Cretaceous Research xxx (2012) 1e10

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

A new peirosaurid from the Upper Cretaceous of Argentina: Implications for

specimens referred to Peirosaurus torminni Price (Crocodyliformes: Peirosauridae)
Agustín G. Martinelli a, *, Joseph J.W. Sertich b, Alberto C. Garrido c, Ángel M. Praderio d
a
UFTM/Complexo Cultural e Científico de Peirópolis, Centro de Pesquisas Paleontológicas L.I. Price, 38001-970, BR-262, Km 784, Bairro Peirópolis, Uberaba, MG, Brazil
b
Denver Museum of Nature and Science, Department of Earth Sciences, 2001 Colorado Boulevard, Denver, CO 80205, USA
c
Museo Provincial de Ciencias Naturales “Profesor Dr. Juan A. Olsacher”, Etcheluz y Ejército Argentino, Zapala 8340, Neuquén, Argentina
d
Centro Regional de Investigación y Desarrollo Cultural (CRIDC), Adrián Illesca N 138, Malargüe, Mendoza, Argentina

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

Article history: Two crocodyliform specimens recovered from the Upper Cretaceous of Argentina and originally referred
Received 29 November 2011 to Peirosaurus torminni Price from Minas Gerais State, Brazil are reassigned to Gasparinisuchus peir-
Accepted in revised form 22 March 2012 osauroides gen. et sp. nov. No unique features are shared between the Brazilian and Argentinian speci-
Available online xxx
mens and many traits, including the presence of five premaxillary teeth, a noticeably smooth perinarial
region, a lateral notch at the premaxillaemaxilla contact for the reception of an enlarged anterior
Keywords:
dentary tooth, and the shape of the external nares, are widely distributed among peirosaurids and, in
Mesoeucrocodylia
some cases, other mesoeucrocodylians. Though overlapping elements are limited to the premaxilla and
Peirosauridae
Late Cretaceous
the dentition, the broad, rounded rostrum, anteroposteriorly short premaxilla, reduced perinareal fossa,
Argentina and short premaxillary interdental spaces in Gasparinisuchus differentiate it from P. torminni. Addition-
Brazil ally, Gasparinisuchus is unique among peirosaurids for its broad rostrum and apparent absence of an
antorbital fenestra. Reassignment of the Argentinian material formerly referred to Peirosaurus therefore
removes one of the few specific links between the Cretaceous terrestrial biotas of Brazil and Argentina.
Ó 2012 Elsevier Ltd. All rights reserved.

1. Introduction (Larsson and Gado, 2000) from the Aptian of Niger, though several
recent studies (e.g., Lapparent de Broin, 2002; Carvalho et al., 2007;
Peirosaurid crocodyliforms represent a relatively common Turner and Buckley, 2008) exclude Stolokrosuchus from Peirosaur-
component of many Cretaceous terrestrial faunas known from idae. The broad-snouted crocodyliform Mahajangasuchus insignis
Gondwana. At present, South American members of Peirosauridae (Buckley and Brochu, 1999; Turner and Buckley, 2008) from the
include at least seven taxa: Barcinosuchus gradilis (Leardi and Pol, Maastrichtian of Madagascar was originally described as a potential
2009), Itasuchus jesuinoi (Price, 1955), Lomasuchus palpebrosus peirosaurid, though more recent phylogenetic analyses exclude it
(Gasparini et al., 1991), Montealtosuchus arrudacamposi (Carvalho from this clade (e.g., Turner, 2006; Carvalho et al., 2007; Larsson
et al., 2007), Peirosaurus torminni (Price, 1955), Pepesuchus deiseae and Sues, 2007; Turner and Buckley, 2008; Sereno and Larsson,
(Campos et al., 2011), and Uberabasuchus terrificus (Carvalho et al., 2009; Turner and Sertich, 2010). An additional taxon from these
2004). These taxa are thus far restricted to the Cretaceous same deposits in Madagascar, Miadanasuchus oblita (Buffetaut and
(AptianeMaastrichtian) of Argentina and Brazil, with the Late Taquet, 1979; Simons and Buckley, 2009), may also represent
Cretaceous taxon Peirosaurus torminni representing the only peir- a peirosaurid taxon, though currently described specimens are too
osaurid common to both countries (Gasparini, 1982; Gasparini fragmentary for a confident assignment.
et al., 1991; but see below). African taxa include Hamadasuchus Within Peirosauridae, little consensus exists regarding ingroup
rebouli (Buffetaut, 1994; Larsson and Sues, 2007) from the relationships, specifically the phylogenetic position of African taxa
AlbianeCenomanian of Morocco and Stolokrosuchus lapparenti relative to South American taxa from which the family was origi-
nally established (Gasparini, 1982). Additionally, the phylogenetic
position of Peirosauridae within Mesoeucrocodylia is still poorly
resolved. Among widely divergent recent phylogenetic hypotheses,
* Corresponding author.
Peirosauridae has been recovered as the sister-group of Neosuchia
E-mail addresses: [email protected] (A.G. Martinelli), jsertich@
dmns.org (J.J.W. Sertich), [email protected] (A.C. Garrido), (e.g., Pol and Apesteguía, 2005; Gasparini et al., 2006; Turner and
[email protected] (Á.M. Praderio). Buckley, 2008; Leardi and Pol, 2009); nested within the clade

0195-6671/$ e see front matter Ó 2012 Elsevier Ltd. All rights reserved.
doi:10.1016/j.cretres.2012.03.017

