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An impressive skeleton of the giant top predator Prestosuchus chiniquensis

(Pseudosuchia: Loricata) from the Triassic of Southern Brazil, with
phylogenetic remarks

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DOI: 10.1080/08912963.2018.1559841

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An impressive skeleton of the giant top predator

Prestosuchus chiniquensis (Pseudosuchia: Loricata)
from the Triassic of Southern Brazil, with
phylogenetic remarks

Lúcio Roberto-Da-Silva, Rodrigo Temp Müller, Marco Aurélio Gallo de França,

Sérgio Furtado Cabreira & Sérgio Dias-Da-Silva

To cite this article: Lúcio Roberto-Da-Silva, Rodrigo Temp Müller, Marco Aurélio Gallo de França,
Sérgio Furtado Cabreira & Sérgio Dias-Da-Silva (2018): An impressive skeleton of the giant top
predator Prestosuchus�chiniquensis (Pseudosuchia: Loricata) from the Triassic of Southern Brazil,
with phylogenetic remarks, Historical Biology, DOI: 10.1080/08912963.2018.1559841

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HISTORICAL BIOLOGY
https://doi.org/10.1080/08912963.2018.1559841

ARTICLE

An impressive skeleton of the giant top predator Prestosuchus chiniquensis

(Pseudosuchia: Loricata) from the Triassic of Southern Brazil, with phylogenetic
remarks
Lúcio Roberto-Da-Silvaa,b, Rodrigo Temp Müller a,c
, Marco Aurélio Gallo de Françad, Sérgio Furtado Cabreirab
and Sérgio Dias-Da-Silvac
a
Programa de Pós-Graduação em Biodiversidade Animal, Universidade Federal de Santa Maria, Santa Maria, Brazil; bLaboratório de Paleontologia,
Universidade Luterana do Brasil, Canoas, Brasil; cCentro de Apoio à Pesquisa Paleontológica da Quarta Colônia, Universidade Federal de Santa
Maria, São João do Polêsine, Brazil; dLaboratório de Paleontologia e Evolução de Petrolina, Campus de Ciências Agrárias, Universidade Federal do
Vale do São Francisco, Petrolina, Brazil

ABSTRACT ARTICLE HISTORY

In the present contribution, we aim to present the osteology of ‘ULBRA-PVT-281’, which comprises the Received 28 August 2018
best-preserved skeleton of Prestosuchus chiniquensis ever found. ULBRA-PVT-281 combines the mor- Accepted 13 December 2018
phology of two classic specimens referred to P. chiniquensis, UFRGS-PV-0156-T and UFRGSPV- 0152-T, KEYWORDS
reunited in a single operational taxonomic unit (OTU) in previous phylogenetic studies. Therefore, the Archosauria; cladistics;
new specimen reinforces the combination of both specimens. We performed a phylogenetic analysis, Paracrocodylomorpha;
combining the information of these three specimens plus braincase data from a fourth specimen, Pinheiros-Chiniquá
UFRGS-PV-0629-T, into a new P. chiniquensis terminal taxon. Moreover, our analysis also included some sequence; ‘Rauisuchia’
new taxa potentially related to P. chiniquensis. As a result, we found a topology slightly distinct from
previous studies, where Ticinosuchus ferox is the basalmost member of Loricata, which also includes the
new combined P. chiniquensis. Our results place P. chiniquensis, Luperasuchus fractus, and Saurosuchus
galilei distributed in a pectinate paraphyletic pattern towards Crocodylomorpha. On the other hand,
a constrained analysis forcing the monophyly of these taxa demands just a single extra step. Therefore,
both scenarios are plausible and agree with the placement of P. chiniquensis within Loricata, whereas
T. ferox nests in Loricata only in the unconstrained analysis.

Introduction the poor knowledge regarding their evolutionary history

(Desojo and Arcucci 2009; Lautenschlager and Desojo
‘Rauisuchia’ is a diverse group of archosaurs with notable
2011; Nesbitt and Deosjo 2017). For instance, there is no
morphological disparity during the Triassic Period (Brusatte
consensus whatsoever if ‘rauisuchians’ are monophyletic,
et al. 2009; Desojo and Arcucci 2009; França et al. 2011;
paraphyletic, or even polyphyletic (Gower 2000; Brusatte
Nesbitt et al. 2013). It comprises huge quadrupedal preda-
et al. 2010; Nesbitt 2011; Nesbitt et al. 2013).
tors (Prestosuchus chiniquensis Huene 1942; Saurosuchus
In South America, the ‘rauisuchian’ record is restricted to
galilei Reig 1959; Fasolasuchus tenax Bonaparte 1981), cur-
Brazil and Argentina. Argentina yields Luperosuchus fractus
sorial bipeds (Poposaurus gracilis Mehl 1915), small edentu-
Romer 1971 (Chañares Formation/Middle-Late Triassic);
lous cursorial bipeds (Shuvosaurus inexpectatus Chatterjee
Sillosuchus longicervix Alcober and Parrish 1997; Saurosuchus
1993 and Effigia okeefeae; Nesbitt and Norell 2006), and
galilei Reig 1959 (both from Ischigualasto Formation), and
partially aquatic animals (Qianosuchus mixtus Li et al.
Fasolasuchus tenax Bonaparte 1981 (Los Colorados Formation).
2006). Moreover, due to their large size, some ‘rauisuchians’
In Brazil, ‘rauisuchians’ are restricted to the Rio Grande do Sul
had become top-tier predators of their ecosystems (e.g.
state in Pinheiros-Chiniquá and Santa Cruz sequences, both
Prestosuchus chiniquensis, Saurosuchus galilei, Postosuchus
included in the Santa Maria Supersequence (Zerfass et al. 2003;
kirkpatricki – Gower and Schoch 2009; Nesbitt et al. 2013).
Horn et al. 2014; Lacerda et al. 2016). In these two units, four valid
Investigation of this group provides important insights
species are so far recorded: Prestosuchus chiniquensis Huene
regarding archosaurian radiation during Mesozoic, as ‘raui-
1942 (Upper Triassic, Dinodontosaurus Assemblage Zone);
suchians’ were widespread in Pangea, with records in almost
Decuriasuchus quartacolonia França et al. 2011 (Upper Triassic,
all continents excepting Antarctica and Australia (Gower
Dinodontosaurus Assemblage Zone); Rauisuchus tiradentes
2000; Brusatte et al. 2010; França et al. 2011; Nesbitt et al.
Huene 1942; Lautenschlager and Rauhut 2014 (Upper Triassic,
2013). However, their phylogenetic relationships are still
Hyperodapedon Assemblage Zone), and Dagasuchus santacruzen-
very controversial. Taxonomic problems and existence of
sis Lacerda et al. 2015 (Upper Triassic, Santacruzodon Assemblage
convergent characters with other archosaurs in part explain

CONTACT Rodrigo Temp Müller rodrigotmuller@hotmail.com Programa de Pós-Graduação em Biodiversidade Animal, Universidade Federal de Santa Maria,
Santa Maria, Brazil
Supplemental data for this article can be accessed here.
© 2018 Informa UK Limited, trading as Taylor & Francis Group

