., Schwarz, D., Wings, O., 2019a. Taxonomic affinities of biología. In: Actas de las II Jornadas Internacionales sobre Paleontología de
the putative titanosaurs from the Late Jurassic Tendaguru Formation of Dinosaurios y su Entorno. Salas de Los Infantes, Espan ~ a, pp. 139e168.
Tanzania: phylogenetic and biogeographic implications for eusauropod dino- Salgado, L., De Souza Carvalho, I., 2008. Uberabatitan ribeiroi, a new titanosaur from
saur evolution. Zoological Journal of the Linnean Society 185, 784e909. the Marília Formation (Bauru Group, Upper Cretaceous), Minas Gerais, Brazil.
Mannion, P.D., Upchurch, P., Jin, X., Zheng, W., 2019b. New information on the Creta- Palaeontology 51 (4), 881e901.
ceous sauropod dinosaurs of Zhejiang Province, China: impact on Laurasian tita- Salgado, L., Apesteguía, S., Heredia, S.E., 2005. A new specimen of Neuquensaurus
nosauriform phylogeny and biogeography. Royal Society Open Science 6, 191057. australis, a Late Cretaceous Saltasaurinae titanosaur from North Patagonia.
Marpmann, J.S., Carballido, J.L., Sander, M.P., Kno € tschke, N., 2015. Cranial anatomy of Journal of Vertebrate Paleontology 25, 623e634.
the Late Jurassic dwarf sauropod Europasaurus holgeri (Dinosauria, Camar- Salgado, L., Gallina, P.A., Paulina Carabajal, A., 2014. Redescription of Bonatitan reigi
asauromorpha): ontogenetic changes and size dimorphism. Journal of Sys- (Sauropoda: Titanosauria), from the CampanianeMaastrichtian of the Río
tematic Palaeontology 13, 221e263. Negro Province (Argentina). Historical Biology 27, 525e548.
Martinelli, A.G., Forasiepi, A.M., 2004. Late Cretaceous vertebrates from Bajo de Sallam, H.M., Gorscak, E., O'connor, P.M., El-Dawoudi, I.A., El-Sayed, S., Saber, S.,
Santa Rosa (Allen Formation), Río Negro province, Argentina, with the Kora, M.A., Sertich, J.J.W., Seiffert, E.R., Lamanna, M.C., 2018. New Egyptian
description of a new sauropod dinosaur (Titanosauridae). In: Revista Museo sauropod reveals Late Cretaceous dinosaur dispersal between Europe and Af-
Argentino de Ciencias Naturales, Nueva Serie, vol. 6, pp. 257e305. rica. Nature Ecology & Evolution 2, 445e451.
Marsh, O.C., 1884. Principal characters of American Jurassic dinosaurs. Pt VII. On the Sander, P.M., 2013. An evolutionary cascade model for sauropod dinosaur
Diplodocidae, a new family of Sauropoda. American Journal of Science 27, 161e167. gigantism-overview, update and tests. PLoS One 8 (10), e78573.
Martínez, R.D.F., Lamanna, M.C., Novas, F.E., Ridgely, R.C., Casal, G.A., Mart, J.E., 2016. Sereno, P.C., Beck, A.L., Dutheil, D.B., Larsson, H.C.E., Lyon, G.H., Moussa, B.,
A Basal Lithostrotian Titanosaur (Dinosauria: Sauropoda) with a complete skull: Sadleir, R.W., Sidor, C.A., Varricchio, D.J., Wilson, G.P., Wilson, J.A., 1999. Creta-
implications for the evolution and paleobiology of Titanosauria. PLoS One 11 ceous sauropods from the Sahara and the uneven rate of skeletal evolution
(4), e0151661. https://doi.org/10.1371/journal.pone.0151661. among dinosaurs. Science 286, 1342e1347.