Please cite this article in press as: Martinelli, A.G., et al., A new peirosaurid from the Upper Cretaceous of Argentina: Implications for specimens
referred to Peirosaurus torminni Price (Crocodyliformes: Peirosauridae), Cretaceous Research (2012), doi:10.1016/j.cretres.2012.03.017
2 A.G. Martinelli et al. / Cretaceous Research xxx (2012) 1e10

Sebecia, closely related to sebecid crocodyliforms (e.g., Sebecus and 1750 PV has become the primary source of phylogenetic infor-
Bretesuchus) (Larsson and Sues, 2007); included in a clade with mation for P. torminni in most analyses of mesoeucrocodylian
Mahajangasuchus, Trematochampsa, and Araripesuchus spp. (Turner, relationships.
2006); recovered within Neosuchia (Sereno and Larsson, 2009); or The holotype of Peirosaurus torminni (DGM-433-R) and the
recovered within Notosuchia (Carvalho et al., 2004; Turner and primary referred Argentinian material (MOZ 1750 PV) unfortu-
Sertich, 2010). The phylogenetic relationships of Peirosauridae nately share very few overlapping elements (the premaxilla and
among mesoeucrocodylians are beyond the scope of this project the dentition). The recent description of several new South
and will not be discussed here. American peirosaurid taxa (e.g., Uberabasuchus, Montealtosuchus)
Originally described by Price (1955), Peirosaurus torminni have demonstrated that previously apomorphic features shared by
(Mesoeucrocodylia, Peirosauridae) is based upon a single frag- the Argentinian and Brazilian specimens of P. torminni have a
mentary specimen, DGM-433-R. This specimen was discovered at broader distribution among mesoeucrocodylians. Detailed rein-
the site known as “Ponto 2 do Price”, near the rural town of Peir- vestigation and re-evaluation of putative features shared between
ópolis (Uberaba County, Minas Gerais State, Brazil), in the Serra da P. torminni (holotype, DGM-433-R) and referred Argentinian
Galga Member, Marília Formation (Bauru Group, Maastrichtian). specimens (MOZ 1750 PV and PV-CRIDC-12) indicate substantial
The holotype of P. torminni (DGM-433-R) consists of the left differences. Thus, critical re-examination of material referred to
premaxilla with five teeth (Fig. 1), various associated, isolated teeth, P. torminni and evaluation of their taxonomic status is undertaken
the right palpebral, and many postcranial elements (i.e., radius, here. Furthermore, a preliminary discussion of the taxonomy of
ulnae, fragments of the ischia and pubis, vertebrae, ribs, chevrons, peirosaurid taxa from Peirópolis (Minas Gerais State, Brazil) is also
and osteoderms). Of these elements, only the premaxilla, selected undertaken.
teeth, and osteoderms have been figured (Price, 1955; Marinho
et al., 2006). Institutional abbreviations. CPPLIP, Centro de Pesquisas Paleon-
Subsequent to the original description of Peirosaurus torminni by tológicas Llewellyn Ivor Price, Peirópolis, Uberaba County, Minas
Price (1955), cranial and postcranial remains (MOZ 1750 PV; Gerais, Brazil; CRIDC, Centro Regional de Investigación y Desarrollo
Figs. 2e5) from Loma de la Lata, Neuquén Province, Argentina (Bajo Cultural, Malargüe, Mendoza, Argentina; DGM, Departamento
de la Carpa Formation, Santonian, Neuquén Group; Calvo and Nacional da Produção Mineral, Museu de Ciências da Terra, Rio de
Porfiri, 2010; Garrido, 2010) were referred to the Brazilian taxon. Janeiro, Brazil; MOZ, Museo Provincial de Ciencias Naturales “Pro-
In addition, incomplete cranial remains (PV-CRIDC-12) were fesor-Dr. Juan A. Olsacher”, Zapala, Neuquén, Argentina.
preliminarily described and tentatively assigned to P. torminni by
Praderio et al. (2009; see Fig. 6). This material, which also includes 2. Systematic palaeontology
undescribed postcranial elements, was recovered from the area
known as Cañadón Amarillo, in the southernmost region of Malar- Archosauria Cope, 1869
güe District, Mendoza Province, Argentina (Praderio et al., 2009), Crocodyliformes Hay, 1930 (sensu Clark, in Benton and Clark, 1988)
from the Santonian Anacleto Formation of the Neuquén Group. Mesoeucrocodylia Whetstone and Whybrow, 1983
The referral of specimen MOZ 1750 PV from the Upper Creta- Peirosauridae Gasparini, 1982
ceous of Argentina to Peirosaurus torminni greatly enhanced the Gasparinisuchus gen. nov.
anatomical and taxonomic understanding of the species,
providing the basis for Gasparini (1982) to define Peirosauridae. Derivation of name. Gasparini, in honour of the accomplished
Following the comprehensive description of this material vertebrate paleontologist Zulma Brandoni de Gasparini for her
(Gasparini, 1982; Gasparini et al., 1991), the presence of P. torminni prolific work on South American crocodyliforms, and suchus from
in both the Santonian of Argentina and Maastrichtian of Brazil was the Greek Souchos, with reference to the Egyptian crocodile-headed
widely accepted. Additionally, the nearly complete skull of MOZ god Sobek.