Published online 24 Dec 2018

2 L. ROBERTO-DA-SILVA ET AL.
Zone). Additonally, two Brazilian taxa rely on fragmentary mate-
rials, needing taxonomic revision: Prestosuchus loricatus Huene
1942 and Procerosuchus celer Huene 1942. Since the first report by
Huene 1942, many specimens were collected and ascribed to
Prestosuchus chiniquensis. However, taxonomic problems persist
involving these specimens (Mastrantônio 2010; Raugust 2014;
Lacerda et al. 2015, 2016). According to Desojo (personal com-
munication), a comprehensive review is currently in progress (for
a detailed discussion, see Lacerda et al. 2016). In the present
contribution, we describe one of the most impressive specimens
of Prestosuchus chiniquensi ever found, together with
a phylogenetic analysis carried out in order to test the potential
implications of the new finding on the topology of loricatans.
Institutional abbreviations
BSPG: Bayerische Staatssammlung für Paläontologie und
Geologie, Munich, Germany; CPEZ: Coleção Paleontológica do
Museu Paleontológico e Arqueológico Walter Ilha, São Pedro do
Sul, Brazil; MCN: Museu de Ciências Naturais da Fundação
Zoobotânica do Rio Grande do Sul, Porto Alegre, Brazil; MCP:
Palaeontology Collection of the Museu de Ciências e Tecnologia
of the Pontíficia Universidade do Rio Grande do Sul, Porto Alegre,
Brazil; MCZ: Museum of Comparative Zoology, Harvard,
Cambridge, United States: MSM: Arizona Museum of Natural
History, Mesa, United States; NHMUK: Natural History
Museum, London, United Kingdom; PULR: Museu de Ciencias
Naturales, Universidade Nacional de La Rioja, La Rioja,
Argentina; PVL: Instituto Miguel Lillo, Tucuman, Argentina;
PVSJ: Division of Vertebrate Palaeontology of the Museo de
Ciencias Naturales de la Universidade Nacional de San Juan, San
Juan, Argentina; SMNS: Staatliches Museum für Naturkunde,
Stuttgart, Germany; UFRGS-PV: Paleovertebrate Collection of
the Laboratório de Paleovertebrados da Universidade Federal do
Rio Grande do Sul, Porto Alegre, Brazil; ZPAL: Institute of
Paleobiology, Polish Academy of Science, Warsaw, Poland.
Figure 1. ULBRA-PVT-281 and the location of the study area. (a) map of the Dona Francisca area, Rio Grande do Sul, Brazil, showing the location of the Posto Site
(modified from Müller et al. 2018). (b) ULBRA-PVT-2815 in the rock block. (c), reconstruction of the preserved portions of the skeleton of ULBRA-PVT-281.
Material and methods
Material
The specimen (Figure 1) is housed at the Universidade
Luterana do Brazil under the code ULBRA-PVT-281. It is
partially exposed in a large rock block, which preserves the
following elements: complete and articulated skull, cervical
series, three dorsal and three caudal vertebrae, some dorsal
osteoderms, some dorsal ribs, gastralia, right scapular girdle,
right ulna, and right hindlimb.
Phylogenetic analysis
We performed a phylogenetic analysis in order to investigate the
putative implications of the new specimen on the position of
Prestosuchus chiniquensis within archosaurs. The specimen rein-
forces the combination of UFRGS-PV-0156-T and UFRGS-PV
-0152-T as a unique operational taxonomic unity (OTU).
Historically, both specimens were treated as a unique OTU in
phylogenetic analyses, named as ‘Combined Prestosuchus chini-
quensis’ (see Nesbitt 2011). Although few skeletal elements over-
lap between f UFRGS-PV-0156-T and UFRGS-PV-0152-T,
ULBRA-PVT-281 connects these two specimens and supports
this historical combination. Moreover, the braincase characters
described for UFRGS-PV-0629-T by Mastrantônio et al. (2013)
also were herein included, in the ‘New Combined Prestosuchus
chiniquensis’. This new OTU was inserted in a modified version
of the data matrix of Nesbitt (2011). The modifications
employed by subsequent studies of Butler et al. (2011), Butler
et al. (2014), 2017), Nesbitt and Butler (2012), Nesbitt et al.
(2014), Baczko et al. (2014), Nesbitt and Deosjo (2017) and
Lacerda et al. (2018) were incorporated in the data matrix.
These modifications totalise the inclusion of eight characters
and five taxa (Luperasuchus fractus, Mandasuchus tanyauchen,
Nundasuchus songeaensis, Venaticosuchus rusconii, and
Yonghesuchus sangbiensis). Erpetosuchids were not included in

