Cretaceous Research 128 (2021) 104957

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

Osteology, paleohistology and phylogenetic relationships of

Pellegrinisaurus powelli (Dinosauria: Sauropoda) from the Upper
Cretaceous of Argentinean Patagonia
Ignacio Cerda a, b, Virginia Laura Zurriaguz a, *, Jose

Luis Carballido c, Romina Gonza

lez d,
a
Leonardo Salgado
a
CONICET-Instituto de Investigaciones en Paleobiología y Geología, Universidad Nacional de Río Negro, Avenida Roca 1242, General Roca 8332, Río Negro,
Argentina
b
CONICET-Museo Carlos Ameghino, Belgrano 1700, Paraje Pichi Ruca (predio Marabunta), Cipolletti R8324, Río Negro, Argentina
c
CONICET, Museo Paleontolo
gico Egidio Feruglio, Trelew 9100, Chubut, Argentina
d
CONICET-UNNE, Centro de Ecología Aplicada del Litoral. Laboratorio de Paleontología de Vertebrados. Ruta 5 Km 2,5, 3400, Corrientes, Argentina

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

Article history: Pellegrinisaurus powelli is a large titanosaurian sauropod from the Upper Cretaceous of northern Pata-
Received 12 January 2021 gonia (Río Negro Province, Argentina). The holotype of this species comprises an incomplete femur, four
Received in revised form dorsal and 26 caudal vertebrae collected from the Lago Pellegrini locality. A single middle caudal vertebra
9 June 2021
from the nearby Cinco Saltos locality has been more recently referred to Pellegrinisaurus. With the
Accepted in revised form 11 July 2021
purpose of increasing our knowledge about the titanosaur faunas of the Upper Cretaceous of Patagonia,
Available online 21 July 2021
here we provide a detailed redescription of the holotype and referred specimens of Pellegrinisaurus,
reviewing its diagnosis and phylogenetic position. A histological analysis of the femur of the holotype is
Keywords:
Titanosauria
also carried out. The diagnostic characters originally proposed for Pellegrinisaurus are here considered
Lithostrotia invalid because they either correspond with diagenetic deformations of the specimen or are present in
Pellegrinisaurus powelli other titanosaurs. The new diagnostic features are related with the presence of large lateral blind fossae
Upper Cretaceous with distinct dorsal rims in the anterior caudal centra and with a low longitudinal ridge on the posterior
Patagonia side of the femur shaft originated from the trochanteric shelf and extending to the distal third of the
Paleohistology shaft. The phylogenetic analysis recovers Pellegrinisaurus as a non-saltasaurid lithostrotian, closely
related with Alamosaurus sanjuanensis. The bone histology indicates that the holotype corresponds to a
subadult (i.e. sexually mature but still growing) individual. Finally, the caudal vertebra previously
attributed to Pellegrinisaurus cannot be referred to this genus and is here assigned to Titanosauria indet.
© 2021 Elsevier Ltd. All rights reserved.

1. Introduction femur, the individual actually preserved many more elements

Pellegrinisaurus powelli is a large titanosaurian sauropod from
the Upper Cretaceous of northern Patagonia (Río Negro Province,
Argentina). The remains of the holotype specimen were discovered
in a sand quarry located at the south margin of the Pellegrini Lake
and recovered by the crew of the Museo Carlos Ameghino of
Cipolletti (Río Negro Province) in 1975 (Salgado 1996; Powell
2003). Although the original description of Salgado (1996) was
based on four dorsal centra, 26 caudal vertebrae and an incomplete
* Corresponding author.
E-mail addresses: nachocerda6@gmail.com (I. Cerda), vzurriaguz@unrn.edu.ar
(V.L. Zurriaguz), carballidojl@gmail.com (J.L. Carballido), romigonzl95@gmail.com
(R. Gonz
alez), lsalgado@unrn.edu.ar (L. Salgado).
when it was discovered (Fig. 1), including several complete dorsal
and sacral vertebrae, which were found in articulation (Powell
2003). Pellegrinisaurus powelli was initially assigned to Epachtho-
saurus sp. on the basis of dorsal vertebrae characters (Powell 1986).
Later, Salgado (1996) assigned its remains to a new genus and
species, proposing autapomorphies based on the relative pro-
portions of the dorsal centra and in the morphology of the neural
arches of the mid-posterior and posterior caudal vertebrae. More
recently, Salgado and Bonaparte (2007) assigned an isolated middle
caudal vertebra from the Cinco Saltos locality (Río Negro Province)
to Pellegrinisaurus. Illustrations of this specimen were however not
provided, as well as the anatomical features supporting its tax-
onomical assignment.
Regarding its phylogenetic affinities, Salgado (1996) considered
Pellegrinisaurus powelli as the probable sister taxon of Saltasaurinae

https://doi.org/10.1016/j.cretres.2021.104957
0195-6671/© 2021 Elsevier Ltd. All rights reserved.
I. Cerda, V.L. Zurriaguz, J.L. Carballido et al.

nica” sand quarry, located in the South margin of the Pellegrini Lake (Río Negro Province, Argentina). The
Fig. 1. Excavation of the Pellegrinisaurus powelli holotype in the “Pala Meca
picture includes dorsal and sacral vertebrae in articulation, which are oriented upside down. Picture courtesy of Jaime Powell (R.I.P).
(¼Saltasaurini sensu Salgado and Bonaparte 2007), but it was not
included in a phylogenetic analysis to support this hypothesis. The
first contribution that included Pellegrinisaurus in a phylogenetic
analysis corresponds to Upchurch et al. (2004). There, this species
was recovered as the sister taxon of Alamosaurus sanjuanensis, with
this clade recovered as the sister group of Saltasauridae. More
recently, Gorscak and O'Connor (2016, 2019) recovered Pelle-
grinisaurus as the sister group of the clade Alamosaurus
sanjuanensis þ Baurutitan britoi. In this data set Pellegrinisaurus is
recovered as a Saltasauridae, with this position the result of an
unusual position of the specifier taxa of this clade (Saltasaurus
loricatus and Opisthocoelicaudia skarzynski). As such, Saltasauridae,
in that study, includes several taxa that are usually excluded from
this clade (e.g., Lognkosauria and basal titanosaurs such as Epach-

lez Riga et al., 2018;
thosaurus sciuttoi; Carballido et al., 2017; Gonza
Mannion et al., 2019). The matrix of Gorscak and O'Connor (2016)
was used by Sallam et al. (2018) to perform a phylogenetic anal-
ysis based on both Bayesian and parsimony approaches. Bayesian
analysis recovered Pellegrinisaurus as the sister group of a clade
formed by Dreadnoughtus schrani þ (Alamosaurus
sanjuanensis þ Baurutitan britoi), in a similar topology to that
recovered by Gorscak and O'Connor (2016, 2019). On the other
hand, the parsimony analysis recovered Pellegrinisaurus in a slightly
more basal position, being the sister group of Ampelosaurus atacis þ
(Paludititan nalatzensis þ (Dreadnoughtus schrani þ (Alamosaurus
sanjuanensis þ Baurutitan britoi))). A basal position of Pelle-
grinisaurus amongst titanosaurs was recovered by Bandeira et al.
(2016). In sum, most analyses coincide in depicting Pelle-
grinisaurus as a non-Saltasauridae lithostrotian, except that of
Gorscak and O'Connor (2016) and data sets derived from it, in
which the “Saltasauridae relationships” of Pellegrinisaurus are
caused by the unusual distant position of Opisthocoelicaudia and
Saltasaurus, as was mentioned above.
In this paper, we provide a comprehensive redescription of the
holotype material Pellegrinisaurus powelli. Our main goals are: to
2
Cretaceous Research 128 (2021) 104957
expand the anatomical description of Pellegrinisaurus powelli, to
revise its current diagnosis, to describe and discuss the assignation
of the specimen mentioned by Salgado and Bonaparte (2007), to
determine the ontogenetic stage of the holotype specimen using
bone histology, and to revise the phylogenetic position of this
taxon. With regard to this last objective, we test previous hypoth-
esis of the phylogenetic affinities of Pellegrinisaurus. This study is
the first that test the phylogenetic position of Pellegrinisaurus based
on first hand study of the material.
Institutional abbreviations: MCS-Pv, Museo de Cinco Saltos,