Fig. 1. Left premaxilla of the holotype of Peirosaurus torminni (DGM-433-R) in A, lateral and B, ventral views. Scale bar represents 2 cm. Abbreviations: amp, anteromedial process of
the premaxilla; en, external naris; M, maxilla; pd, paracanine depression or fossa; Pm, premaxilla; pn, perinareal region of the premaxilla; 1e5, tooth position. Horizontal line in B
indicates the estimated midline of the skull.

Please cite this article in press as: Martinelli, A.G., et al., A new peirosaurid from the Upper Cretaceous of Argentina: Implications for specimens
referred to Peirosaurus torminni Price (Crocodyliformes: Peirosauridae), Cretaceous Research (2012), doi:10.1016/j.cretres.2012.03.017
 A.G. Martinelli et al. / Cretaceous Research xxx (2012) 1e10 3

Fig. 2. Skull of the holotype of Gasparinisuchus peirosauroides gen. et sp. nov. (MOZ 1750 PV) with interpretive line drawing in dorsal view. Scale bar represents 5 cm. Grey areas
indicate broken and reconstructed surfaces. Abbreviations: F, frontal; J, jugal; L, lacrimal; M, maxilla; N, nasal; P, parietal; pd, paracanine depression or fossa; Pf, prefrontal; Pm,
premaxilla; sf, supratemporal fossa; Sq, squamosal; tr, tooth root; ?, reconstructed surface with unassociated bone fragment.

Type and only known species. Gasparinisuchus peirosauroides sp. nov. Remarks. Postcranial remains included within MOZ 1750 PV cited
by Gasparini (1982) and Gasparini et al. (1991) are currently
Diagnosis. As for the species. missing.
Gasparinisuchus peirosauroides sp. nov.
Referred specimen. PV-CRIDC-12, right premaxilla and maxilla
Figs. 2e6
(Fig. 6), isolated teeth, and part of the postcranial skeleton (origi-
Derivation of name. Peirosauroides, with reference to the signifi- nally tentatively referred to Peirosaurus torminni; Praderio et al.,
cance of the holotype specimen as the basis for the designation of 2009).
the Family Peirosauridae by Gasparini (1982).
Locality and horizon. MOZ 1750 PV was collected from Loma de la
Holotype. MOZ 1750 PV, partial skull, lower jaws (Figs. 2e5), and Lata locality, Neuquén Province, Argentina; Bajo de la Carpa
various associated postcranial remains (originally referred to Peir- Formation, Neuquén Group (Santonian, Upper Cretaceous; Calvo
osaurus torminni; Gasparini, 1982; Gasparini et al., 1991). and Porfiri, 2010; Garrido, 2010). PV-CRIDC-12 was collected from

Please cite this article in press as: Martinelli, A.G., et al., A new peirosaurid from the Upper Cretaceous of Argentina: Implications for specimens
referred to Peirosaurus torminni Price (Crocodyliformes: Peirosauridae), Cretaceous Research (2012), doi:10.1016/j.cretres.2012.03.017
4 A.G. Martinelli et al. / Cretaceous Research xxx (2012) 1e10

Fig. 3. Skull of the holotype of Gasparinisuchus peirosauroides gen. et sp. nov. (MOZ 1750 PV) with interpretive line drawing in ventral view. Scale bar represents 5 cm. Grey areas
indicate broken and reconstructed surfaces. Abbreviations: Bs, parabasisphenoid; Bo, basioccipital; c, occipital condyle; M, maxilla; mEf, median Eustachian foramen; Pa, palatine;
Pm, premaxilla; pd, paracanine depression or notch; Q, quadrate; sin, septum of the internal nares. Numbers refer to tooth position.

Cañadón Amarrillo, south Malargüe city, Mendoza Province, absence of contact between nasal and lacrimal; parabasisphenoid
Argentina; Anacleto Formation, Neuquén Group (early Campanian, and basioccipital well-exposed in ventral view; alveolar border of
Upper Cretaceous; Leanza and Hugo, 2001; Garrido, 2010). mandible festooned at level of fourth (hypertrophied) and thir-
teenth teeth; mandibular symphysis extending posteriorly to level
Diagnosis. Peirosaurid crocodyliform diagnosed by the following of eighth tooth; splenials occupying approximately 20% of
autapomorphies (marked with an asterisk) and combination of symphyseal length in dorsal aspect; five premaxillary and at least
characters: snout almost as wide as temporal region; facial portion 14 maxillary teeth; 18 mandibular teeth; moderate degree of
of premaxilla anteroposteriorly short; shallow, lateral groove at variation of tooth morphology along tooth row.
premaxillomaxillary suture for reception of hypertrophied fourth
mandibular tooth; large, convex alveolar border of maxilla at level Remarks. The phylogenetic position of the holotype specimen of
of hypertrophied third tooth; slightly convex alveolar border at Gasparinisuchus peirosauroides was previously tested in several
level of eighth and ninth teeth; absence of antorbital fenestra*; recent analyses under the taxon name Peirosaurus torminni (e.g.,
nasal anteroposteriorly long, occupying 50% of skull length; Gasparini et al., 2006; Larsson and Sues, 2007; Turner and Buckley,

Please cite this article in press as: Martinelli, A.G., et al., A new peirosaurid from the Upper Cretaceous of Argentina: Implications for specimens
referred to Peirosaurus torminni Price (Crocodyliformes: Peirosauridae), Cretaceous Research (2012), doi:10.1016/j.cretres.2012.03.017
 A.G. Martinelli et al. / Cretaceous Research xxx (2012) 1e10 5

Fig. 4. Mandible of the holotype of Gasparinisuchus peirosauroides gen. et sp. nov. (MOZ 1750 PV) with interpretive line drawing in dorsal view. Scale bar represents 5 cm. Grey areas
indicate broken surfaces. Abbreviations: D, dentary; Sp, splenial. Numbers refer to tooth position.