reason of their behaviour as wildcard taxa (Butler et al. 2014).
Modifications on the characters scores employed by the afore-
mentioned studies were also carried out here. Therefore, the
final data matrix comprises 85 OTUs and 420 characters (see the
Supplemental material). All characters received the same weight
and 19 (32, 52, 75, 121, 137, 139, 156, 168, 188, 198, 223, 247,
258, 269, 271, 291, 297, 328, 356, 377, 399, and 416) were treated
as ordered following the studies of Nesbitt (2011) and Butler
et al. (2014). The phylogenetic analysis was conducted in the
software TNT v1.1 (Goloboff et al. 2008). Mesosuchus browni
was used to root the most parcimonious trees (MPTs), which
were recovered with a ‘Traditional search’ using 1000 replicates,
with 10 Hold, Tree Bisection and Reconnection (TBR) collap-
sing of zero-length branches. In addition, we carried out
a constrained analysis forcing the monophyly of a group com-
posed by Luperasuchus fractus, Prestosuchus chiniquensis, and
Saurosuchus galilei. This analysis was conducted to access the
required number of extra steps to recover this clade, which was
firstly recovered by Nesbitt and Deosjo (2017). The same para-
meters of the unconstrained analysis were employed.
Geological setting
ULBRA-PVT-281 was recovered at the ‘Posto de Gasolina’
outcrop (21°37ʹ38ʹ’S 53°22ʹ07ʹ’W), nearby the main access to
the municipality of Dona Francisca, central region of Rio
Grande do Sul, Brazil (Figure 1(a)). This outcrop is situated
in the Faxinal do Soturno structural block of sedimentary
rocks (Da-Rosa and Faccini 2005) and belongs to the
Pinheiros-Chiniquá Sequence of Horn et al. (2014), charac-
terized by the prevalence of reddish mudstones with subaerial
exposure and carbonate concretions (Rubert and Schultz
2004). The faunal content of the outcrop suggests its inclu-
sion at the Dinodontosaurus Assemblage Zone (França et al.
2013; Pavanatto et al. 2016), biostratigraphically correlated
with the Los Chañares Formation in Argentina, which is
early Carnian in age (Marsicano et al. 2016). The ‘Posto de
Gasolina’ also had yielded other specimens ascribed to
Loricata, including several skeletons of Decuriasuchus quarta-
colonia (França et al. 2011), additional material referred to
Prestosuchus chiniquensis (UFRGSPV-0629-T; Mastrantônio
2010), and isolated elements without a less inclusive taxo-
nomic ascription.
Systematic palaeontology
Archosauria Cope 1869 sensu Nesbitt 2011
Pseudosuchia Zittel 1887 sensu Sereno 2005
Loricata Merrem 1820 sensu Nesbitt 2011
Prestosuchus chiniquensis Huene 1942 (Figs. 1–9)
Lectotype (sensu KREBS 1976)
BSPG 1933L 1–3/5–11/28–41/41 (Excavation 34; Huene 1942)
comprising splenial, anterior portion of the surangular, ante-
rior portion of the angular, prearticular, right partial maxilla,
fragmentary dentary, three incomplete cervical vertebrae,
fragmentary ribs, one sacral vertebra, two sacral ribs, five
anterior caudal vertebrae with chevron bones, 14 middle
HISTORICAL BIOLOGY 3
and posterior caudal vertebrae, both scapulocoracoids, inter-
clavicle and clavicle, distal left humerus, right proximal and
distal humerus, distal radius, fragmentary ulna, one manual
phalanx, an incomplete ilium, fragmentary ischia, pubes and
a complete left hind limb.
Paralectotype (sensu KREBS 1976)
BSPG 1933L/7 (Excavation 41; Huene 1942): an articulated
vertebral sequence, composed of two sacral vertebrae with
sacral ribs, incomplete last dorsal and the first caudal verte-
bra, dorsal portion of the right ilium, a series of osteoderms
articulated with neural spines.
Referred specimens
UFRGS-PV-0156-T, complete skull and partial axial skele-
ton (Barberena 1978); UFRGS-PV-0629-T, almost complete
skull, complete presacral vertebrae sequence, two sacral and
three caudal vertebrae, scapular girdle, partial forelimb, pel-
vic girdle, and partial hindlimb (Mastrantônio 2010);
UFRGS-PV-0152-T, incomplete skull, partial axial skeleton,
vertebral sequence composed of cervical, dorsal, sacral and
caudal elements, scapular girdle, partial forelimb, pelvic
girdle, and partial hindlimb (Nesbitt 2011); CPEZ-239b,
fragmentary and incomplete skeletons of at least two indi-
viduals, which includes cranial and post-cranial elements
(Lacerda et al. 2016).
New referred specimen
ULBRA-PVT-281, complete and articulated skull, cervical
series, three dorsal and three caudal vertebrae, some dorsal
osteoderms, some dorsal ribs, gastralia, right scapular girdle,
right ulna, and right hindlimb.
Diagnosis
Three autapomorphies are described for Prestosuchus chini-
quensis: (i) an anterior notch between the scapula and cor-
acoid (Desojo and Rauhut 2008); (ii) a longitudinal ridge on
the dorsal surface of the ilium (Desojo and Rauhut 2008); and
(iii) presence of a sharp ridge leading from the glenoid to
anteroventral corner of the coracoid (Nesbitt 2011).
Description and comparison
Rock matrix partially encloses ULBRA-PVT-281 with
most bones exposed in right lateral view (Figure 1(b)).
Hence, the present description mainly focuses on this side
of the skeleton. ULBRA-PVT-281 is covered by a hard con-
cretion layer of calcium carbonate and iron oxide, charac-
teristic of many materials found in the Pinheiros-Chiniquá
Sequence. Moreover, most bones of this specimen are articu-
lated, suggesting little or no hydraulic transport.
Skull
Premaxilla. The right premaxilla is lateromedially com-
pressed, with a subquadrangular main body in lateral view
(Figure 2, 3), in that the dorsoventral and anteroposterior
length are subequals, resembling those other specimens of
Prestosuchus chiniquensis (UFRGS-PV-0629-T, UFRGS-PV
-0156-T, BSPG AS XXV 28), (Barberena 1978; Mastrantônio
4 L. ROBERTO-DA-SILVA ET AL.
Figure 2. Skull of Prestosuchus chiniquensis (ULBRA-PVT-281) in right lateral view. Abbreviations: an, angular; d, dentary; j, jugal; l, lacrimal; mx, maxilla; n, nasal;
o, osteoderm; pal, palpebral; pmx, premaxilla; pmxg, premaxillary groove; po, postorbital; q, quadrate; qj quadratojugal; as, surangular; scl, sclerotic ring; sf,
subnarial foramen; sq, squamosal.
2010). In contrast, some basal loricatans this bone is subrec-
tangular as in Decuriasuchus quartacolonia, Polonosuchus
silesiacus, Fasolasuchus tenax, Batrachotomus kupferzellensis,
Postosuchus kirkpatricki, Vivaron haydeni (França et al. 2013;
Lessner et al. 2016). However, intraspecific variations are
recognized for Saurosuchus galilei (either subquadrangular
or subrectangular, see Alcober 2000). The main body of
premaxilla is subrectangular in CPEZ-239b, considered
a juvenile of Prestosuchus chiniquensis (Lacerda et al. 2016),
probably also representing an instraspecific variation or an
ontogenetic variation.
The two dorsal projections of premaxilla (anterodorsal
and posterodorsal processes) are slender, resembling those
of UFRGS-PV-0156-T and UFRGS-PV-0629-T (Lacerda
et al. 2016). In contrast, in Fasolasuchus tenax and
Saurosuchus galilei these processes are dorsoventrally wide
(Bonaparte 1981; Alcober 2000). In ULBRA-PVT-281, both
processes meet the nasal, forming both anterior and poster-
ior margins of the external naris as occurs in Saurosuchus
galilei (PVSJ-32: Alcober 2000). The posteroventral margin
of the posterodorsal process contacts the dorsal process of
the maxilla as in others referred to Prestosuchus chiniquensis,
as CPEZ-239b (Lacerda et al. 2016). This process forms an
angle of 55° in relation to the body of the premaxilla, con-
trasting with 45° observed in Decuriasuchus quartacolonia
(França et al. 2013). The anterodorsal process is semicircular
in cross-section, forming an angle of 45º with main body of
premaxilla. This region on UFRGS-PV156-T is slender, lack-
ing an acute corner.
The anterior margin of the main body of the premaxilla is
straight and vertical (not sinuous). ULBRA-PVT-281 shows
a well-marked groove which starts at the premaxillar-maxillar
suture together with the subnarial foramen, which runs ante-
roventrally to the premaxillary body. Previously, this premax-
illary groove was not described for any other known
specimen of Prestosuchus chiniquensis (Roberto-da-Silva
et al. 2016). However, Prestosuchus chiniquensis specimen
UFRGS-PV-0156-T might present a similar structure, still
undescribed. The premaxillary groove is absent in many
other ‘rauisuchians’, such as Luperosuchus fractus,
 HISTORICAL BIOLOGY 5

Figure 3. Snout of the skull of Prestosuchus chiniquensis (ULBRA-PVT-281). (a) anterior end in right lateral view. (b) anterior end in anterolateral view. (c) antorbital
region in right lateral view. Abbreviations: adp, anterodorsal process; aof, antorbital fenestra; d, dentary; dp, dorsal process; l, lacrimal; mx, maxilla; n, nasal; o,
osteoderm; pdp, posterodorsal process; pmx, premaxilla; sf; subnarial foramen.