n de paleontología de vertebrados, Río Negro, Argentina;
coleccio
MPCA-Pv, Paleovertebrate collection of the Museo Provincial ‘Carlos
Ameghino’, Cipolletti, Río Negro, Argentina.
2. Materials and methods
2.1. Materials
The materials analyzed comprise the holotype of Pelle-
grinisaurus powelli (MPCA-Pv 1500), which consists of four
incomplete dorsal vertebrae (MPCA-Pv 1500/1-4), 26 fragmentary
caudal vertebrae (MPCA-Pv 1500/5-30) and an incomplete left fe-
mur (MPCA-Pv 1500/6-31). Unfortunately, other discovered parts of
the skeleton (e.g. sacrum, dorsal ribs, neural arches of the dorsal
vertebrae) were badly ruined in the field during the excavation
procedures. Different from the original description of Salgado
(1996), we realized that two anterior caudal vertebrae (the first
and the fourth or the fifth) described in the original paper are
missing from the MPCA collection. Unexpectedly, despite the
absence of these two elements, the number of caudal vertebrae is
still the same as that reported by Salgado (1996). Since all the
reviewed caudal vertebrae share the same features, including de-
gree of preservation, labeling and mode of restoration, we consider
that all belong to the same individual. Therefore, we infer that the
original number of preserved caudal vertebrae was, at least, 28
I. Cerda, V.L. Zurriaguz, J.L. Carballido et al.
bones. It is worth to note that, in his descriptions, Powell (1986,
2003) mentioned that “a series of 27 specimens [caudal vertebrae]
were preserved in succession almost without interruption”. Besides
the Pellegrinisaurus powelli holotype specimen, we describe a
middle caudal vertebra (MCS-Pv 53) assigned to this taxon by
Salgado and Bonaparte (2007).
2.2. Geographical and stratigraphic provenance
The holotype specimen of Pellegrinisaurus powelli was recovered
from the “Pala Meca
nica” commercial sand quarry, located at the
south margin of the Pellegrini Lake, an artificial reservoir situated
around 10 km North to the city of Cinco Saltos, Río Negro Province
(Fig. 2). All the Pellegrini Lake sand quarries have been originally
considered as outcrops of the lower member of the Allen Formation
(Bonaparte and Novas 1985; Powell 1986, 2003; Bonaparte 1991).
The Allen Formation corresponds with the base of the Malargüe
Group and is currently considered upper Campanianelower
Maastrichtian in age (Rodríguez et al., 2007). The correspondence
of the Pellegrini Lake sand quarries with the Allen Formation was
subsequently discussed by Heredia and Salgado (1999), who pro-
posed that these outcrops actually correspond to the Anacleto
Formation (Río Colorado Formation in the original publication). The
Anacleto Formation, which also is exposed in the Cinco Saltos lo-
cality, corresponds with the top of the Neuque
n Group and has
been assigned to the lower Campanian (Dingus et al., 2000).
Although this last geological unit assignment for the Pellegrini Lake
sand quarries has been followed by several authors (e.g. Salgado
Fig. 2. Map of location and geological outcrops of the region of Pellegrini Lake and Cinco Saltos (Río Negro Province, Argentina), where MPCA-Pv 1500 (red arrowhead) and MCS-Pv
53 (white arrowhead) were collected. (For interpretation of the references to colour in this figure legend, the reader is referred to the Web version of this article.)
3
Cretaceous Research 128 (2021) 104957
2003; Coria 2007; Rodríguez et al., 2007; Salgado and Bonaparte
2007; Garrido 2010; Cerda et al., 2015; Otero and Salgado 2015;
García et al., 2017), Delaloye and Garrido (2017) have again
considered the original correspondence of these outcrops to the
Allen Formation. Given that this last change of stratigraphic
assessment has not been to date formally proposed in a publication,
we follow the more broadly accepted proposal of Heredia and
Salgado (1999), i.e. that the Pellegrinisaurus type specimen came
from the Anacleto Formation.
In the case of the specimen MCS-Pv 53, this was recovered from
outcrops exposed in the northwestern scarps near Cinco Saltos city,
adjacent to the former Indupa chemical factory (now Imextrade
S.A) (Fig. 2). These outcrops correspond to the top of the Anacleto
Formation, from which other non-avian dinosaur remains have
been recovered (Coria and Salgado 1996; Salgado et al., 1997a,
2005).
2.3. Methods
For the anatomical description we use “Romerian” terms (Wilson
2006) for the structures (e.g. “centrum”, not “corpus”) and their
orientation and relative position (e.g. “anterior”, not “cranial”).
Nomenclature of neural laminae and fossae follows Wilson (1999)
and Wilson et al. (2011a) respectively. For the pneumatic struc-
tures we use the terminology proposed by Wedel (2003).
For histological analysis, the femur of the holotype (MPCA-Pv
1500/31) was sampled following a modification of methodology
proposed by Sander (2000). In this regard, we extracted two cores,
I. Cerda, V.L. Zurriaguz, J.L. Carballido et al.
one from the anterior and the posterior sides, using a diamond
studded drill bit mounted on a domestic power drill. We used
30 mm diameter bits with their ends covered with diamond grit.
Thin sections from each core were obtained in the Paleohistological
laboratory of the Museo Provincial Carlos Ameghino (Cipolletti, Río
Negro Province, Argentina) and the methodology followed the
procedures of Cerda et al. (2020). The thin sections were studied
under transmitted light, both normal and polarized with a micro-
scope Zeiss Axioskop 40 with an attached Canon (PowerShot A640)
digital camera. The terminology follows Francillon-Vieillot et al.
(1990).
3. Systematic paleontology
Dinosauria Owen 1842
Saurischia Seeley 1887
Sauropoda Marsh, 1878
Titanosauria Bonaparte and Coria 1993
Lithostrotia Upchurch et al., 2004
Pellegrinisaurus powelli Salgado 1996
Emended diagnosis. Lithostrotian titanosaurian characterized by
the following combination of characteristics (autapomorphies
highlighted with asterisks): presence of a prominent blind lateral
fossa (“pleurocoel”) with well-developed dorsal rim in the anterior
caudal centra*; femur with longitudinal ridge located on the
posterior surface of the shaft and laterally displaced, originated
from the trochanteric shelf and extending to the distal third of the
shaft.
Locality and horizon. South margin of the Pellegrini Lake, around
10 km North to the city of Cinco Saltos, Anacleto Formation (lower
Campanian, Dingus et al., 2000).
4. Anatomical description
4.1. Dorsal vertebrae (MPCA-Pv 1500/1-4)
Four incomplete dorsal vertebrae centra are preserved (Fig. 3).
There is not current information regarding to their relative position
in the column. Also, it is not possible to determine if they are
continuous elements. Salgado (1996) assigned these elements to
posterior dorsal vertebrae based on the absence of parapophyses.
For descriptive purpose, we assign tentatively these elements to
seventh (MPCA-Pv 1500/1), eighth (MPCA-Pv 1500/2), ninth
(MPCA-Pv 1500/3) and tenth (MPCA-Pv 1500/4) dorsal vertebrae
(Supplementary Table 1). The sequential order is based on the
centrum length, which decreases from the seventh to the tenth
dorsal vertebrae in other titanosaurs (e.g. Powell 2003; Coria et al.,
2013). Different degrees of anatomical deformation are evident in
the four elements, probably from post-depositional stress. Also,
several portions of the elements have been broken away and, in
several instances, covered with plaster, which makes a compre-
hensive morphological description difficult. Since the four centra
share several anatomical features, we describe them together,
indicating differences when applicable.
All the centra are opisthocoelous. The condyle is more promi-
nent in MPCA-Pv 1500/2 than in the other elements. A poorly
developed anterior rim bounds the anterior condyle from the main
body of the centrum in MPCA-Pv 1500/3 and MPCA-Pv 1500/4. The
relationship between the width and the height of the centrum is
not the same in all the centra, mainly due to plastic deformation. In
this regard, the width is not much greater than the height in the
least altered centrum (MPCA-Pv 1500/1). On the other hand, the
centra of MPCA-Pv 1500/2 and MPCA-Pv 1500/4 are clearly wider
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Cretaceous Research 128 (2021) 104957
than high, particularly in the last one. However, this feature seems
to be related to a very substantial dorsoventral compressional
deformation suffered by the centra, which is evidenced by promi-
nent displaced fractures on the anterior and posterior sides of each
element. In the case of the specimen MPCA-Pv 1500/3, it presents a
different type of deformation, in which the centrum is distorted
with regard to the sagittal plane. Such distortion results in an
inclination of around 22 degrees from the sagittal plane, leaving the
floor of the neural canal displaced to the left side of the centrum
and giving a roughly ‘parallelepiped shape’ in anterior and poste-
rior views. Other gross anatomical features of the centra (e.g.
relationship between anterior and posterior height, condylar
shape) are difficult to establish with confidence due to the
noticeable degree of deformation of these bones. The ventral sur-
faces lack any ridge and they are both transversely and ante-
roposteriorly flat or slightly concave (e.g. MPCA-Pv 1500/1). All the
elements exhibit large, “eye shaped” lateral pneumatic foramina,
which occupy half or more of the total length of the centra. The
lateral pneumatic foramen is posteriorly displaced in MPCA-Pv
1500/4. The lateral pneumatic foramina located on the right side
of the four centra are larger than those on the left side. Such vari-
ation, however, can be due to taphonomic deformation. Dividing
laminae are absent inside the lateral pneumatic foramen. Natural
breakages reveal a pneumatic internal structure, which corre-
sponds with camellate tissue.
4.2. Caudal vertebrae (MPCA-Pv 1500/5-30)
Twenty six caudal vertebrae are preserved (Fig. 4A-U). As pre-
viously mentioned, these do not strictly correspond to the twenty
six elements mentioned by Salgado (1996), who included the first
caudal and other anterior vertebrae, possibly the fourth or the fifth.
As with the dorsal centra, the caudal vertebrae are mostly badly
preserved, showing in some cases substantial plastic deformation.
Neural arches are usually incomplete or missing. The centra are
broken in several parts and have been reconstructed with abundant
plaster (this is particularly evident in the element MPCA-Pv 1500/
29, in which plaster covers almost the entire bone).
In general terms, with the exception of the biconvex first caudal
vertebrae figured by Salgado (1996), all caudal vertebral centra are
procoelous. The degree of procoely decreases gradually from the
anterior to the posterior caudal vertebrae. The position of the
condylar apex migrates from the dorsal (i.e. above the horizontal
midline of the centrum) to the middle of the posterior articular
surface area. The length of the centra in relation with its width and
height increases toward the posterior caudals. The neural arches
are located above the anterior halves of the centra. The neural
spines are anteroposteriorly short. Their height decreases posteri-
orly in the caudal sequence. The inclination of the prezygapophyses
in relation to the centrum becomes less pronounced toward the
posterior caudal vertebra, where they become parallel. The pre-
zygapophyses project beyond the anterior margin of the centrum.
The postzygapophyses are mostly not preserved or are covered
with the prezygapophyses of the following vertebra. Fracture sur-
faces reveal the absence of internal pneumatic structure both in
neural arches and centra. In this regard, the bone tissue varies
between compact and cancellous depending on the vertebral areas.
Along with these general features, the caudal vertebrae exhibit
particular anatomical characteristics, which are described below.
Following previous works (e.g., Mannion et al., 2013; Carballido and
Sander 2014) the classification of the caudal vertebrae in anterior,
middle and posterior elements was based on anatomical features,
as for example the presence of transverse processes, the develop-
ment of neural spines, pre- and postzygapophysis, and the relative
length of their centra. Additionally, and given the incomplete
I. Cerda, V.L. Zurriaguz, J.L. Carballido et al.
Fig. 3. Dorsal vertebrae of Pellegrinisaurus powelli MPCA 1500. A, MPCA-Pv 1500/1. B, MPCA-Pv 1500/2. C, MPCA-Pv 1500/3. D, MPCA-Pv 1500/4. The vertebrae are showed in right
lateral (1), anterior (2) and posterior (3) views. Dashed white lines in C.3 indicate the position of the neural canal floor. Scale bars equal to 10 cm. Abbreviations: lpf: lateral
pneumatic foramen.
preservation of these structures in MPCA-Pv 1500 we also followed
other more complete sequence known for other related titanosaurs
(e.g., Baurutitan britoi; Kellner et al., 2005). For the present
description and analysis, the caudal sequence of Pellegrinisaurus
was divided in three sections: anterior (caudals 1-6), middle
(caudals 7-16), and posterior (caudals 17-28) caudal vertebrae.
4.2.1. Anterior caudal vertebrae (MPCA-Pv 1500/5-8)
The three anteriormost preserved caudal vertebrae are strongly
procoelous, and they exhibit anteroposteriorly short centra
(Figs. 4A-C, 5A-C). The ventral and lateral faces of the centra are
slightly concave. While the centrum height appears to be greater
than the width in MPCA-Pv 1500/5-6 (these elements are incom-
plete), MPCA-Pv 1500/7 is slightly wider than high. Nevertheless,
this difference seems to be related to a deformation by lateral
compression in the first two elements. Excluding the articular
condyle, the centrum is shorter than high. The posterior condyle is
prominent, occupying 28e34% of the entire length of the centrum
in MPCA-Pv 1500/5-6 (it is broken in MPCA-Pv 1500/7). The pos-
terior condyle is delimited from the main body of the centrum by a
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Cretaceous Research 128 (2021) 104957
distinct border. The ventral portion of the condyle, which is only
well preserved in MPCA-Pv 1500/5, exhibits a concave outline in
lateral view. A well-developed blind fossa is present on the lateral
sides of the centra (Figs. 4A-C, 5B). They are anteroposteriorly
elongate (almost twice longer than high) and located in the middle
of the centra. Since the right side of these vertebrae has been
eroded, the following dimensions (length x height) of the lateral
blind fossae correspond to the left side: 5.4  3 cm (MPCA-Pv 1500/
5), 5.9  3.2 cm (MPCA-Pv 1500/6) and 5.6  1.9 cm (MPCA-Pv
1500/7). The ventral faces of the centra lacks any ridge and are
slightly anteroposteriorly and transversely concave. The dorsal
margin of the lateral blind fossae of MPCA-Pv 1500/5 and MPCA-Pv
1500/6 exhibits a distinct rim. The same is also present in MPCA-Pv
1500/7, but it is much less pronounced than in the other centra.
Distinct neurovascular foramina open below the lateral blind fossae
and also on the ventral side of the centra. These are large (reaching
20 mm diameter), anteroposteriorly enlarged and irregularly
distributed. Transverse processes are mostly broken. Part of the
base of the left transverse process is preserved in MPCA-Pv 1500/6,
which is placed along the dorsal margin of the centrum. It is robust
I. Cerda, V.L. Zurriaguz, J.L. Carballido et al. Cretaceous Research 128 (2021) 104957