2008; Leardi and Pol, 2009; Turner and Sertich, 2010). Therefore, peirosaurid relationships in light of recent discoveries (e.g., Uber-
the position of Gasparinisuchus, the phylogenetic resolution of abasuchus, Montealtosuchus, Hamadasuchus, Stolokrosuchus). Mate-
peirosaurid interrelationships, and the position of Peirosauridae rial here referred to G. peirosauroides (PV-CRIDC-12) was briefly
within Mesoeucrocodylia will not be tested here. described by Praderio et al. (2009) and will be discussed in detail
elsewhere (Praderio, in prep.).
Description. A description of MOZ 1750 PV was provided by
Gasparini (1982) and Gasparini et al. (1991). A brief re-description Skull. The skull of Gasparinisuchus is heavily damaged, bearing
of the material, including additional comments and figures, several cracks that obscure many external sutures. The external
is provided here with emphasis on morphology relevant to surface of the skull is strongly ornamented with irregular pits and

Fig. 5. Left postdentary mandibular fragment of the holotype of Gasparinisuchus peirosauroides gen. et sp. nov. (MOZ 1750 PV) with interpretive line drawing in dorsal view. Scale
bar represents 5 cm. Grey areas indicate broken surfaces. Abbreviations: An, angular; Ar, articular; rart, retroarticular process; ?Sa, possible fragment of surangular.

Please cite this article in press as: Martinelli, A.G., et al., A new peirosaurid from the Upper Cretaceous of Argentina: Implications for specimens
referred to Peirosaurus torminni Price (Crocodyliformes: Peirosauridae), Cretaceous Research (2012), doi:10.1016/j.cretres.2012.03.017
6 A.G. Martinelli et al. / Cretaceous Research xxx (2012) 1e10

Fig. 6. Left premaxilla of the referred specimen of Gasparinisuchus peirosauroides gen. et sp. nov. (PV-CRIDC-12) in A, lateral and B, ventral views. Scale bar represents 2 cm.
Abbreviations: M, maxilla; L, lacrimal. Numbers refer to tooth position.

grooves, especially on the cranial table. The rostrum is nearly as The maxilla is exposed laterally and dorsally, occupying much
mediolaterally wide as the posterior portion of the skull, with of the external surface of the rostrum. In lateral view, the alveolar
a roughly ovoid outline in dorsal view (Fig. 2) differing from the border is ventrally convex at the level of the third, hypertrophied,
subtriangular, mediolaterally narrow rostra of other peirosaurids, tooth. In addition, moderate enlargement of the eighth and ninth
such as Uberabasuchus and Montealtosuchus. maxillary alveoli creates a second, much smaller convexity along
The premaxilla is anteroposteriorly short, convex and the the posterior maxillary margin, though this region of the maxilla
external surface is less heavily ornamented than the rest of the is damaged. Anteriorly, a large wedge-like anterior process of the
skull. The perinarial fossa is partially preserved, although it is maxilla is present on the premaxillaromaxillary suture, as in
smaller than those of Peirosaurus and Uberabasuchus as a result of other South American peirosaurids. Dorsally, the maxilla contacts
the overall short anteroposterior length of the premaxilla. The facial the nasal along a straight, anteroposteriorly directed suture
process of the premaxilla extends posteriorly, wedging between (Fig. 2). In ventral view, the palatal process of the maxilla is broad
the maxilla and nasal. In ventral view, the palatal process of the and smooth, with the alveolar portion projecting ventrally. On the
premaxilla is pierced by small vascular foramina near the alveolar right side, a fracture obscures a portion of the suture between
margin. The relatively small incisive foramen is bordered anteriorly the maxilla and palatine, though it appears that the maxilla
and laterally by the premaxillae and posteriorly by a small contri- contributes to the anterolateral margin of the suborbital fenestra
bution of the maxillae (Fig. 3). The premaxillomaxillary suture is (Fig. 3).
oblique in palatal view, directed anteromedially from the para- The nasals are paired and anteroposteriorly elongate with
canine depression to the incisive foramen. The margins of the almost parallel lateral margins, occupying approximately 50% of the
external nares are damaged but are directed anteriorly and slightly skull length (Fig. 2). The anteriormost portion of the nasals is
laterally. A laterally opened paracanine depression or fossa at the missing, and posteriorly only a portion of the suture with the right
premaxillomaxillary suture for reception of the fourth, hypertro- prefrontal and frontal can be clearly discerned. The nasal does not
phied mandibular tooth is present, as in other peirosaurids. This contact the lacrimal.
notch is moderately damaged on both sides of the skull but appears An antorbital fenestra or fossa cannot be clearly delineated in
to have been relatively shallow. MOZ 1750 PV as both sides of the skull are significantly damaged in