Batrachotomus kupferzellensis, Postosuchus kirkpatricki,
Decuriasuchus quartacolonia, and Rauisuchus tiradentes
(Weinbaum 2002, Lautenschlager 2008, Desojo and Arcucci
2009, França et al. 2011, 2013). ULBRA-PVT-281 shares the
premaxillary groove with Saurosuchus galilei (PVSJ 32; PVL
2062). The left premaxilla of UFRGS-PV-156-T bears this
structure, whereas on the right side the presence is not clear.
Maxilla. The maxilla contributes to the ventral, dorsal and ante-
rior margin of the antorbital fenestra in ULBRA-PVT-281 (Figure
2, 3), and resembles that from Saurosuchus galilei (Alcober 2000).
The robust maxilla bears slender rugosities and foramina close to
its ventral margin on lateral surface. In ULBRA-PVT-281,
the posterior process and main body of maxillae lack the
anteroposterior bulbous ridge present on Postosuchus (Long and
Murry 1995, Alcober 2000, Weinbaum 2002) and Rauisuchus
(Lautenschlager & Rauhut 2015). The absence of a ridge is shared
by others specimens of Prestosuchus chiniquensis (UFRGS-PV
-0152-T, UFRGS-PV- 0156-T, UFRGS-PV-0629-T). The maxilla
of ULBRA-PVT-281 does not contribute to the external naris. In
contrast, in Batrachotomus kupferzellensis the maxilla forms part
of narial margin (Gower & Schoch 1999; Lautenschlager and
Rauhut 2014). The anterior margin of maxilla is almost straight
as occurs in UFRGS-PV0156-T. Conversely, in other specimens,
as UFRGS-PV-0629-T and CPEZ-239b, the anterior margin is
slightly angular. The maxillar dorsal process is posterodorsally
displaced and contact the ventral part of the nasal as in CPEZ-
239b (Lacerda et al. 2016). This process contributes to form the
6 L. ROBERTO-DA-SILVA ET AL.
anterodorsal margin of antorbital fenestra. At this point, the
specimen differs from Mandasuchus tanyauchen, which bears an
anteroposterior narrow process (Butler et al. 2017).
The ventral margin of the maxilla is convex on the anterior
region, with sinous morphology on the posterior region. The
shape of the antorbital fossa is subtriangular, with broad
contribution of the dorsal process of maxilla on dorsal region.
Similarly, the antorbital fenestra is subtriangular in shape, but
forming an acute angle on anterior region, just on contact of
dorsal process and main body of maxilla, very similar to other
Prestosuchus chiniquensis specimens (UFRGS-PV-0156-T;
CPEZ-239b). ULBRA-PVT-281 presents a recess in poster-
odorsal margin of antorbital fenestra accompanied by a small
process. However, these structures might also be explained as
artefacts of preservation. The other Prestosuchus chiniquensis
specimens already cited here do not possess these structures.
In addition, the antorbital fenestra in UFRGS-PV156-T is
more dorsoventral narrow than ULBRA-PVT-281, mainly
on the posterior region. The posterior process of maxilla
gently tapers posteriorly to articulate with jugal.
The subnarial foramen. Barberena (1978) did not mention
the presence of a subnarial foramen/fenestra when firstly
describing a complete skull of Prestosuchus chiniquensis.
Further descriptions reported the presence of this structure
in this taxon, but with no information regarding its shape and
position (Mastrantônio 2010, França et al. 2013, Lacerda et al.
2016). Nesbitt (2011) in his extensive phylogenetic analysis of
archosaurs, stated that this structure is absent in Prestosuchus
chiniquensis. Recently, with basis in ULBRA-PVT-281,
a contribution corroborated the presence of subnarial fora-
men in Prestosuchus chiniquensis and discussed its phyloge-
netic significance for Archosauria (Roberto-da-Silva et al.
2016). The subnarial foramen in ULBRA-PVT-281 (Figure
3) is a ‘drop-like’ shaped opening between premaxilla/maxilla
bordered by the main body and dorsal process of the maxilla
as occurs in Saurosuchus galilei (Alcober 2000; Roberto-da-
Silva et al. 2016).
Nasal. The nasal of ULBRA-PVT-281 is an elongated element
comprising about 60% the total length of the skull roof (Figure 2,
4). An enlongated nasal also occurs in Postosuchus kirkpatricki
and other Prestosuchus chiniquensis specimens (UFRGS-PV-
0152-T; UFRGS-PV-0156-T; CEPZ-239b). Its anterior region
bears two processes: anteroventral and posteroventral. The ante-
roventral process contacts the anterodorsal process of the premax-
illa, establishing the dorsal margin of the external naris. The
posteroventral process contacts the posterodorsal process of pre-
maxilla. Dorsally the nasal shows a massive and rugous ridge as
observed in Batrachotomus kupferzellensis (Gower 1999),
Postosuchus kirkipatricki (Weinbaum 2011), Polonosuchus
silesiacus (Brusatte et al. 2009), and Rauisuchus tiradentes
(Lautenschlager and Rauhut 2014). This ridge runs almost entirely
along the nasal body. In Batrachotomus kupferzellensis the ridge is
well-developed. Conversely, it is less prominent in ULBRA-PVT-
281, like others Prestosuchus chiniquensis (UFRGS-PV156-T;
UFRGS-PV0629T; Mastrantônio 2010). In Decuriasuchus quarta-
colonia this ridge is present but appears as a lateral laminar margin
(França et al. 2013). The anterodorsal process is not covered by the
dorsal process of the maxilla and its lateral margin is laminar. The
convexity between the external naris and the antorbital fenestra
named by Romer (1971) as ‘roman nose’ observed in many
‘rauisuchians’ is present in ULBRAPVT-281 and shared with
Decuriasuchus quartacolonia (MCN-PV10.105a), Luperosuchus
fractus (PULR-04), other Prestosuchus chiniquensis (e.g. UFRGS-
PV-0156-T; CEPZ-239), Saurosuchus galilei (PVL-2062; PVSJ-
32), and Riojasuchus tenuisceps (PVL-3827) (Desojo and Arcucci
2009; França et al. 2013; Lacerda et al. 2016; Nesbitt and Deosjo
2017). However, the convexity is more prominent in UFRGS-
PV156-T than ULBRA-PVT-281. In addition, the external naris
of UFRGS-PV156-T extends to the half-length of ‘roman nose’,
whereas in ULBRA-PVT-281 it does not reach the level of ‘roman
nose’. Accordingly, the external naris on ULBRA-PVT-281 is
anteroposterior narrow. Posteriorly, the nasal articulates with
the frontal, prefrontal, and lacrimal.
Lacrimal. This bone shows an inverted L-shape, is laterally
compressed and forms the posterodorsal margin and poster-
ior border of the antorbital fenestra (Figure 2, 4), as in
Saurosuchus galilei and Postosuchus kirkpatricki (Alcober
2000; Weinbaum 2011). The lacrimal is composed of an
anterior ramus and the ventral process. The anterior ramus
is longer than the ventral process, as in Decuriasuchus quar-
tacolonia (França et al. 2013) and other Prestosuchus chini-
quensis specimens (UFRGS-PV-0629-T; UFRGS-PV-0156-
T). In contrast, in Postosuchus kirkpatricki and
Polonosuchus silesiacus the anterior ramus is short (França
et al. 2013). The anterior ramus is laminar, becoming most
robust posteriorly. On the posterior region of anterior ramus
of ULBRA-PVT-281 the dorsal margin of antorbital fenestra
is slightly convex, contrasting with straight margin observed
on UFRGS-PV-0156-T. The lacrimal foramen is not visible
in ULBRA-PVT-281, but described in UFRGS-PV-0629-T
(Mastrantônio 2010). The ventral process is slender, con-
tacts medially the jugal, and composes the posterior margin
of the anterorbital fossa/fenestra as occurs in Decuriasuchus
quartacolonia (França et al. 2013). Ventrally, it contacts the
jugal, on a firm attachment as occurs in other Prestosuchus
chiniquensis specimen (UFRGS-PV-0156-T; Barberena 1978)
and Decuriasuchus quartacolonia (França et al. 2013), con-
trasting with what was described for UFRGS-PV-0629-T and
Saurosuchus galilei (Alcober 2000; Mastrantônio 2010).
Palpebral. ULBRA-PVT-281 possesses a palpebral bone that
forms the orbital roof and prevents the frontal from contri-
buting to the orbital rim (Figures 2, 4). The bone is triangular
and contacts the frontal medially, the prefrontal anterome-
dially, and the postfrontal posteromedially. This bone element
occurs in many ‘rauisuchid’ specimens according to Nesbitt
et al. (2013), such as Postosuchus kirkpatricki, Postosuchus
alisonae, Saurosuchus galilei, Polonosuchus silesiacus, and
Batrachotomus kupferzellensis. More recently, Nesbitt and
Deosjo (2017) described a palpebral bone in Luperosuchus
fractus and Prestosuchus chiniquensis (UFRGS-PV-0156-T).
According to Nesbitt and Deosjo (2017) this element differs
from surrounding bones in reason of the presence of stria-
tions and pits. However, the dorsal surface of the new speci-
men is covered by carbonate concrations, so the pattern of