Fig. 4. Caudal vertebrae of Pellegrinisaurus powelli MPCA-Pv 1500. The sequence include anterior (AeD), middle (EeN) and posterior (OeU) elements in lateral left side. A, MPCA-Pv
1500/5. B, MPCA-Pv 1500/6. C, MPCA-Pv 1500/7. D, MPCA-Pv 1500/8. E, MPCA-Pv 1500/9. F, MPCA-Pv 1500/10. G, MPCA-Pv 1500/11. H, MPCA-Pv 1500/12. I, MPCA-Pv 1500/13. J,
MPCA-Pv 1500/14. K, MPCA-Pv 1500/15. L, MPCA-Pv 1500/16. M, MPCA-Pv 1500/17. N, MPCA-Pv 1500/18. O, MPCA-Pv 1500/19 and 20. P, MPCA-Pv 1500/21 and 22. Q, MPCA-Pv
1500/23 and 24. R, MPCA-Pv 1500/25. S, MPCA-Pv 1500/26. T, MPCA-Pv 1500/27 and 28. U, MPCA-Pv 1500/30. For comparative purpose we have inverted the image showed in I,
which actually corresponds with the right side. Scale bars equal to 10 cm. Abbreviations: bf: lateral blind fossa; dt: dorsal tuberosity; nvf: neurovascular foramen; pvp: poster-
oventral protuberance.