Please cite this article in press as: Martinelli, A.G., et al., A new peirosaurid from the Upper Cretaceous of Argentina: Implications for specimens
referred to Peirosaurus torminni Price (Crocodyliformes: Peirosauridae), Cretaceous Research (2012), doi:10.1016/j.cretres.2012.03.017
 A.G. Martinelli et al. / Cretaceous Research xxx (2012) 1e10 7

this region. However, in the referred specimen (PV-CRIDC-12) there third. Tooth positions one, two, and three are closely spaced,
is no evidence of an antorbital fenestra. whereas the remaining teeth are separated by a small interdental
On the right side of the snout of MOZ 1750 PV, a bone fragment space. The premaxillary teeth are conical and circular in cross-
may have been mistakenly attached during preparation of the section. In contrast, the maxillary teeth are conical to sub-conical,
specimen (labelled “?”. Fig. 2), though this portion is shown as with a moderately flat lingual face and a convex labial surface.
missing by Gasparini et al. (1991). Dorsally, the suture between the The first and second maxillary teeth are relatively small; the third
frontal and parietal is transversally directed and interdigitated. The tooth is hypertrophied and represents the largest tooth of the upper
frontal appears to contribute to the margin of the supratemporal dentition (Fig. 3). Posterior to the third tooth, the maxillary teeth
fenestra, as in most peirosaurids, but the degree of contribution decrease in size to the seventh position. The eighth and ninth
cannot be discerned. A small portion of the parietal-squamosal positions bear teeth slightly larger crowns. The last four teeth are
suture can be observed on the right side, though the squamosal is small, with short mediolaterally compressed crowns.
heavily damaged. The supratemporal fenestrae are incomplete but The first 13 mandibular teeth are conical to sub-conical, with
appear to be anteroposteriorly elongated. All other sutures and nearly planar lingual faces and convex labial surfaces. The fourth
structures on the dorsal aspect of the skull cannot be discerned. tooth is hypertrophied, preceded by three teeth of similar size and
Ventrally, the palatines, pterygoids, choanal margins, and much followed by six teeth that decrease slightly in size to the tenth tooth
of the basicranial region are damaged. Only part of the anterior half position (Fig. 4). Teeth 11e13 are relatively large, with the tooth in
of the suborbital fenestra is preserved, with its anterior margin position 13 being the largest (but not as large as the fourth). As in
extending to the level of the eleventh maxillary tooth. The para- the maxilla, the last five teeth are small, with short spatulate
basisphenoid is large, triangular shaped and, together with crowns. When preserved, both the upper and lower teeth possess
the basioccipital, well-exposed in ventral view (Fig. 3). The finely denticulate anterior and posterior carina.
basioccipital-parabasisphenoid contact defines an interdigitated
and elevated suture at the level of the ventral portion of the basal 3. Discussion
tubera. A large rounded opening is present between the parabasi-
sphenoid and basioccipital corresponding to the median Eusta- 3.1. Features shared by Peirosaurus and Gasparinisuchus (MOZ
chian foramen (Fig. 3). Only a small fragment of quadrate is 1750 PV) and their distribution among mesoeucrocodylians
preserved. In posterior (occipital) view, sutures between the otoc-
cipital, supraoccipital, parietal, and squamosal are not evident. The In order to re-evaluate systematically the previous referral of
foramen magnum is ovoid, bounded dorsally and laterally by the Argentinian specimen MOZ 1750 PV to Peirosaurus torminni
otoccipital and ventrally by the basioccipital. The occipital condyle (Gasparini, 1982; Gasparini et al., 1991), features used to unite these
is large, especially in comparison with Lomasuchus, protruding specimens are assessed here. Because overlap between these two
posteroventrally with a shallow basal constriction forming a specimens is limited to the dentition and premaxilla, many features
short neck. previously used to unite them (Gasparini, 1982; Gasparini et al.,
1991) are now recognized to be widely distributed among basal
Mandible. The mandible is externally ornamented with pits and mesoeucrocodylians.
furrows. Anteriorly, it is dorsoventrally low, with the ramus
becoming dorsoventrally taller posterior to the symphysis. The Smooth perinarial fossa. The perinarial fossa is defined as the
alveolar border is dorsally expanded at the level of both the portion of the external surface of the premaxilla lacking orna-
hypertrophied fourth tooth and the moderately enlarged thirteenth mentation and bordered anteriorly by the ventral and/or posterior
tooth. margin of the external naris, ventrally by the alveolar margin, and
In dorsal view, the mandibular symphysis is relatively wide, posteriorly by the ornamented facial portion of the premaxilla. One
posteriorly reaching to the level of the eighth tooth (Fig. 4). of the most conspicuous features on the premaxilla of Peirosaurus is
The splenials participate in the symphysis, occupying approxi- the presence of a large perinarial fossa located ventral and posterior
mately 20% of the symphyseal length in dorsal view and extending to the posteroventral margin of the external naris (Figs. 1, 7A).
anteriorly to the level of the sixth alveolus. Posterior to the Generally, the perinarial fossa of Peirosaurus has an inclined tear-
symphysis, the ventral border of the madibular ramus protrudes drop shape, with the convex posterior margin reaching the level of
lateroventrally. the root of the fourth premaxillary tooth. Posteriorly, the margin of
The region of the mandibular fenestra is broken and recon- the perinarial fossa is strongly delimited by the raised, heavily
structed with plaster. The posterior portion of the left lower jaw ornamented facial lamina of the premaxilla. Anteriorly, the peri-
was briefly described and figured by Gasparini (1982; Gasparini narial fossa slopes ventrally, reaching the narial margin and the
et al., 1991). At present, this portion is separated from the rest of alveolar border dorsal to the first two alveoli. In Gasparinisuchus,
the mandible and is badly preserved (Fig. 5). The glenoid of the both premaxillae are damaged in lateral view, obscuring this region
articular is axially concave and transversely broad. Anteriorly, the and limiting comparative information (Fig. 7B).
prearticular process of the articular is triangular in dorsal view. The A distinct perinarial fossa is widely distributed among meso-
retroarticular process is broken but apparently it was ante- eucrocodylians including sebecosuchians (e.g., Iberosuchus,
roposteriorly elongate, posteriorly protruding, and directed below Baurusuchus, Wargosuchus), Simosuchus (Buckley et al., 2000),
the level of the glenoid (Fig. 5). A portion of angular and possibly Araripesuchus, and peirosaurids (e.g., Uberabasuchus, Mon-
a fragment of surangular are preserved but their fragmentary tealtosuchus, Hamadasuchus). Among peirosaurids, the configura-
nature precludes any anatomical description. In lateral and ventral tion of the perinarial region of Peirosaurus is most similar to that of
views, the postdentary elements are heavily damaged. Uberabasuchus.