Figure 4. Orbital region of the skull of Prestosuchus chiniquensis (ULBRA-PVT-281). (a) in right lateral view. (b) in dorsal view. Abbreviations: ar, anterior ramus;
aof, antorbital fenestra; j, jugal; l, lacrimal; mx, maxilla; n, nasal; o, osteoderm; p, parietal; pal, palpebral; pmx, premaxilla; po, postorbital; sq, squamosal; vp,
ventral process.
ornamentation is not visible. The palpebral of Luperosuchus
fractus is completely fused to the frontal and postfrontal
(Nesbitt and Deosjo 2017). Indeed, in ULBRA-PVT-281 the
sutures are poorly visible, but in lateral view the palpebral
present an anteroposterior hump, similar to Saurosuchus
galilei (Nesbitt et al. 2013).
Frontal. This element is covered by the sedimentary matrix
(Figure 4(b)). However, it comprises most of the skull roof
and is excluded from the orbit laterally in response to the
presence of the palpebral.
Postfrontal. The postfrontal is a subrectangular element from
which a small portion contributes to the dorsal border of orbit,
as in other specimens of Prestosuchus chiniquensis (UFRGS-PV
-0629-T; UFRGS-PV-0156-T; CEPZ-239b), Arizonasaurus bab-
bitti, Saurosuchus galilei, and Decuriasuchus quartacolonia
(Alcober 2000; Nesbitt 2005; Mastrantônio 2010; França et al.
2013; Lacerda et al. 2016). In contrast, in Postosuchus kirkpa-
tricki this bone does not reach the lateral margin of the skull
roof (Weinbaum 2011). Anteriorly, the postfrontal is limited
by the frontal, medially by the parietal, and posteriorly by the
postorbital. Moreover, these limits are not visible as described
to Postosuchus kirkpatricki (Weinbaum 2011).
HISTORICAL BIOLOGY 7
Parietal. The parietal constitutes the posterior portion of
skull roof and most of the medial border of the supratemporal
fenestra (Figure 4(b)). This element contacts the frontal ante-
riorly, both postorbital and postfrontal anterolaterally, and
the supraoccipital posteroventrally. Posteriorly, it shows
a vertical crest that posteromedially borders the upper tem-
poral fenestra, it rises in the dorsal margin and continues
ventrally, as in Decuriasuchus quartacolonia, Saurosuchus
galilei, Arizonasaurus babbitti, and Postosuchus kirkpatricki
(Alcober 2000; Nesbitt 2005; Mastrantônio 2010; Weinbaum
2011; França et al. 2013). This sagittal ridge is also visible in
other Prestosuchus chiniquensis specimens (e.g. UFRGS-PV
-0156-T).
Postorbital. The shape of this element is typical of that from
other rauisuchians as (see Weinbaum 2011), in which is ‘T’-
shaped in lateral view, with medial, posterior and ventral
processes (Figure 2, 5(a)). The medial process contacts the
posterior margin of the postfrontal and the anterolateral mar-
gin of the parietal as in UFRGS-PV-0629-T (Mastrantônio
2010). In dorsal view, both posterior and ventral processes
are not visible. The lateral margin of the postorbital shows
a rugose area, a condition shared with UFRGS-PV-0629-T
and Decuriasuchus quartacolonia (Mastrantônio 2010; França
8 L. ROBERTO-DA-SILVA ET AL.

Figure 5. Posterior region of the skull and lower jaw of Prestosuchus chiniquensis (ULBRA-PVT-281). (a) posterior and in right lateral view. (b) detail of the scletorical
ring. (c) right lower jaw in right lateral view. (d) lower jaws in ventral view. Abbreviations: an, angular; d, dentary; hy, hyoid; j, jugal; l, lacrimal; mx, maxilla; o,
osteoderm; pal, palpebral; po, postorbital; q, quadrate; qj, quadratojugal; sa, surangular; scl, sclerotic ring; sp, splenial; sq, squamosal; ul, ulna.

et al. 2013). In contrast, a comparatively denser rugose area is
present in Postosuchus kirkpatricki (Weinbaum 2011) and
Batrachotomus kupferzellensis (SMNS-52970; SMNS-80260)
(Gower 1999; França et al. 2013). The posterior process con-
tacts the squamosal, both constituting the anterodorsal border
of the lower temporal and the lateral border of the upper
temporal fenestra, as in Decuriasuchus quartacolonia and
Saurosuchus galilei (Alcober 2000; França et al. 2013). The
ventral process forms a typical ‘key-hole’ shaped orbit in raui-
suchids, as this structure advances into the orbit, for instance,
in Postosuchus kirkipatricki, Batrachotomus kupferzellensis, and
Saurosuchus galilei (Gower 1999; Alcober 2000; Weinbaum
2011). The ventral process forms the postorbital bar together
with the dorsal process of jugal.
Squamosal. This element comprises four rami constituted by
anterior, posterior, anteroventral, and anteromedial processes
(Gower 1999; Alcober 2000; Mastrantônio 2010). The anterior
process is short and semicircular (Figure 2, 5(a)). In cross-section,
it articulates with the postorbital and forms a bar, which splits the
temporal fenestra in two parts as in both UFRGS-PV-0156-T
and UFRGS-PV-0152-T, Decuriasuchus quartacolonia, and
Saurosuchus galilei (PVSJ-32) (Mastrantônio 2010; França et al.
2013; Raugust 2013). In contrast, a suboval shaped anterior pro-
cess occurs in Batrachotomus kupferzellensis (SMNS-80260) and
Polonosuchus silesiacus (ZPAL/abIII-563), (França et al. 2013).
The squamosal contribution to the posterodorsal margin of
bar is small, as in UFRGS-PV-0629-T (Mastrantônio 2010),
Decuriasuchus quartacolonia (França et al. 2013), and Saur-
osuchus galilei (Alcober 2000). Conversely, a larger contribution
is observed in Postosuchus kirkpatricki (Chatterjee 1985;
Weinbaum 2011) and Batrachotomus kupferzellensis (SMNS-
80260) (França et al. 2013). The lateral surface of the anterior
process does not present any sign of rugosity as occurs in