and directed laterally and slightly posteriorly. There is a distinct
lateral prominence at the level of the dorsal margin line of the
neural canal (Fig. 5A-C), which resembles the “dorsal tuberosity”
described for other titanosaurians (e.g. Bonitasaura salgadoi, Baur-
utitan britoi; Kellner et al., 2005; Gallina and Apesteguía 2015). The
bases of the neural arches are partially preserved and they occupy
most of the anteroposterior length of the centrum (discounting the
6
hemispherical posterior condyle). The neural canals are roughly
higher than wide, and their shape becomes wider ventrally.
The specimen MPCA-Pv 1500/8 (Fig. 4D), which has been inter-
preted as the sixth caudal vertebra, was originally described and
figured by Salgado (1996, figure 4). However, some important aspects
of that original description need to be reinterpreted. The element
preserves the centrum and part of the neural arch, including the base
I. Cerda, V.L. Zurriaguz, J.L. Carballido et al.
Fig. 5. Anterior caudal vertebra of Pellegrinisaurus powelli (MPCA-Pv 1500/6) in anterior (A), left lateral (B) and posterior views. Striped areas indicate broken and/or covered with
plaster surfaces. Scale bars equals to 10 cm. Abbreviations: bf: lateral blind fossa; bfr: dorsal rim of the blind fossa; dt: dorsal tuberosity; nc: neural canal; nvf: neurovascular
foramen; tp: transverse process.
of the left prezygapophysis. The centrum has a less pronounced
posterior condyle compared to the most anterior centra. The condylar
apex of the posterior surface is dorsally displaced. In its original
description, Salgado (1996) indicated that the lateral sides of the
centrum are downward oriented, leaving a narrow ventral surface.
However, the centrum is strongly deformed in a similar way (but
much more pronounced) that the previously described in the dorsal
vertebra MPCA-Pv 1500/3 (i.e. oblique distortion of the element with
regard to the sagittal plane). Such deformation causes the right and
left lateral sides to be exposed in dorsal and ventral views respec-
tively. Hence, the “downward oriented” right lateral side actually
corresponds to the ventral surface and the “very narrow ventral
surface” is therefore the boundary between the left lateral and ventral
sides. The original shape of the centrum seems to be roughly
quadrangular. Although an incomplete left postzygapophysis seems
to be preserved, its identification is dubious because of the strong
deformation suffered by the specimen.
4.2.2. Middle caudal vertebrae (MPCA-Pv 1500/9-18)
From the anterior middle caudals (i.e. MPCA-Pv 1500/9-14,
Fig. 4E-J), which exhibit similar anatomical features, the best pre-
served element corresponds to specimen MPCA-Pv 1500/9, which
has been interpreted as the seventh caudal vertebra (Figs. 4E, 6).
The centra of the anterior middle caudals are slightly wider than
high, and exhibit a rather quadrangular shape in posterior view.
Excluding the articular condyle, the centra are slightly longer than
high. The posterior condyle is less developed than in the anterior
caudals, occupying less than 16% of the entire length centrum in the
best preserved elements. The apex of the posterior condyles is
located on the dorsal half of the articular surface. The lateral sides
are almost straight, just gently excavated. A small but distinct
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Cretaceous Research 128 (2021) 104957
protuberance is placed in the posteroventral area of the lateral side
of MPCA-Pv 1500/9 centrum (Fig. 4E). This structure is not observed
in other centra. Such absence, nevertheless, may be related to the
general poor preservation of these elements. The distinct blind
lateral fossae described in the anterior caudals are absent in these
vertebrae. Large neurovascular foramina, however, are distinct in at
least the lateral surfaces of the centrum MPCA-Pv 1500/10. The
ventral surfaces exhibit a shallow depression, which is delimited
laterally by slightly pronounced ridges oriented anteroposterioly.
Chevron facets are mostly not preserved. Nevertheless, they appear
to be poorly developed. Only the base of the left transverse process
is preserved in one element (MPCA-Pv 1500/9). The same, however,
does not appear to be well developed. A weakly pronounced but
distinct “dorsal tuberosity” is observed in MPCA-Pv 1500/9, at the
level of the neural canal (Fig. 6A-C). Possible traces of this structure
are recorded in the same position in specimens MPCA-Pv 1500/10-
12. As in the anterior caudal vertebrae, the bases of the neural
arches occupy most of the anteroposterior length of centrum
(discounting the posterior condyle). Almost complete neural spines
are only preserved in elements MPCA-Pv 1500/9 and MPCA-Pv
1500/14. The neural arch of MPCA-Pv 1500/9 is relatively
high, being approximately half of the complete vertebral height
(Fig. 6A-C). The neural spine of this vertebra is laterally compressed,
with their anterior and posterior edges placed at the same level (i.e.
the anterior edge of the neural spine tip is not located above or
below the level of the posterior edge of the neural spine tip, Fig. 6B).
The dorsal margin of the spine is slightly convex in lateral view. In
this vertebra, the long axis of the spine (i.e. spine main axis) is
perpendicular to the anteroposterior axis of the centrum. The
posterior margin of the spine exhibits at distinct wrinkled border,
possibly related to the presence of an interspinous ligament. The
I. Cerda, V.L. Zurriaguz, J.L. Carballido et al.
neural spine of specimen MPCA-Pv 1500/14 (possibly the 12th) is
less preserved than in MPCA-Pv 1500/9. Its main difference is in the
orientation, which in posteriorly inclined, forming an angle of 40
degrees with the main axis of the centrum (Fig. 4J). Another dif-
ference with the neural spine of MPCA-Pv 1500/9 is in the relative
position of the posterior edge, which is lower with respect to the
anterior (i.e. the overall dorsal margin slopes to face poster-
odorsally). Complete prezygapophyses are preserved only in the
specimen MPCA-Pv 1500/9. These are strongly elongated, occu-
pying almost half of the complete vertebral length. They are in-
clined forming a low angle (of approximately 15 degrees) with the
centrum long axis. Such inclination appears to be maintained in the
specimens MPCA-Pv 1500/10-14. The articular facets of the pre-
zygapophyses are elongated and medially oriented. The post-
zygapophyses are located above the midpoint of the centrum
(excluding the posterior condyle). While the left postzygapophysis
is broken, the right one is covered with the anterior end of the
prezygapophysis of the following vertebra.
Posterior middle caudals (MPCA-Pv 1500/15-18) exhibit more
elongated centra in comparison with the preceding middle ele-
ments (Fig. 4K-N). The width and height of the centra is roughly the
same in these elements, which exhibit a slightly hexagonal shape in
anterior view. The centrum length is twice longer than high. A
shallow dorsal concavity is present on the posterior portion of the
Fig. 6. Anterior middle caudal vertebra of Pellegrinisaurus powelli (MPCA-Pv 1500/9) in anterior (A), right lateral (B) and posterior views. A fragment of the prezygapophysis of the
following vertebra is marked with an asterisk. Striped areas indicate broken and/or covered with plaster surfaces. Scale bars equal to 10 cm. Abbreviations: dt: dorsal tuberosity.
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Cretaceous Research 128 (2021) 104957
centra, with the bony rim that limits the posterior end very pro-
nounced in most of the elements. The posterior condyles occupy
only a small portion (less than 13%) of the total centrum length in
the most complete vertebrae. These condyles are bounded from the
main body of the centrum by a distinct border. This border is much
more pronounced on the lateral sides of the condyles. The condylar
apices are located in the middle of the articulation, slightly
ventrally displaced in MPCA-Pv 1500/18. The lateral surfaces
exhibit a slightly developed longitudinal ridge on the midline of the
centra. The same divides the surfaces in two halves, which are
slightly laterodorsally and lateroventrally oriented. The ventral
surfaces exhibit a shallow depression. The best preserved remain of
a chevron facet is recorded in the posterior margin of the element
MPCA-Pv 1500/17. The same appears to be poorly developed. The
lateroventral protuberance described in the posterior margin of the
specimen MPCA-Pv 1500/9 is absent in this segment of the tail.
Lateral blind fossae and large neurovascular foramina are also ab-
sent. Transverse processes are absent. Possible remains of a small
“dorsal tuberosity” are observed in MPCA-Pv 1500/15. Neverthe-
less, this structure is absent in the other elements.
The bases of the neural arches of the posterior middle caudals
are restricted to the anterior half of the centrum. Neural arches are
low, being less than 40% of the vertebral height. The neural spines
are posteriorly inclined, forming an angle of around 35 degrees
I. Cerda, V.L. Zurriaguz, J.L. Carballido et al.
with respect to the main axis of the centrum. These are dorso-
ventrally and laterally compressed. As pointed out by Salgado
(1996), the neural spines are anteroposteriorly elongated, with
their anterior edges located above the level of the posterior ones.
Although the posterior margins of the neural spines are posteriorly
displaced, they never reach the anterior margin of the posterior
articular surface. The dorsal margin of the neural spines is straight.
Prezygapophyses are incompletely preserved. They surpass the
anterior margin of the centra and appear to be oriented forming a
low angle (around 12 degrees) with the main axis of the centrum.
Postzygapophyses are posteriorly displaced, located above the
posterior half of the centrum. They are elliptical, with their longest
axes anteroposteriorly oriented. Their posterior edges do not sur-
pass the posterior margins of the neural spines. The post-
zygapophyseal facets are lateroventrally oriented.
4.2.3. Posterior caudal vertebrae
The posterior caudal vertebrae are represented by the last
twelve preserved elements of the caudal series (MPCA-Pv 1500/19-
30). The general morphology of the centra is simpler than in the
anterior and middle caudal vertebrae (Figs. 4OeU, 7AeD). In this
regard, transverse processes and the associated dorsal tuberosity,
lateral blind fossae and large neurovascular foramina are absent.
The centra are cylindrical and twice (or more) longer than high. The
width and height of the centra is rather the same in all the ele-
ments. However, the anterior and posterior contours are difficult to
evaluate because the elements are incomplete. Both lateral and
ventral faces of the centra are convex. This feature becomes more
pronounced towards the posteriormost elements. The posterior
articular condyle is only preserved in MPCA-Pv 1500/20, where it
occupies about 12% of the centrum length. Its apical border is
located in the middle of the articulation. Chevron facets are not
recorded in the posterior caudals. A particular feature is recorded
on the right lateral side of the elements MPCA-Pv 1500/21 and
MPCA-Pv 1500/21, in the place of the articulation between these
elements. In this area, a distinct protuberance of both elements is
recorded, which is more clearly observed in dorsal and ventral
Fig. 7. Posterior caudal vertebrae of Pellegrinisaurus powelli (MPCA-Pv 1500/21 and 22) in anterior (A), left lateral (B), posterior (C) and dorsal (D) views. Note the bulging
appearance of the articular area in the right side (black arrow). Scale bars equals to 5 cm.
9
Cretaceous Research 128 (2021) 104957
views (Fig. 7D). Contrary to other deformations recorded in this
individual, this particular osteological feature does not appear to be
a preservation artifact, being possibly related with an osseous
pathology.
As in the middle caudal vertebrae, the bases of the neural
arches are restricted to the anterior half of the centrum. However,
the anterior border of the bases is comparatively more posteriorly
displaced, leaving a wider space between them and the dorsal
margin of the anterior centrum edge. Therefore, the surface of the
centrum occupied by the base of neural arch reduces gradually
towards the posterior caudal vertebrae. The neural arches are
proportionally lower than in the middle caudal vertebrae, being
equal or less than 33% of the vertebral height. As in the middle
caudal vertebrae, the neural spines are laterally compressed and
anteroposteriorly elongated. The dorsal margin of the neural
spines is straight. The anterior margin of the neural spines is just
above the level of the posterior one, at least up to the element
MPCA-Pv 1500/23. The dorsal margin of the neural spines of the
posteriormost vertebrae is approximately at the same level as the
dorsal margin of the prezygapophyses. The posterior margin of the
neural spines is posteriorly displaced, located at the level of the
anterior border of the posterior condyle. The preserved pre-
zygapophyses are very elongated, with their anterior margins
almost reaching the mid length of the preceding vertebra. The
same are oriented parallel to the main axis of the centrum. Pre-
zygapophyseal facets are medially oriented. Postzygapophyses are
mostly not preserved, but appear to be located above the posterior
half of the centrum.
4.3. Femur
The only appendicular bone of the holotype corresponds to an
incomplete left femur (MPCA-Pv 1500/31, Fig. 8A-C). Both proximal
and distal ends of the femur are missing. By overlapping the
silhouette of the femur with the one of the titanosaur Patagotitan
mayorum, we estimated that the preserved portion corresponds to
about 61% of the total bone length, which approximate measures
I. Cerda, V.L. Zurriaguz, J.L. Carballido et al. Cretaceous Research 128 (2021) 104957