Mcintosh, J.S., Berman, D.S., 1975. Description of the palate and lower jaw of the Sereno, P.C., Wilson, J.A., Witmer, L.M., Whitlock, J.A., Maga, A., Ide
, O.A., Rowe, T.B.,
sauropod dinosaur Diplodocus (Reptilia: Saurischia) with remarks on the nature 2007. Structural extremes in a Cretaceous dinosaur. 1. PLoS One 2, e1230.
of the skull of Apatosaurus. Journal of Paleontology 49, 187e199. Silva Junior, J.C.G., Martinelli, A.G., Lori, F.V., Marinho, T.S., Hechenleitner, M.E.,
McLoughlin, S., 2001. The breakup history of Gondwana and its impact on pre- Langer, M.C., 2021. Reassessment of Aeolosaurus maximus, a titanosaur dinosaur
Cenozoic floristic provincialism. Australian Journal of Botany 49, 271e300. from the Late Cretaceous of southeastern Brazil. Historical Biology. https://
McPhee, B.W., Mannion, P.D., De Klerk, W.J., Choiniere, J.N., 2016. High diversity in doi.org/10.1080/08912963.2021.1920016.
the sauropod dinosaur fauna of the Lower Cretaceous Kirkwood Formation of Stevens, K.A., Parrish, J.M., 2005. Neck posture, dentition, and feeding strategies in
South Africa: Implications for the Jurassic-Cretaceous transition. Cretaceous Jurassic sauropod dinosaurs. In: Tidwell, V., Carpenter, K. (Eds.), Thunderlizards:
Research 59, 228e248. the Sauropodomorph Dinosaurs. Indiana University Press, Bloomington,
Moore, A.J., Mo, J., Clark, J.M., Xu, X., 2018. Cranial anatomy of Bellusaurus sui pp. 212e232.
(Dinosauria: Eusauropoda) from the Middle-Late Jurassic Shishugou Formation Torcida Ferna
ndez-Baldor, F., Canudo, J.I., Huerta, P., Montero, D., Pereda
of northwest China and a review of sauropod cranial ontogeny. PeerJ 6, e4881. Suberbiola, X., Salgado, L., 2011. Demandasaurus darwini, a new rebbachisaurid
Novas, F.E., 2009. The Age of Dinosaurs in South America. Indiana University Press, sauropod from the Early Cretaceous of the Iberian Peninsula. Acta Palae-
Indiana, pp. 1e536. ontologica Polonica 56, 535e552.
Novas, F.E., De Valais, S., Vickers-Rich, P., Rich, T., 2005. A large Cretaceous theropod Tschopp, E., Mateus, O., 2017. Osteology of Galeamopus pabsti sp. nov. (Sauropoda:
from Patagonia, Argentina, and the evolution of carcharodontosaurids. Natur- Diplodocidae), with implications for neurocentral closure timing, and the
wissenschaften 92, 226e230. cervico-dorsal transition in diplodocids. PeerJ 5, e3179.
Novas, F.E., Agnolín, F.L., Ezcurra, M.D., Porfiri, J., Canale, J.I., 2013. Evolution of the Tschopp, E., Mateus, O., Benson, R.B.J., 2015. A specimen-level phylogenetic analysis
carnivorous dinosaurs during the Cretaceous: the evidence from Patagonia. and taxonomic revision of Diplodocidae (Dinosauria, Sauropoda). PeerJ 3, e857.
Cretaceous Research 45, 174e215. https://doi.org/10.7717/peerj.857.
Ouyang, H., Ye, Y., 2002. The First Mamenchisaurian Skeleton with Complete Skull Whitlock, J.A., 2011. Inferences of diplodocoid (Sauropoda: Dinosauria) feeding
Mamenchisaurus youngi. Sichuan Science and Technology Press, Chengdu. behavior from snout shape and microwear analyses. PLoS One 6, e18304.

rez Moreno, A., Carballido, J.L., Otero, A., Salgado, L., Calvo, J.O., 2022. The axial
Pe Whitlock, J.A., Wilson, J.A., Lamanna, M.C., 2010. Description of a nearly complete
skeleton of Rinconsaurus caudamirus (Sauropoda: Titanosauria) from the Late juvenile skull of Diplodocus (Sauropoda: Diplodocoidea) from the Late Jurassic
Cretaceous of Patagonia, Argentina. Ameghiniana 59 (1), 1e46. of North America. Journal of Vertebrate Paleontology 30, 442e457.