Postosuchus kirkpatricki (Chatterjee 1985; Weinbaum 2011),
Batrachotomus kupferzellensis (SMNS-80260), and Polonosuchus
silesiacus (ZPAL/abIII-563), as in these taxa a dense rugose ridge is
present (França et al. 2013). The posterior process is short, with
two concavities towards its dorsal surface, and separated by a crest,
as in UFRGS-PV-0629-T and UFRGS-PV-0152-T (Mastrantônio
2010; Raugust 2013). The anteromedial process is vertical and
contacts the lateral margin the occipital process of the parietal.
The anteroventral process bears a ridge that perforates the lower
temporal fenestra, as occurs in Saurosuchus galilei (Nesbitt 2011).
Distinctly, in Postosuchus kirkpatricki, Polonosuchus silesiacus and
Rauisuchus an anterior process on the ventral process perforates,
but also bisects the lower temporal fenestra (Nesbitt 2011).
Jugal. The jugal is a flattened bone formed by the anterior,
dorsal, and posterior processes (Figure 2, 4, 5). Neither ridge
nor swelling occurs on the lateral surface of ULBRA-PVT-281
and other Prestosuchus chiniquensis specimens (UFRGS-PV
-0629-T; UFRGS-PV-0156-T). In contrast, this bone displays
a large swelling on its lateral surface in Postosuchus kirkpatricki
(Chatterjee 1985; Weinbaum 2011). Regarding Decuriasuchus
quartacolonia, it presents the jugal process an antero-posteriorly
directed ridge on its ventral portion of the lateral surface (França
et al. 2013). The anterior process contacts the maxilla anteriorly
and the lacrimal dorsally, contributing to the antorbital fenestra.
The anterior process bears a dorsal blade that articulates to the
lacrimal/prefrontal bar, also contributing with anteroventral
margin of orbit as in the Prestosuchus chiniquensis specimen
UFRGS-PV-0629-T (Mastrantônio 2010). The dorsal process is
long, antero-posteriorly slender and dorsoposteriorly directed.
It also possesses an expanded base that tapers dorsally as in
the specimens UFRGS-PV-0629-T and UFRGS-PV-0156-T
(Mastrantônio 2010). The entire anterior surface of the dorsal
process forms an articular facet for the ventral process of the
postorbital as occurs in Prestosuchus chiniquensis (UFRGS-PV
-0629-T) and Decuriasuchus quartacolonia (França et al. 2013).
This articulation extends along the entire length of the process as
in UFRGS-PV-0629-T, Decuriasuchus quartacolonia, and
Saurosuchus galilei (França et al. 2013). Additionally, in
Batrachotomus kupferzellensis the dorsal process has a higher
participation in the orbital margin (França et al. 2013). The
posterior process is flattened and tapers both dorsally and ven-
trally, as in Saurosuchus galilei (Alcober 2000). Moreover, it is
devoid of a rugose ridge as in Decuriasuchus quartacolonia
(França et al. 2013). In contrast, a rugose ridge is present in
Postosuchus kirkpatricki (Chatterjee 1985; Weinbaum 2011). In
ULBRA-PVT-281 the posterior process is broken and cannot be
located in the ‘expected’ position in which it would contact the
quadratojugal, dorsally overlapping the anterior process of this
bone.
Quadratojugal. The quadratojugal is a ‘V’ shaped element
constituted by an anterior and a dorsal process (Figure 2, 5
(a)). The anterior process articulates to the posterior process
of the jugal forming the posteroventral border of the lower
temporal fenestra. This process forms a 45º angle with the
dorsal process. The dorsal process is a large, plate-like bone as
that of Postosuchus kirkpatricki (Weinbaum 2011). The pro-
cess extends slightly more than the half-length of posterior
HISTORICAL BIOLOGY 9
bar of lower temporal fenestra, like in Saurosuchus galilei
(PVSJ-32). This element posteromedially contacts the dorsal
process of the quadrate and, between the contact areas, the
quadrate foramen should be present, but this structure is not
visible in ULBRA-PVT-281, probably as an artefact of pre-
servation. In another specimen of Prestosuchus chiniquensis
(UFRGS-PV-0629-T) the quadrate foramen is present and in
Postosuchus kirkpatricki and Batrachotomus kupferzellensis
(Gower 1999; Mastrantônio 2010; Weinbaum 2011).
Quadrate. The quadrate is a vertical bone, which possesses
two basic portions: a columnar posterior portion and a plate-
like anterior portion (Figure 2, 5(a)). This morphology is
similar to many other ‘rauisuchians’ as Postosuchus kirkpa-
tricki (Weinbaum 2011), Saurosuchus galilei (Alcober 2000),
and Decuriasuchus quartacolonia (França et al. 2011, 2013).
As usual, the ventral end articulates with the lower jaw. The
dorsal end presents the so-called dorsal head, an attachment
area to the ventral surface of squamosal as in Postosuchus
kirkpatricki (Chatterjee 1985; Weinbaum 2011). Anteriorly,
the quadrate possesses two ridges, one medial and other
ventral. The medial one is not visible in ULBRA-PVT-281
as it is articulated, preventing direct observation, but there is
no reason to assume that it would not be present in the
specimen. Its medial ridge is shorter than the ventral one
and both form a notch that attaches to the dorsal process of
the quadratojugal as in Saurosuchus galilei (Alcober 2000). In
contrast, Decuriasuchus quartacolonia shows a thin blade to
attach the quadratojugal (França et al. 2013). The ventral end
of this element is constituted of two separated condyles due to
the presence of well-marked groove, as in UFRGS-PV-0629-T
(Mastrantônio 2010), Postosuchus kirkpatricki (Chatterjee
1985; Weinbaum 2011), and Decuriasuchus quartacolonia
(França et al. 2011, 2013).
Sclerotic ring. ULBRA-PVT-281 preserves two ossicles from
the sclerotic ring, the first evidence of these bones for
Prestosuchus. (Figure 2, 5(b)). These elements show a plate-
like format and are strongly curvated. These two plates are
articulated and placed dorsally, close to dorsal border of the
orbit. They constituted the sclera, a subunit that contributes
to visual acuity in many vertebrates. The sclerotic ring serves
to accommodate the eye and as an attachment area to ciliary
muscles.
Dentary. The dentary is an antero-posteriorly is a slender and
elongate bone, which mostly forms the mandibular element
(Figure 2, 5). Laterally, it meets the surangular posterodorsally,
the angular posteroventrally, as occurs in UFRGS-PV-
0629-T; UFRGS-PV-0156-T and Decuriasuchus quartacolonia
(Mastrantônio 2010; França et al. 2013). Its ventral margin is
slightly convex as in Decuriasuchus quartacolonia and
Saurosuchus galilei (Alcober 2000; França et al. 2013). In con-
trast, the Prestosuchus chiniquensis specimen BSPG/ASXXV-1,
Batrachotomus kupferzellensis (SMNS-80260), Arganasuchus,
and Arizonasaurus babbitti display a straight margin (Nesbitt
2005; Jalil and Peyer 2007; França et al. 2013). Additionally, in
the ventral margin of the dentary of Postosuchus kirkpatricki
there is a sinuous margin (Chatterjee 1985; Weinbaum 2011).
10 L. ROBERTO-DA-SILVA ET AL.
Surangular. This element is firmly articulated to the articular,
as in Decuriasuchus quartacolonia (MCN-PV10.105a; França
et al. 2013). In ULBRA-PVT-281, it is mediolaterally com-
pressed and visible only in lateral view (Figure 5(c)). It mostly
forms the posterolateral region of the mandible as in
Postosuchus kirkpatricki (Chatterjee 1985; Weinbaum 2011).
The surangular composes the posterodorsal margin of the
mandibular fenestra, as described in UFRGS-PV-0156-T,
UFRGS-PV-0152-T, UFRGS-PV-0629-T, Decuriasuchus quar-
tacolonia, and Postosuchus kirkpatricki (Chatterjee 1985;
Mastrantônio 2010; Weinbaum 2011; Raugust 2014). It con-
tacts the dentary anterodorsally, the angular ventrally, the
articular posteriorly and medially as in Decuriasuchus quarta-
colonia (França et al. 2013). Moreover, it presents an elongated
crest in both lateral and dorsal views, as in Decuriasuchus
quartacolonia and UFRGS-PV-0629-T (Mastrantônio 2010;
França et al. 2013). According to França et al. (2013), this
crest is the so-called lateral shelf of Sampson and Witmer
(2007). In dorsal view, this protuberance also forms a narrow
flat surface as in Decuriasuchus quartacolonia (França et al.
2013). Just below the protuberance, there is a small excavated
area probably to the insertion of M. adductor mandibullae
externus (Sampson and Witmer 2007; França et al. 2013).
Angular. As in many ‘rauisuchians’ like Postosuchus kirkpa-
tricki, Prestosuchus chiniquensis (UFRGS-PV-0156-T;
UFRGS-PV-0629-T), and Decuriasuchus quartacolonia,
(Barberena 1978; Chatterjee 1985; Mastrantônio 2010;
França et al. 2011, 2013, Weinbaum 2011), this element
contributes to the mandibular fenestra (Figure 5(c)).
Anteriorly, the angular contacts the dentary and, poster-
odorsally, the surangular as in Postosuchus kirkpatricki and
Decuriasuchus quartacolonia (Weinbaum 2011; França et al.
2013). Dorsally it meets the surangular as in Postosuchus
kirkpatricki (Weinbaum 2013). The dorsoventrally height of
angular on level of mandibular fenestra is slightly more
robust in ULBRA-PVT-281 than UFRGS-PV-0156-T.
Articular. The articular is a robust and shows five distinct
articular projections and forms the largest area of the
retroarticular process, as in Postosuchus kirkpatricki
(Chatterjee 1985; Weinbaum 2011), Decuriasuchus quarta-
colonia (França et al. 2011, 2013), and other specimens of
Prestosuchus chiniquensis (UFRGS-PV-0629-T, UFRGS-PV
-0152-T) (Mastrantônio 2010; Raugust 2014). Many details
are not visible due to the fact that mandible is attached to
the skull. However, its dorsal face shows two glenoids that
contact the correspondent two condyles from the ventral,
posterior end of the quadrate, as in Rauisuchus tiradentes
(Lautenschlager and Rauhut 2014), Decuriasuchus quarta-
colonia (França et al. 2011, 2013), UFRGS-PV-0629-T, and
UFRGS-PV-0152-T (Mastrantônio 2010; Raugust 2014).
Hyoids. To this date, the hyoid apparatus was unknown in
Prestosuchus chinquensis, this becomes the first report and
description of this structure. The right hyoid element is pre-
served in ULBRA-PVT-281, but only its most caudal region
portion is visible in ventral view (Figure 5(d)). This element is
long and slightly curved. Its corpus shows a ‘rod-like’ shape as
in Decuriasuchus quartacolonia and Qianosuchus mixtus (Li
et al. 2006; França et al. 2013). In addition, its morphology is
similar to the dinosaurs Syntarsus kayentakatae and
Coelophysis bauri (Tykoski 1998). Unfortunately, the relation
of hyoid and mandibular length is not visible.
Dentition. Three premaxillary teeth are preserved in right
side of ULBRA-PVT-281 (Figure 5(c)), plus one empty
alveolus, making four tooth positions on this bone.
These elements have relatively long alveoli, curved and
slightly cylindrical in comparison with the maxillary
teeth and comparatively longer than those of Rauisuchus
tiradentes (Lautenschlager and Rauhut 2014). The pre-
sence of four premaxillary teeth is also present on other
specimen referred to Prestosuchus chiniquensis (UFRGS-
PV-0629-T; UFRGS-PV-0156-T; CPEZ-239b), and shared
with several loricatans, such as Saurosuchus, Fasolasuchus,
Batrachotomus, Postosuchus, Polonosuchus, Rauisuchus
(Barberena 1978; Bonaparte 1981; Long and Murry 1995;
Gower 1999; Weinbaum 2002; Sulej 2005; Mastrantônio
2010; Lautenschlager & Rauhut 2015; Lacerda et al. 2016).
Distinctly, Vivaron haydeni shows five alveoli, a condition
shared with basal crocodylomorphs (Nesbitt 2011; Lessner
et al. 2016). The medial surface on ULBRA -PVT-281 is
not visible because the lower jaw is in occlusion, preclud-
ing morphological description of the medial surface, inter-
dental plates and palatal process of premaxilla/maxilla.
The maxilla bears eight teeth plus three alveoli, compris-
ing 11 dental elements (Figure 5(c)). These teeth are
compressed, posteriorly curved and present a gentle serra-
tion. The first three are very large compared to the
remaining ones that become gradually smaller. In other
Prestosuchus chiniquensis specimens, the number of
alveoli is variable. As stated, at least 11 are present in
UFRGS-PV-0156-T whereas 13 are present in UFRGS-PV
-0629-T (Barberena 1978; Mastrantônio 2010). This con-
trasts with Decuriasuchus quartacolonia in which at least
17 alveoli are observed (França et al. 2011). Saurosuchus
galilei shows the same number of the 11 teeth seen in
ULBRA-PVT-281.
Axial skeleton
Cervical vertebrae. ULBRA-PVT-281 shows a series of eight
vertebrae preserved and articulated with their adjacent osteo-
derms (Figure 1(b), 6(a)). Following the last cervical element,
there are three transitional vertebrae between cervical and
dorsal series. Unfortunately, their preservation state prevents
a detailed description. Osteoderms are dorsally articulated to
their respective neural spines. Moreover, at least two cervical
ribs are placed close to their original articulation with the C6
(?) and C7 (?).
The axis is preserved but partially obscured by the skull.
The thin neural spine is laterally exposed, with the dorsal
edge convex as in Rauisuchus tiradentes and Polonosuchus
silesiacus (Gower and Schoch 2009). In contrast, in
Batrachotomus kupferzellensis and Fasolasuchus tenax the
dorsal edge is concave (Bonaparte 1981; Gower and Schoch
2009). Unfortunately, the centrum and the intercentrum,
prezygapophyses and postzygapophyzes cannot be observed