83 cm, and indicating a total reconstructed size of 1.4 m. The
minimum circumference of the shaft is 55.5 cm. A prominent
lateral bulge is developed between the level of the femoral head
and the beginning of the fourth trochanter. The posterior surface of
the proximal end exhibits a distinct trochanteric shelf on the lateral
region. This structure is oriented parallel to the lateral bulge, sur-
passing the distal end of the fourth trochanter. The distinct fossa
located between the lateral bulge and the trochanteric shelf ex-
hibits a rough surface. The fourth trochanter is reduced to a long,
poorly developed ridge with a rough surface. It is located in the
posteromedial margin of the shaft, on the proximal third of the
femur. Lateral to the fourth trochanter and on the posterior aspect
of the femoral shaft there is a well-developed muscle scar, possibly
related to the insertion site of the caudofemoralis brevis (Borsuk-

Fig. 8. Incomplete left femur of Pellegrinisaurus powelli (MPCA-Pv 1500/31) in anterior (A), medial (B) and posterior views. Striped areas indicate broken and/or covered with plaster
surfaces. Scale bars equal to 10 cm. Abbreviations: cas: caudofemoralis brevis attachment site; fc: femorotibialis crest; ft: fourth trochanter; lb: lateral bulge; plr: posterolateral
ridge; ts: trochanteric self.