Dentition. The dentition consists of five premaxillary, 14 maxillary, Anteromedial process of premaxilla. The anteromedial portion of the
and 18 mandibular teeth (Figs. 3, 4). Most crowns are broken or premaxilla of Peirosaurus possesses an acute, strongly ante-
damaged in MOZ 1750 PV, missing much of the surface enamel in rodorsally projecting process contributing to the formation of an
preserved crowns. Premaxillary teeth increase in size from the first internarial bar (Fig. 1). This process is not preserved in Gaspar-
to the fourth (largest) position, the fifth is similar in size to the inisuchus (MOZ 1750 PV). In other peirosaurids such as

Please cite this article in press as: Martinelli, A.G., et al., A new peirosaurid from the Upper Cretaceous of Argentina: Implications for specimens
referred to Peirosaurus torminni Price (Crocodyliformes: Peirosauridae), Cretaceous Research (2012), doi:10.1016/j.cretres.2012.03.017
8 A.G. Martinelli et al. / Cretaceous Research xxx (2012) 1e10

interdental spaces can vary ontogenetically, thus limiting its taxo-

nomic value. Regardless, the condition of Peirosaurus is more
similar to that of other known Brazilian peirosaurids than to
Gasparinisuchus.
Price (1955) noted finely denticulated mesial and distal carninae
in the second through fifth teeth of Peirosaurus, but only a distally
denticulate carina in first premaxillary tooth. Unfortunately, this
feature cannot be observed in Gasparinisuchus (MOZ 1750 PV)
owing to the absence of crowns on the first teeth.

Rostrum morphology. As initially highlighted by Gasparini (1982;

also Gasparini et al., 1991), Gasparinisuchus possesses a broad,
relatively dorsoventrally compressed rostrum (Fig. 3) differing from
the tall pointed rostra of other peirosaurids such as Lomasuchus,
Montealtosuchus, Uberabasuchus, Stolokrosuchus, and Hama-
dasuchus. Though the complete rostrum is not preserved in Peir-
osaurus (DGM-433-R), the morphology of the premaxilla (Price,
1955, fig. 1; Fig. 1) indicates a transversely narrow anterior
rostrum similar to that of these other taxa. Therefore, the wide
snout of Gasparinisuchus is unique among known peirosaurids and
differs from the likely narrow, subtriangular snout of Peirosaurus.