Perry, S.F., Christian, A., Breuer, T., Pajor, N., Codd, J.R., 2009. Implications of an Wilson, J.A., 2005. Redescription of the Mongolian sauropod Nemegtosaurus mon-
avian-style respiratory system for gigantism in sauropod dinosaurs. Journal of goliensis Nowin
ski (Dinosauria: Saurischia) and comments on Late Cretaceous
Experimental Zoology Part A 311, 1e11. sauropod diversity. Journal of Systematic Palaeontology 3, 283e318.
Poropat, S.F., Kear, B.P., 2013. Photographic atlas and three-dimensional recon- Wilson, J.A., Pol, D., Carvalho, A.B., Zaher, H., 2016. The skull of the titanosaur
struction of the holotype skull of Euhelopus zdanskyi with description of addi- Tapuiasaurus macedoi (Dinosauria: Sauropoda), a basal titanosaur from the
tional cranial elements. PLoS One 8, e79932. Lower Cretaceous of Brazil. Zoological Journal of the Linnean Society 178 (3),
Poropat, S.F., Kundra
t, M., Mannion, P.D., Upchurch, P., Tischler, T.T., Elliott, D.A., 611e662. https://doi.org/10.1111/zoj.12420.
2021. Second specimen of the Late Cretaceous Australian sauropod dinosaur Woodruff, D.C., Carr, T.D., Storrs, G.W., Waskow, K., Scannella, J.B., Norde
n, K.K.,
Diamantinasaurus matildae provides new anatomical information on the skull Wilson, J.P., 2018. The smallest diplodocid skull reveals cranial ontogeny and
and neck of early titanosaurs. Zoological Journal of the Linnean Society 192, growth-related dietary changes in the largest dinosaurs. Scientific Reports 8,14341.
610e674. Xu, X., Upchurch, P., Mannion, P.D., Barrett, P.M., Regalado-Fernandez, O.R., Mo, J.,
Poropat, S.F., Mannion, P.D., Rigby, S.L., Duncan, R.J., Pentland, A.H., Bevitt, J.J., Ma, J., Liu, H., 2018. A new Middle Jurassic diplodocoid suggests an earlier
Sloan, T., Elliott, D.A., 2023. A nearly complete skull of the sauropod dinosaur dispersal and diversification of sauropod dinosaurs. Nature Communications.
Diamantinasaurus matildae from the Upper Cretaceous Winton Formation of https://doi.org/10.1038/s41467-018-05128-1.
Australia and implications for the early evolution of titanosaurs. Royal Society Zaher, H., Pol, D., Carvalho, A.B., Nascimento, P.M., Roccomini, C., Larson, P., Jua
rez-
Open Science 10, 221618. https://doi.org/10.1098/rsos.221618. Valieri, R.D., Pires-Domingues, R., Da Silva, N.J., Campos, D.A., 2011. A complete
Powell, J.E., 1992. Osteología de Saltasaurus loricatus (SauropodaeTitanoesauridae) skull of an Early Cretaceous sauropod and the evolution of advanced titano-
del Creta
cico Superior del Noroeste Argentino. In: Sanz, J.L., Buscalioni, J.L. saurians. PLoS One 6, e16663.
(Eds.), Los Dinosaurios y su entorno bio
tico, Actas del Segundo Curso de Pale-
ontología de Cuencas. Instituto “Juan De Valde
s”, Cuencas, pp. 165e230.
Powell, J.E., 2003. Revision of South American Titanosaurid Dinosaurs: Palae-
obiological, Palaeobiogeographical and Phylogenetic Aspects. Recors of the
Queen Victoria Museum, Launceston, p. 173pp. Appendix A. Supplementary data
Rauhut, O.W.M., Carballido, J.L., Pol, D., 2015. A diplodocid sauropod dinosaur from
the Late Jurassic Can ~ ado

n Calc

areo Formation of Chubut, Argentina. Journal of Supplementary data to this article can be found online at https://doi.org/10.
Vertebrate Paleontology 35, e982798. 1016/j.cretres.2023.105754.

14