10
I. Cerda, V.L. Zurriaguz, J.L. Carballido et al.
Białynicka 1977). There is a very low, dorsoventrally elongated crest
along the midline of the anterior surface of the shaft, which begins
at the level of the fourth trochanter and appears to correspond with
the femorotibialis crest (i.e. linea intermuscularis cranialis) (Otero
2010). The posterior side of the shaft exhibits a low but distinct
longitudinal ridge, which is laterally displaced and corresponds to
the distal extension of the trochanteric self. The midshat is elliptical
in cross-section, with a transverse diameter of about 162% the
anteroposterior diameter.
5. Femur histology
The samples show a poor preservation; however, the main
histological features are discernible. In transverse section, the
posterior side of the sample displays both compact and cancellous
bone (Fig. 9A). The medullar region exhibits thick trabeculae
enclosing intertrabecular spaces of varying sizes (Fig. 9B). These
cavities are surrounded by centripetally deposited layers of sec-
ondary lamellar bone. The trabecular bone contains remains of
secondary osteons. Some primary osteons are also visible. The
transition into the cortical bone is gradual. The cortex mostly
consists of secondary tissue formed during at least three genera-
tions (Fig. 9C). The secondary osteons predominate from the inner
to the mid cortex, and their density decrease towards the outer
cortex. In fact, no secondary osteons are recorded in the sub-
periosteal margin. Remains of parallel fibered bone tissue are
visible between the secondary osteons of the mid cortex (Fig. 9D).
This tissue is similar to the “modified laminar bone” (MLB) re-
ported by Klein et al. (2012). According to these authors, the MLB
consists of a primary tissue where the scaffolding of the fibrola-
mellar bone, which usually is laid down as matrix of woven bone,
is laid down as parallel fibered or lamellar bone matrix. The sec-
ond feature of this MLB is the arrangement of the vascular spaces,
which corresponds with a laminar pattern. Radial vascular spaces
are also present in the sampled element. Additionally, remains of
longitudinal primary osteons coated with a thick layer of lamellar
tissue are visible in some areas between the secondary bone
(Fig. 9E). Primary osteons in early stages of development (i.e.
containing few layers of lamellar bone) are observed at the
outermost portion of the cortex (Fig. 9F).
A peculiar feature is recorded in the outer cortex of the
sample, on the posterior side of the shaft. In this region, several
large (from 1 to 3 mm of diameter) erosion cavities of irregular
shape are recorded (Fig 9A). The gross microanatomy of this
unusual tissue is easily observed even with the naked eye and
consists of irregular spaces bounded by thin bony struts which
give it an appearance of cancellous bone. The spaces between the
bony struts are coated by a thin layer of lamellar bone tissue. The
remains of primary bone tissue located in the non-remodeled
areas between the large cavities consist of fibrolamellar bone.
Remains of secondary osteons are also visible in this portion of
the cortex (Fig. 9G). Osteocyte lacunae are numerous and are
densely grouped.
The outer cortex is interrupted by at least four single and double
lines of arrested growth (LAGs), which are recognized by a thin line
that interrupts the cortex (Fig 9H). Sharpey's fibers are present in
several parts of the outer cortex (Fig. 9I).
6. Phylogenetic analysis
In order to better understand the phylogenetic position of Pelle-
grinisaurus powelli amongst Titanosauria, we include the taxon into
the data matrix recently published by Gallina et al. (2021), expanding
the taxon sampling with the incorporation of four genera (Nullotitan
glaciaris, Diamantinasaurus matildae, Choconsaurus baileywillisi and
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Cretaceous Research 128 (2021) 104957
Pellegrinisaurus powelli), two new characters (419 and 420), and the
inclusion of an additional state in two already used character (236 and
251) (see Supplementary Material). Scores of Choconsaurus and Nul-
lotitan are based on publications (Simo
n et al., 2017; Novas et al.,
2019) and personal observations of the materials (JLC and LS), and
those from Diamantinasaurus were based on the holotype specimen
described by Poropat et al. (2015). Choconsaurus scores are solely
based on the holotype specimen. The resultant data matrix with 98
taxa and 420 characters was analyzed under equally weighted
parsimony analysis using TNT 1.5 (Goloboff and Catalano 2016). From
the 420 characters, 24 were treated as ordered (characters 14, 61, 100,
102, 109, 115, 127, 132, 135, 136, 167, 180, 196, 257, 260, 277, 279, 280,
282, 300, 304, 347, 353, and 355). We conducted two tree searches,
the first one under equally weighted (EWA; Equally Weighted Anal-
ysis) and a second one under implied weighted (IWA; Implied
Weighted Analysis).
The first tree search was performed, under equally weighted,
using the New Technologies search tool, conducting the search up
find 25 MPTs (Most Parsimonious Trees). The procedure retrieved
114 MPTs of 1508 steps, which were subjected to a second round
of TBR finding >400.000 MPTs. The large amount of MPTs recov-
ered is the result of the presence of 12 polytomies in the strict
consensus tree and the large number of possible resolutions of
them. Five of them include seven or more lineages, one of them is
present amongst rebbachisaurids (caused by the multiple posi-
tions that Rayososaurus agrioensis can take amongst rebbachi-
saurids). The other four major polytomies amongst
Titanosauriformes includes unresolved relationships amongst
brachiosaurids, basal somphospondylans, lognkosaurs, and
derived lithostrotians (Fig. S1). Pellegrinisaurus is recovered as part
of the polytomy that includes five lithostrotians, with this lineage
being part of a larger polytomy that includes several non-
colossosaur titanosaurians. The unstable positions of Malargüe-
saurus florenciae, and Ninjatitan zapatai are causing the polytomies
amongst basal somphospondylans and lognkosaurs respectively
(Fig. S2). Once these three taxa were pruned from the MPTs and
the repeated MPTs were deleted from the trees in memory (using
the command unique) a total of 81,279 trees were conserved. The
polytomy recovered within derived titanosaurs, which includes
Pellegrinisaurus, and 11 taxa, can be completely resolved if
Tapuiasaurus macedoi and Nemegtosaurus mongoliensis are
excluded from the strict consensus, which can be recovered in
multiple positions (Fig. 10A). Pellegrinisaurus is recovered as a
derived lithostrotian, closely related to Nullotitan, with Alamo-
saurus and Baurutitan as successive sister taxa (see below for a
more complete discussion).
The second analysis was performed under extended implied
weightings (Goloboff 2014) and using a k (concavity constant; see
Goloboff 1993) value of 9, instead of using the default value of 3 (see
Goloboff et al., 2018; Mannion et al., 2019). The tree search was
conducted using the New Technologies search tool, up find 25
MPTs. This procedure retrieves 88,560 MPTs of 74.07751 steps. The
complete strict consensus tree present seven polytomies, being the
major one present amongst rebbachisaurids (Fig. S3), which is be-
ing caused by the multiple position in which Rayososaurus can be
placed. Once Rayososaurus is pruned from the MPTs and repeated
trees are deleted a total of 17,977 MPTs were kept in memory.
Pellegrinisaurus is recovered in a polytomy that includes this taxon
and eight more derived lithostrotians. If Nemegtosaurus is
excluded from the consensus this polytomy is completely resolved
(Fig. 10B). Pellegrinisaurus is recovered as a derived lithostrotian,
closely related to the clade formed by Trigonosaurus and
Alamosaurus and with Baurutitan as the sister taxon of
Pellegrinisaurus þ (Trigonosaurus þ Alamosaurus) (see below for a
more complete discussion).
I. Cerda, V.L. Zurriaguz, J.L. Carballido et al. Cretaceous Research 128 (2021) 104957

Fig. 9. Bone histology of the left femur of Pellegrinisaurus powelli (MPCA-Pv 1500/31). A, general view of the posterior cross section of the femur. Note the large resorption cavities in
the outer cortex and three line of arrested growth (black arrows head). B, trabeculae of the cancellous bone at the medullar region. Note the intertrabecular spaces surrounded by

12
I. Cerda, V.L. Zurriaguz, J.L. Carballido et al. Cretaceous Research 128 (2021) 104957

Fig. 10. Phylogenetic relationships Pellegrinisaurus powelli. A, equally weighted analysis, reduce consensus tree after pruning Tapuiasaurus and Nemegtosaurus, which possible
positions are indicated with the letters a and b respectively. B, implied weighted analysis, reduce consensus tree after pruning Nemegtosaurus, which possible positions are indicated
with the letter a. For the complete results see Figs. S1eS3 in the Supplementary Materials.

7. Anatomical description of caudal vertebra MCS-Pv 53 8. Discussion

This specimen consists of a single element, interpreted here as a
middle caudal, which includes an almost complete centrum and an
incomplete neural arch (Fig. 11A-C). Natural fractures reveal that
pneumatic cavities are absent in both the centrum and the neural
arch. The vertebral centrum is strongly procoelous, with a deep
concave anterior face that is slightly wider than high. Excluding the
articular condyle, the centrum is slightly longer than high. The
posterior articular face has a heart-shaped outline, with its dorsal
margin wider (almost twice) than the ventral one. The condyle is
well developed, occupying almost 30% of the entire centrum length.
The apical margin of the condyle is dorsally displaced. The anterior
margin of the centrum is inclined, with its ventral portion poste-
riorly displaced and forming an angle of approximately 63 degrees
with the main axis of the centrum. The lateral surfaces of the
centrum are slightly concave and strongly lateroventrally oriented.
This inclination of the lateral surfaces results in a narrow ventral
surface. Chevron facets are distinct in the posterior face, where they
enclose a ventral furrow. Due to the incompleteness of the element,
it is not possible to determine the presence of transverse processes.
The base of the neural arch occupies the anterior half of the
centrum. The neural canal has an oval shape, being slightly wider
than high. Although the neural spine is not preserved, it appears to
be posteriorly inclined. Only the bases of the prezygapophyses are
preserved. These are robust and inclined, forming and angle of 12
degrees with the main axis of the centrum. There is a small but
deep fossa (spinoprezygapophyseal fossa) at the base of the neural
spine, between the bases of the prezygapophyses. The post-
zygapophyses are not preserved.
8.1. Diagnostic features of Pellegrinisaurus powelli
The anatomical review of the holotype of Pellegrinisaurus powelli
provided here allows us to review the original diagnosis of Salgado
(1996). Considering dorsal vertebrae, Salgado (1996) proposed two
diagnostic features: 1- dorsal centra strongly depressed and, 2- trans-
verse width of the centrum of the posterior dorsal vertebrae approxi-
mately twice the maximum dorsoventral depth. These characters,
however, are only recorded in two of the dorsal vertebrae analyzed
(MPCA-Pv 1500/2 and MPCA-Pv 1500/4). Furthermore, such features
are attributed to a substantial dorsoventral compressional deformation
suffered by the centra, which is evidenced by prominent displaced
fractures on the anterior and posterior sides of each element.
Regarding the caudal vertebrae, two diagnostic characters were
proposed by Salgado (1996): 1- midposterior and posterior caudals
with anteroposteriorly elongated and dorsoventrally depressed
neural spines and 2- anterior end of the neural spine placed at a
higher position than the posterior end in mid posterior and pos-
terior caudals. The first character is widely distributed amongst
titanosaurs (e.g. Malawisaurus dixeyi Ophistocoelicaudia skarzynskii,
Alamosaurus sanjuanensis, Baurutitan britoi; Gilmore 1946; Borsuk-
Bialynicka 1977; Gomani 2005; Kellner et al., 2005), so we consider
that this is not diagnostic of Pellegrinisaurus. The second character,
which is always present in combination with the first one, is also
present in at least two taxa: Alamosaurus sanjuanensis and Baur-
utitan britoi (Gilmore 1946; Kellner et al., 2005). Due to the close
phylogenetic relationship between Pellegrinisaurus and Alamosau-
rus, which form a clade with Nullotitan or Trigonosaurus (depending