Figure 6. Axial elements of Prestosuchus chiniquensis (ULBRA-PVT-281). (a), cervical series in right lateral view. (b), two mid-dorsal vertebrae in left lateral view; (c),
dorsal ribs, caudal vertebrae, and gastralia associated. Abbreviations: c, centrum; dp, diapophysis; g, gastralia; ifl, infradiapophyseal lamina; na, neural arch; ns, neural
spine; o, osteoderm; pp, parapophysis; pz, prezygapophysis; rb, rib.
in ULBRA-PVT-281. In C3 (?), C4 (?) and C5 (?), the neural
spines are partially visible. The centra of these elements are
subrectangular in lateral view. The distal end of neural spines
is poorly expanded transversally, with a slight T-shaped
expansion, as in Saurosuchus galilei and Batrachotomus kup-
ferzellensis (Gower and Schoch 2009; Trotteyn et al. 2011).
Conversely, in Postosuchus kirkpatricki this expansion is dis-
tinctly “heart-shaped‟ (Weinbaum 2013). No hyposphene-
hypantrum, prezygapophyseal and postzygapophyseal lami-
nae are visible, but the lateral fossa and the ventral keel are
present in the centra. The parapophysis is anteroventrally
placed in the vertebral centrum and the diapophysis above
the distal end of the transverse process. Moreover, this trans-
verse process is ventrally oriented. The transverse process is
short and increases in size posteriorly. The diapophyses and
parapophyses are placed in the limit between the centrum and
HISTORICAL BIOLOGY 11
the neural arch as in UFRGS-PV-0629-T (Mastrantônio
2010). The direct observation of the cervical centrum is not
possible but, given its proportions we infer they are laterally
constricted and anterior-posteriorly shorter than middle dor-
sal centrum.
Cervical ribs. Two right cervical ribs are preserved in
ULBRA-PVT-281 (Figure 6(a)). The largest one is at least
two times longer than the centrum, resembling the fifth
cervical rib of Postosuchus alisonae (Peyer et al. 2008). The
smaller one is short and consequently, the distance between
the articular surfaces is shorter. Both cervical ribs show
flattened and relatively wide articular surfaces, resembling
those ofBatrachotomus kupferzellensis (Gower and Schoch
2009). In contrast, the articular surfaces of the cervical ribs
of Postosuchus alisonae are circular in outline (Peyer et al.
12 L. ROBERTO-DA-SILVA ET AL.
2008). The capitulum and tuberculum of cervical ribs
tapers just after their articular surfaces, resulting in a very
sharp distal end, as in UFRGS-PV-0629-T (Mastrantônio
2010). An accessory head in the cervical ribs is reported for
Batrachotomus kupferzellensis (Gower and Schoch 2009),
a feature absent in the preserved elements of ULBRA-
PVT-281and other Prestosuchus chiniquensis specimens
(UFRGS-PV-0629-T; UFRGS-PV-0152-T; Mastrantônio
2010; Raugust 2014).
Dorsal vertebrae. Six elements are preserved in ULBRA-
PVT-281 (Figure 1(b), 6(b)): one set of three vertebrae
articulated to the posterior element from the cervical series,
therefore being transitional (Mastrantônio 2010), called
similar dorsal elements articulated with the cervical series
(‘transitional’) and three posterior articulated, but isolated
from the first set. In the posterior elements, the parapophy-
sis is inserted on the transverse process, which is laterally
oriented, as in UFRGS-PV-0629-T and Batrachotomus kup-
ferzellensis (Gower and Schoch 2009; Mastrantônio 2010). In
lateral view, the diapophyses are laterally oriented and
placed on the transverse process, whereas the parapophysis
reaches the height of the neural canal, as in Saurosuchus
galilei (Trotteyn et al. 2011). A paradiapophyseal lamina lies
between the diapophysis and parapophysis, resembling
Postosuchus alisonae (Peyer et al. 2008). The centra of mid-
dle dorsal elements of ULBRA-PVT-281 are amphicoelous
and quadrangular as in UFRGS-PV-0152-T (Raugust 2014).
A well-marked lateral fossa is present in the lateral surface
of the centrum as in Batrachotomus kupferzellensis,
Postosuchus kirkpatricki, Postosuchus alisonae, Rauisuchus
tiradentes, and other Prestosuchus chiniquensis (UFRGS-PV
0629T, UFRGS-PV0152T), (Peyer et al. 2008; Gower and
Schoch 2009; Mastrantônio 2010; Weinbaum 2011;
Lautenschlager and Rauhut 2014; Raugust 2014). In ventral
view, the centrum of ULBRA-PVT-281 is devoid of a keel,
which resembles Batrachotomus kupferzellensis, Postosuchus
kirkpatricki, Postosuchus alisonae, Rauisuchus tiradentes,
and UFRGS-PV0629T, UFRGS-PV0152T (Peyer et al. 2008;
Gower and Schoch 2009; Mastrantônio 2010; Weinbaum
2011; Lautenschlager and Rauhut 2014; Raugust 2014). The
neural spines are not visible, as the scapula conceals them.
Dorsal ribs. Nine dorsal ribs are preserved (Figure 5(c)). The
first one is placed near to the articulation with the last
‘transitional’ vertebra. Strangely, this rib folds under the sca-
pula and its distal end enfolds the centrum of a dorsal verte-
bra positioned below. This is indicative that probably all
cervical region and skull suffered some degree of distortion
before or during the diagenetic process. This element is
slightly flattened close to the proximal end and becomes
cylindrical toward to distal end. The remaining ribs are asso-
ciated with the gastralia basket, but some elements are not
complete. Two ribs are completely preserved. Their morphol-
ogy is very similar to that from Batrachotomus kupferzellensis
(Gower and Schoch 2009). The flattened part corresponds to
the tuberculum and a rounded capitulum is present, as in
Postosuchus alisonae and Batrachotomus kupferzellensis
(Peyer et al. 2008; Gower and Schoch 2009).
Caudal vertebrae. Three caudal elements are present in
ULBRA-PVT-281 (Figure 6(c)). They are overlapped by dor-
sal ribs and a layer of concretion. Only one is complete,
whereas the other two elements are composed solely by the
centrum. All centra are spool-shaped. In lateral view, with no
signs of fossa. Conversely, in Prestosuchus chiniquensis (BSPG
1933L/7) and Batrachotomus kupferzellensis a lateral fossa is
evident (Gower and Schoch 2009).
Gastralia. Placed in the ventral surface of the trunk, ULBRA-
PVT-281 presents parts of the gastralia basket (Figure 6(c)).
These elements are delicate and slender in ULBRA-PVT-281,
as seen in Postosuchus kirkpatricki (Weinbaum 2013). Only
a few pairs of gastralia are preserved over a sequence of dorsal
vertebrae and ribs. These elements are cylindrical and show
a well-marked curvature on their distal end as observed in
Postosuchus alisonae (Peyer et al. 2008). The observed curva-
ture serves to meet the distal ends of the thoracic ribs. Its
posterior end is flattened as in Postosuchus kirkpatricki
(Weinbaum 2013). According to Peyer et al. (2008) the ‘gas-
tralian’ morphology of Postosuchus alisonae resembles the one
of Ticinosuchus ferox, being similar to ULBRA-PVT-281.
Osteoderms. Only one paramedian osteoderm plus those in
association with the cervical series are preserved in ULBRA-
PVT-281. The paramedian osteoderm is displaced, being
located on the jugal bone. It shows two districts faces:
a ‘plate-like’ and a ventrolateral, which forms an angle,
resembling Batrachotomus kupferzellensis (Gower and
Schoch 2009). The paramedian osteoderm is leaf-shaped
similar in both form and size as that of Batrachotomus
kupferzellensis and UFRGS-PV-0629-T (Mastrantônio
2010) (Lautenschlager and Desojo 2011). The remaining
preserved osteoderms change their morphology throughout
the cervical series. Unfortunately, a diagenetic carbonate
layer covered the dorsal surface of the paramedian osteo-
derm preventing accessing its pattern of ornamentation. In
UFRGS-PV-0629-T (Mastrantônio 2010) the ornamentation
shows a radial pattern, constituted of lower ridges that begin
in a dorsal protuberance, resembling Saurosuchus galilei
(Trotteyn et al. 2015). At this point, there is no reason to
assume that, in the paramedian element of ULBRA-PVT
-281, ornamentation would be different. The cervical osteo-
derms are wider than long and show a slight morphologic
transition among the series, becoming wider close to the
dorsal series. These osteoderms are arranged in pairs and
imbricated to each other, being two pairs anteroposteriorly
displaced for each vertebral element. Their morphology is
different from that present in Postosuchus alisonae and
Rauisuchus tiradentes which are longer than wide (Peyer
et al. 2008). The ornamentation is similar to that from
UFRGS-PV-0629-T (Mastrantônio 2010).
Appendicular skeleton
Scapula. The right scapula is preserved in ULBRA-PVT-281
(Figure 7). It is dorsally convex, ventrally concave, elongated,
and narrow. Its central region presents a constriction as in
UFRGS-PV-0629-T, Rauisuchus tiradentes, Batrachotomus kup-
ferzellensis, and Postosuchus kirkpatricki (Gower and Schoch