Facial lamina of premaxilla. The posterior region of facial lamina of

the premaxilla of Peirosaurus (DGM-433-R) is damaged relative to
the original condition of the specimen as described by Price (1955,
fig. 1). Regardless, the facial surface of the premaxilla is ante-
roposteriorly elongate (Fig. 7), similar to the facial lamina of Uber-
abasuchus and Montealtosuchus but differing from Gasparinisuchus
in which the facial lamina of the premaxilla is relatively short
(Fig. 7).
Posterior to the last premaxillary alveolus, the premaxilla of
Peirosaurus is concave (Figs. 1, 7A), probably indicating the anterior
portion of the notch on the premaxillaemaxilla contact for the
reception of an enlarged dentary tooth. This condition is also
present in MOZ 1750 PV, but is widely distributed among croc-
odyliforms including other peirosaurids, sebecosuchians, trem-
atochampsids, and some basal crocodyliforms (e.g., Orthosuchus)
(Nash, 1975; Buffetaut, 1976; Carvalho et al., 2004, 2005; Larsson
Fig. 7. Comparison of A, the premaxillae of Peirosaurus torminni (DGM-433-R) in left and Sues, 2007; Martinelli and Pais, 2008).
lateral view and B, Gasparinisuchus peirosauroides (MOZ 1750 PV) in reversed lateral
view. Scale bar represents 2 cm. Abbreviations: D, dentary. Numbers refer to tooth
position in the premaxilla. The arrow indicates the paracanine depression in both Antorbital fenestra. An antorbital fenestra is present in most peir-
specimens. osaurids (e.g., Lomasuchus, Uberabasuchus, Hamadasuchus;
Gasparini et al., 1991; Carvalho et al., 2004; Larsson and Sues,
2007). Though the region of the antorbital fenestra is damaged in
Montealtosuchus and Hamadasuchus (although slightly reduced in MOZ 1750 PV, its apparent absence was used as a diagnostic feature
this latter taxon), the anteromedial process of the premaxilla is also by Gasparini (1982) and Gasparini et al. (1991). Alternatively, in
well-developed, though not as anteriorly directed as in Peirosaurus. several phylogenetic studies (e.g., Pol and Apesteguía, 2005;
Larsson and Sues, 2007; Turner and Buckley, 2008; Sereno and
Premaxillary teeth. Both Peirosaurus and Gasparinisuchus share a Larsson, 2009; Turner and Sertich, 2010), the specimen MOZ 1750
total of five premaxillary teeth (Figs. 1, 3, 7), a feature widely was scored as missing data for the presence/absence of the antor-
distributed among mesoeucrocodylians including the peirosaurids bital fenestra. Based upon the referred specimen from Mendoza
Uberabasuchus, Montealtosuchus, Hamadasuchus, Stolokrosuchus Province (PV-CRIDC-12), we conclude that in Gasparinisuchus an
(Gasparini et al., 1991; Carvalho et al., 2004; Larsson and Sues, antorbital fenestra is absent. Unfortunately, the presence/absence
2007), in addition to Araripesuchus wegeneri (Sereno and Larsson, of an antorbital fenestra in Peirosaurus cannot be evaluated
2009) and Elosuchus (Lapparent de Broin, 2002). Likewise, the from available material and is therefore useless for comparisons
close-set anteriormost two premaxillary teeth of Gasparinisuchus between taxa.
and Peirosaurus, a feature previously used to unite the two taxa, is
also present in Uberabasuchus (Carvalho et al., 2004). The fourth Maxillary and mandibular teeth. Isolated teeth associated with
premaxillary tooth of Gasparinisuchus is posteroventrally directed, Peirosaurus may pertain to maxillary and/or mandibular teeth. All
whereas in Peirosaurus, all premaxillary teeth are ventrally teeth possess a short and spatulated crown, with flat lingual and
directed. Relative to Peirosaurus, Gasparinisuchus possesses short convex labial surfaces. In lateral view, the crown of unworn teeth
interdental spaces between the premaxillary alveoli (Fig. 3). The has an equilateral triangular shape, whereas in worn teeth, the
widely spaced thirdefifth premaxillary alveoli of Peirosaurus more crown is low and the tip rounded. The crowns of these teeth
closely resemble the condition observed in the Brazilian peir- possess finely denticulate mesial and distal carinae, the crown
osaurids Uberabasuchus and Montealtosuchus. However, the size of distinctly separated from the root by a constricted neck. The roots

Please cite this article in press as: Martinelli, A.G., et al., A new peirosaurid from the Upper Cretaceous of Argentina: Implications for specimens
referred to Peirosaurus torminni Price (Crocodyliformes: Peirosauridae), Cretaceous Research (2012), doi:10.1016/j.cretres.2012.03.017
 A.G. Martinelli et al. / Cretaceous Research xxx (2012) 1e10 9