lamellar bone. Polarized light. C, detail of the mid cortex showing three generations of secondary osteons (white arrows head). Normal transmitted light. D, detail of the mid cortex
showing parallel fibered bone under polarized light. Note the remodeling by secondary osteons. E, primary osteons of the mid cortex which exhibit thick layers of lamellar bone.
Polarized light. F, modified laminar bone at the outer cortex. Note that the primary osteons exhibit few layers of lamellar bone surrounding the vascular space. Polarized light. G,
detailed view of the bony struts of the unusual tissue. Note the remains of woven bone and a secondary osteon. Normal transmitted light. H, outer cortex showing lines of arrested
growth (white arrows head). Polarized light. I, outer cortex showing extensive areas dark (white arrows) by the presence of Sharpey's fibers. Polarized light. Scale bars equal:
1 mm (A), 0.5 mm (B, I), 0.3 mm (C, G-H) and 0.2 mm (DeF). Abbreviations: RC: resorption cavity; IS: intertrabecular space; LB: lamellar bone; SO: secondary osteons; PFB: parallel-
fibered bone; PO: primary osteons; WB: woven bone.

13
I. Cerda, V.L. Zurriaguz, J.L. Carballido et al.
Fig. 11. Middle caudal vertebra MCS-Pv 53 in anterior (A), right lateral (B) and posterior (C) views). Scale bar equals to 10 cm.
on the analysis) (Fig. 10), the second character is optimized as a
synapomorphy of this clade.
The anatomical redescription of the holotype of Pellegrinisaurus
powelli allows us to propose three new diagnostic characters for
this taxon. The first one is observed in the anterior caudal vertebrae
and consists of the presence of prominent blind fossae (“pleuro-
coels”) with well-developed dorsal rims in their dorsal margins.
Although the presence of lateral fossae in anterior caudal vertebra
has been reported in other titanosaurs, such as Patagotitan
(Carballido et al., 2017), Nullotitan (Novas et al., 2019) and Ninjatitan
(Gallina et al., 2021), the distinct rim in the dorsal border is unique
to Pellegrinisaurus. The other diagnostic character of Pelle-
grinisaurus is recorded in the posterior side of the femoral shaft and
femur and consist with the presence of a low longitudinal ridge
located on the posterolateral area (Fig. 8C). An elongated longitu-
dinal ridge (i.e. extending to the distal third of the shaft) on the
posterolateral side of the femur has been only recorded in Rape-
tosaurus krausei, Jainosaurus septentrionalis, and an indeterminate
titanosaur from the Maastrichtian of the southern Pyrenees (Curry
Rogers 2009; Wilson et al., 2011b; Vila et al., 2012).
8.2. Ontogenetic stage of Pellegrinisaurus powelli MPCA-Pv 1500
Bone histology provides valuable information regarding the
ontogenetic stage of Pellegrinisaurus powelli MPCA-Pv 1500. The
absence of an Outer Circumferential Layer (OCL, i.e. peripheral band
of lamellar or parallel fibered bone with closely packed growth
lines, also referred as External Fundamental System) in the sampled
femur reveals that the individual was not somatically mature at
time of death (Chinsamy Turan 2005). Nevertheless, since the
possibility that an OCL could be present in the specimen but lost
prior to histological sampling (e.g. during field extraction) can not
entirely ruled out, the subadult condition of the individual is
tentative. Using the criteria proposed by Klein and Sander (2008)
for histological ontogenetic stage (HOS) determination in
sauropod dinosaurs, the microstructure of MPCA-Pv 1500 corre-
sponds with a HOS 10. Since sexual maturity is apparently reached
at the HOS 8, the current data indicates that the specimen MPCA-Pv
1500 correspond with a sexually mature subadult individual.
Recent studies have proposed that titanosaurs show a higher
degree of Haversian remodeling compared with other sauropods
(Klein et al., 2009, 2012; Stein et al., 2010; Company 2011). This
feature was documented only in the largest (i.e., oldest) specimens
of basal neosauropods and basal macronarians (Klein and Sander
2008). According to this, the histology of the femur of MPCA-Pv
1500 exhibits extensive secondary reconstruction from the inner
to the mid cortex (with at least three generation of secondary
osteons), in an apparently not fully grown individual (subadult).
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Cretaceous Research 128 (2021) 104957
Thus, our observations concur with the idea of the intense Haver-
sian remodeling in Titanosauria.
To date, the unusual tissue observed in the outer cortex of the
sample has not been reported in any other sauropod. The position of
the large resorption cavities near of the outer cortex is unexpected
because the remodeling is often more intense deeper within the
bone cortex. The reason for such intense remodeling in this partic-
ular region of the cortex is elusive. A possible explanation could be
that is the product of microcracks (microdamage) in the bone cortex.
Several studies have shown that microdamage is a significant factor
in the initiation of intracortical bone remodeling (Burr et al., 1985;
Mori and Burr 1993; Bentolila et al., 1998; Lee et al., 2002). Other
stimuli such as hormonal and metabolic changes also cause bone
remodeling (Lee et al., 2002). Nevertheless, as previously
mentioned, the resorption cavities depart from the typical second-
ary osteons morphology. The causes of the origin of the unusual
tissue should be explored with more detail in future studies.
8.3. Taxonomical assignment of MCS-Pv 53
In their review of Patagonian sauropodomorphs, Salgado and
Bonaparte (2007) referred to Pellegrinisaurus powelli a single mid-
dle caudal vertebra (MCS-Pv 53), which has been here described
and figured. Such assignment, however, is not supported by the
anatomical features recorded in the element. First, diagnostic
characters of Pellegrinisaurus are absent in the element, irrespective
of which diagnosis is considered (the original provided by Salgado
1996 or the new here proposed). Second, the MCS-Pv 53 strongly
differs from the morphology of middle caudal vertebrae of Pelle-
grinisaurus holotype, so it cannot be considered as cf. Pelle-
grinisaurus. Comparing with other titanosaurs from the same
locality (i.e. Neuquensaurus australis and Laplatasaurus araukanicus)
MCS-Pv 53 clearly differs from the caudal vertebrae of Neu-
quensaurus, which are dorsoventrally compressed, with a wide and
concave ventral surface and pneumatized neural arches (Powell
2003; Salgado et al., 2005; Zurriaguz and Cerda 2017). Regarding
Laplatasaurus araukanicus, the only specimen referable to this taxon
correspond with the lectotype, which only consist of a tibia and
fibula (Gallina and Otero 2015), precluding a proper comparison
with MCS-Pv 53. In sum, considering the absence of diagnostic
characters at a generic level, we refer specimen MCS-Pv 53 to
Titanosauria indet.
8.4. Phylogenetic relationships of Pellegrinisaurus powelli
In the two analyses here presented (under equally and extended
implied weighted) Pellegrinisaurus is recovered as a non-saltasaurid
titanosaurian, as in most phylogenetic analyses in which it was
I. Cerda, V.L. Zurriaguz, J.L. Carballido et al.
included, with the main exception of the data set used by Gorscak
and O'Connor (2016) and analyses derived from it (Sallam et al.,
2018; Gorscak and O'Connor 2019). Nevertheless, in these ana-
lyses the inclusion of Pellegrinisaurus as a Saltasauridae, is not based
on the different derived position of it but on the unusual positions
in which Saltasaurus and Opisthocoelicaudia (the specifier taxa of
Saltasauridae) are recovered. In the present analyses Pelle-
grinisaurus is recovered as a closely related taxon to Alamosaurus
and Baurutitan, with Baurutitan recovered in both analyses as the
basal taxon of this clade (Fig. 10A, B), which depends on the ana-
lyses could include to Nullotitan or Pellegrinisaurus. In the equally
weighted analysis Nullotitan (from the Campanian of Santa Cruz) is
recovered as the sister taxon to Pellegrinisaurus, and with Alamo-
saurus (from the Maastrichtian of North America) as the sister clade
of them (Fig. 10A). Under the extended implied weighted analysis
Pellegrinisaurus is recovered as the sister taxon of Alamosaurus plus
Trigonosaurus (from the Campanian-Maastrichtian of Brazil).
Whereas in both analyses Nullotitan and Trigonosaurus are recov-
ered as lithostrotians, Notocolossus is recovered as a lithostrotian in
the EWA or as a colossosaur titanosaurian, closely related to the
origin of Lognkosauria in the extended IWA (Fig. 10). The position
here recovered for Nullotitan (included for the first time in a
phylogenetic analysis) contrasts with that originally proposed by
Novas et al. (2019) as a colossosaur titanosaurian.
In both analyses the clade formed by Baurutitan and more
derived lithostrotians (MDL) is supported by two common syna-
pomorphic characters. 1- presence of a dorsal tuberosity on the
transverse processes of anterior caudal vertebrae (character 236).
Amongst this clade such dorsal tuberosity if present in Baurutitan,
Alamosaurus, and Pellegrinisaurus, it is also present in Trigonosaurus
(which is part of this clade in the IWA) and is not present in Nul-
lotitan (part of this clade in the EWA). Outside this clade, the dorsal
tuberosity is convergently acquired in Epachthosaurus and some
euhelopodids (Huabeisaurus and Xiangshanosaurus). 2- ante-
roposteriorlly elongated neural spine (around the half of the
centrum length) which anterodorsal edge is above the poster-
odorsal one (character 420). Amongst scored taxa this character is
solely present in Baurutitan, Alamosaurus and Pellegrinisaurus.
Under the IWA the clade formed by Pellegrinisaurus and MDL is
supported by one character, vertically oriented neural spines in
posteriormost anterior and middle caudal vertebrae (character
257). Whereas the neural spines of most titanosaurs are posteriorly
directed, in Pellegrinisaurus and Alamosaurus it is vertically ori-
ented. The more derived clade formed by Alamosaurus and Trig-
onosaurus is supported by one derived character, the presence of a
slightly developed dorsal tuberosity on the transverse processes of
anterior caudal vertebrae (character 419). This less development of
the dorsal tuberosity contrast to the high development of such
structure in Pellegrinisaurus and Baurutitan, convergently strongly
developed in Epachthosaurus and Xianshanosaurus.
Under the EWA the clade formed by Alamosaurus and MDL is
supported by two synapomorphies. 1- anterior caudal vertebrae
with a lateral pneumatic foramen (character 228). The presence of a
lateral pneumatic foramen in anterior caudal vertebrae is an un-
common character amongst titanosaurs, and among this clade is
present in Alamosaurus, Pellegrinisaurus and Nullotitan (under this
analysis a monophyletic clade) and is convergently acquired in
Saltasaurus and in Ninjatitan and Patagotitan. 2- elongated ventral
longitudinal hollows in anterior and middle caudal vertebrae
(character 251). This character is present in Alamosaurus and Pel-
legrinisaurus, differing from the shorter ventral hollows of Nulloti-
tan. Elongated ventral hollows are convergently acquired in
Saltasaurus and Opisthocoelicaudia. The sister relationships of Nul-
lotitan and Pellegrinisaurus, obtained in the EWA is based on the
presence of two synapomorphies: 1- presence of large blind fossae
15
Cretaceous Research 128 (2021) 104957
(here scored as “pleurocoels”) on the lateral surface of the centrum
in anterior caudal vertebrae (character 232) and; 2- transversely
concave ventral surface of the caudal centrum. As previously
mentioned, the presence of large blind fossae is an uncommon
character amongst titanosaurs, only reported in Patagotitan
(Carballido et al., 2017), Nullotitan (Novas et al., 2019), Ninjatitan
(Gallina et al., 2021), Savannasaurus (Poropat et al., 2020) and Pel-
legrinisaurus. Markedly concave ventral surface of the caudal
centrum (character 233) is observed, amongst titanosaurians, in a
few taxa such as Saltasaurus, Opisthocoelicaudia, Drusilasaurus, and
Andesaurus.
Irrespective of the criterium followed for the analyses here
presented (EWA or IWA) the strength relationships between
Baurutitan, Pellegrinisaurus and Alamosaurus were recovered in
both searches. As mentioned above most previous analyses co-
incides in recovered Pellegrinisaurus from the Upper Cretaceous of
Argentina with Alamosaurus from the Upper Cretaceous of North
America. Other analyses that do not include Pellegrinisaurus in their
taxon sampling showed a closer relationship between the north
American titanosaur and saltasaurid titanosaurians (a clade defined
by Opisthocoelicaudia from the Upper Cretaceous of Asia and Sal-
tasaurus from the Upper Cretaceous of Argentina) (e.g., Salgado
et al., 1997b; Wilson 2002; Upchurch et al., 2004; Carballido and
Sander 2014; Poropat et al., 2015; Carballido et al., 2017). The
phylogenetic results here presented have a direct impact on the
present hypothesis surrounding the sauropod hiatus in North
America (see D'Emic et al., 2010; Mannion and Upchurch 2011; and
D'Emic and Foreman 2012 for a recent discussion on it). Based on
the supposed relationships between Alamosaurus and other tita-
nosaurs from South America and Asia it was suggested that derived
titanosaurs dispersed to North America from either Asia or South
America (Lucas and Hunt 1989; Wilson and Sereno 1998; D'Emic
et al., 2010). Based on our results it is much more plausible to
explain the presence of Alamosaurus as a dispersion from South
America than Asia, as such hypothesis is supported by the re-
lationships of Alamosaurus with the Argentinean and Brazilian
Upper Cretaceous titanosaurs Baurutitan, Pellegrinisaurus and
perhaps Nullotitan or Trigonosaurus (depending on the criterium
followed for the analysis).
9. Conclusions
A complete preparation and detailed study of the holotype
specimen of Pellegrinisaurus powelli MPCA 1500 provides a wealth
of new information on the anatomy, diagnosis, and phylogenetic
position of this taxon. The diagnostic characters originally proposed
by Salgado (1996) are here considered invalid because, either they
are due to diagenetic deformations, or they are widely distributed
amongst titanosaurs. The new diagnostic features are the presence
of large lateral blind fossae with distinct dorsal rims in the anterior
caudal centra, and the presence of a low longitudinal ridge on the
posterior side of the femur shaft. Regarding its phylogenetic posi-
tion, Pellegrinisaurus is recovered as non-saltasaurid lithostrotian,
closely related with Alamosaurus sanjuanensis. The recognized
microstructural features suggest a subadult (i.e. sexually mature
but still growing) ontogenetic stage for Pellegrinisaurus powelli
MPCA 1500. Finally, anatomical review of a caudal vertebra previ-
ously assigned to Pellegrinisaurus reveals that the same can only be
assigned to Titanosauria indet.
Acknowledgements
The authors would like to thank Carlos Mun ~ oz (MPCA), Natalia

n Amigos del Museo of Cinco
Cides (MCS) and all the staff of Comisio
Saltos for access to the specimens under their care. Jaime Powell
I. Cerda, V.L. Zurriaguz, J.L. Carballido et al.
(R.I.P) provided unpublished pictures of the Pellegrinisaurus exca-

Aravena for the preparation of the material.
vation. We thank Jose
Lorenzo Novak found and brings us the attention of specimen MCS-
Pv 53. Jorge Blanco has skillfully made the illustrations on Figs. 5, 6
and 8. The constructive comments of Agustín Martinelli and Philip
Mannion have greatly improved this manuscript. Sci-hub library
and Wikipaleo group shared valuable publications for this research.
Part of this project was developed with the financial support of
Agencia Nacional de Promocio
n Científica y Tecnolo
gica (PICT-
2015-2021 to I.A.C and PICT-1925 to J.L.C).
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Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.org/10.
1016/j.cretres.2021.104957.