are variable in size, probably dependent on position. The mesio- directed dorsomedial (internarial) process of the premaxilla, and
distal length of the root is similar or slightly larger than the crown. the sharply defined and elevated margin of the perinarial fossa.
Overall, the teeth of Peirosaurus are similar to the teeth of Loma- Minor differences between the two taxa include differences in the
suchus, Uberabasuchus, Montealtosuchus, Hamadasuchus, and Gas- relative thickness of dorsal osteoderms as noted by Marinho et al.
parinisuchus (Gasparini et al., 1991; Carvalho et al., 2004, 2007; (2006), though the taxonomic value of this feature was not
Larsson and Sues, 2007), all of which possess at least some degree explored widely and may be individually variable. Moreover, the
of variation along the tooth row. absence of a denticulate mesial carina on the first premaxillary
tooth in Peirosaurus is not present Uberabasuchus (Carvalho et al.,
Remarks. Peirosaurus and Gasparinisuchus do not seem to share 2004, p. 981).
unique derived features that warrant their synonymy. Previously Postcranial comparisons between the two taxa would greatly
described traits used to unite Argentinian material here assigned to contribute to this discussion; unfortunately, postcranial materials
Gasparinisuchus with P. torminni (Gasparini, 1982; Gasparini et al., of both specimens are not yet formally described (Vasconcellos,
1991) are now recognized to be widely distributed among peir- 2006). Until this information is available, we tentatively consider
osaurids (e.g., Uberabasuchus, Montealtosuchus, Lomasuchus) and both Peirosaurus and Uberabasuchus valid taxa. Future work is
other mesoeucrocodylians. Moreover, the inference of a sub- clearly necessary to definitively resolve the taxonomy of peir-
triangular, relatively narrow snout in Peirosaurus, together with the osaurids from the Marília Formation of Peirópolis (Minas Gerais
shorter facial length of the premaxilla, reduced smooth perinareal State, Brazil).
region, and shorter interdental spaces among premaxillary teeth in
Gasparinisuchus are features that support the taxonomic distinction 4. Conclusions
between Gasparinisuchus and P. torminni.
A partial peirosaurid specimen (MOZ 1750 PV) from the Upper
3.2. Exclusion of PV-CRIDC-12 from Peirosaurus torminni Cretaceous of Neuquén, Argentina originally referred to the Bra-
zilian taxon Peirosaurus torminni (Gasparini, 1982; Gasparini et al.,
A partial skull and postcranial skeleton (PV-CRIDC-12) from 1991; Praderio et al., 2009), can be morphologically distinguished
Mendoza Province, Argentina was tentatively assigned to Peir- on the basis of its broad and rounded rostrum, anteroposteriorly
osaurus torminni by Praderio et al. (2009) on the basis of similar- short premaxilla, reduced perinareal fossa, and short premaxillary
ities to MOZ 1750 PV. Unfortunately, the premaxilla of PV-CRIDC- interdental spaces. These morphological differences, together
12 is poorly preserved, precluding any meaningful comparison with its unique combination of characters including the absence
with the holotype of P. torminni (DGM-433-R). The preserved of an antorbital fenestra, allow the recognition of MOZ 1750 PV
maxilla of PV-CRIDC-12 shares a number of derived features with as a new taxon, Gasparinisuchus peirosauroides gen. et sp. nov.
Gasparinisucus (MOZ 1750 PV), indicating a tall, broad snout Another peirosaurid specimen from the Upper Cretaceous of
(Fig. 6); maxillary teeth implanted in discrete alveoli (in Loma- Mendoza, Argentina (PV-CRIDC-12) is also referred to
suchus some middle and posterior maxillary alveoli are confluent); Gasparinisuchus.
a convex alveolar maxillary edge at the level of the third tooth; Peirosaurus torminni has long been considered one of the few
and similar position and relative size of the maxillary teeth. specific links between the Late Cretaceous faunas of Argentina and
Specimen PV-CRIDC-12 is here referred to Gasparinisuchus Brazil (e.g., Gasparini et al., 1991, Candeiro and Martinelli, 2006; Pol
peirosauroides. and Gasparini, 2007). Close scrutiny of peirosaurid specimens from
Argentina no longer support the presence of the genus Peirosaurus
3.3. Taxonomic status of Peirosaurus torminni in both countries. However, close phylogenetic relationships
among South American peirosaurids (e.g., Carvalho et al., 2004;
The exclusion of Gasparinisuchus from Peirosaurus leaves the Larsson and Sues, 2007; Leardi and Pol, 2009) and other vertebrate
holotype, DGM-433-R, as the only known specimen of P. torminni. groups (e.g., titanosaurian and abelisaurid dinosaurs) support
However, morphological similarities between DGM-433-R and the observations of general faunal similarities across the Upper Creta-
holotype of Uberabasuchus (CPPLIP 630) suggest possible synonymy ceous of South America. Further resolution of peirosaurid interre-
of the two taxa (briefly discussed by Larsson and Sues, 2007). Both lationships, including taxa from the Cretaceous of mainland Africa
specimens were collected from the same stratigraphic level of the and Madagascar, will undoubtedly contribute substantially to our
Maastrichtian Marília Formation, from localities separated by less understanding of the palaeobiogeographic history of Gondwana
than 1 km (“Ponto 1 do Price” and “Ponto 2 do Price”, respectively). during the Cretaceous.
Though limited to a premaxilla, teeth, and various postcranial Though the composition of Peirosauridae and its phylogenetic
remains (the latter still not formally described), the holotype of position within Mesoeucrocodylia remains controversial (e.g., Pol
P. torminni (DGM-433-R) shares a number of derived features with and Apesteguía, 2005; Gasparini et al., 2006; Carvalho et al.,
Uberabasuchus to the exclusion of other peirosaurids that strongly 2007; Larsson and Sues 2007; Sereno and Larsson, 2009; Sertich,
supports synonymy. 2011), new discoveries from Argentina and Brazil have high-
The holotype of Uberabasuchus terrificus (CPPLIP 630) consists of lighted the ecological and biogeographic significance of peir-
a nearly complete and partially articulated skeleton including osaurids from the Cretaceous of South America. At present, at least
a nearly complete skull (Carvalho et al., 2004; Vasconcellos et al., three peirosaurid taxa are recognized from Argentina [Barcino-
2004; Marinho et al., 2006; Vasconcellos, 2006). The well suchus gradilis (Leardi and Pol, 2009), Lomasuchus palpebrosus
preserved premaxillae of Uberabasuchus permit thorough and (Gasparini et al., 1991), Gasparinisuchus peirosauroides] and five
direct comparisons to Peirosaurus. Though many of the features of from Brazil [Itasuchus jesuinoi (Price, 1955), Montealtosuchus arru-
the premaxilla and dentition of Peirosaurus are widespread among dacamposi (Carvalho et al., 2007), Peirosaurus torminni (Price, 1955),
peirosaurids and many mesoeucrocodylians as noted above, several Pepesuchus deiseae (Campos et al., 2011), Uberabasuchus terrificus
are shared exclusively with Uberabasuchus. Among the features (Carvalho et al., 2004)]. However, the potential synonymy of the
shared by Peirosaurus and Uberabasuchus and not observed in other Brazilian taxa Peirosaurus torminni and Uberabasuchus terrificus
peirosaurid taxa are the relatively long anteroposterior facial length from Peirópolis (Uberaba County, Minas Gerais State) warrants
of the premaxilla, the elongate and strongly anteromedially future investigation.

Please cite this article in press as: Martinelli, A.G., et al., A new peirosaurid from the Upper Cretaceous of Argentina: Implications for specimens
referred to Peirosaurus torminni Price (Crocodyliformes: Peirosauridae), Cretaceous Research (2012), doi:10.1016/j.cretres.2012.03.017
10 A.G. Martinelli et al. / Cretaceous Research xxx (2012) 1e10

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Please cite this article in press as: Martinelli, A.G., et al., A new peirosaurid from the Upper Cretaceous of Argentina: Implications for specimens
referred to Peirosaurus torminni Price (Crocodyliformes: Peirosauridae), Cretaceous Research (2012), doi:10.1016/j.cretres.2012.03.017