2009; Mastrantônio 2010; Weinbaum 2013; Lautenschlager and
Rauhut 2014). The dorsal expansion end is similar on compar-
ison with other specimens of Prestosuchus chiniquensis
(UFRGS-PV-0629-T), although it is more expanded than
UFRGS-PV-0152-T. Additionally, in Batrachotomus kupferze-
lensis (Gower and Schoch 2009) the dorsal expansion is more
marked than in specimens of Prestosuchus chiniquensis. This
contrast on expansion length might result from the large size
of ULBRA-PVT-281 or intraspecific variation. The ventral end
of scapula forms the glenoid, which becomes thinner dorsally as
in Rauisuchus tiradentes (Lautenschlager and Rauhut 2014),
with posterolateral orientation. The scapula is firmly attached
to the coracoid and their suture is well visible as in Postosuchus
alisonae, Postosuchus kirkpatricki, and Rauisuchus tiradentes
(Peyer et al. 2008; Weinbaum 2013; Lautenschlager and
Rauhut 2014). Additionally, in Postosuchus kirkpatricki (TTU-
P9002; TTU-P9000) a firm contact between these two elements
is also mentioned, but there is no fusion due to the juvenile
ontogenetic stages in these specimens (Weinbaum 2013).
ULBRA-PVT-281 is apparently a fully grown mature individual
and there is no sign of fusion between scapula and coracoid,
similarly to the condition found in Batrachotomus kupferzellen-
sis (Gower and Schoch 2009). A notch between those bones
marks the posterior margin of this articulation. In lateral view,
the scapula shows a convex surface along its total length as in
Rauisuchus tiradentes (Lautenschlager and Rauhut 2014). Close
to the posteroventral margin, there is a tubercle placed dorsally
to the supraglenoid lip, which attaches the triceps muscle
(Mastrantônio 2010). This structure is elongated laterally and
forms a slightly concave area lateroventrally, which is different
from other specimens of Prestosuchus chiniquensis specimens
(UFRGS-PV0629-T; and BSPHG AS 1933L-12- see Huene 1942;
Mastrantônio 2010). In Prestosuchus chiniquensis UFRGS-PV
0629-T and BSPHG AS 1933L-12, this tubercle is poorly promi-
nent in comparison to ULBRA-PVT-281. In contrast,
Arizonasaurus babbitti presents a suboval tubercle with slender
rugosities (Nesbitt 2005). Additionally, the format of this tuber-
cle in ULBRA-PVT-281 resembles that from Rauisuchus tira-
dentes (Lautenschlager and Rauhut 2014). Distally, the scapula
shows a slightly prominent acromial process, as in UFRGS-PV
0629-T (Mastrantônio 2010) and Batrachotomus kupferzellensis
(Gower and Schoch 2009), which serves as a point of attachment
to the clavicle (Romer 1956). In ULBRA-PVT-281, both clavicle
and interclavicle are associated to the scapula in almost the
actual expected position. The ventral end of the scapula bears
the articular facet to the coracoid. Anteriorly, it is slender as in
BSPHG AS 1933L-12 and Rauisuchus tiradentes (BSPG AS XXV
91) (Mastrantônio 2010, Lautenschlager and Rauhut 2014).
Posterodorsally, the ventral end contributes to most part of
glenoid fossa. Dorsally to glenoid fossa, there is a well-marked
supraglenoid lip as in UFRGS-PV0629-T (Mastrantônio 2010).
Coracoid. The coracoid is subrectangular and about twice longer
than tall (Figure 7), what is observed in UFRGS-PV0629-T
(Mastrantônio 2010). Conversely, in Postosuchus alisoneae and
Postosuchus kirkpatricki the coracoid is more than twice taller than
long (Peyer et al. 2008; Weinbaum 2013). Its lateral surface is
convex and medially concave as in Batrachotomus kupferzellensis
(Gower and Schoch 2009). The coracoidal foramen is placed
HISTORICAL BIOLOGY 13
within the coracoid corpus as in several specimens of
Prestosuchus chiniquensis (UFRGS-PV0152-T; UFRGS-PV0629-
T; BSPHG AS 1933L-12), Batrachotomus kupferzellensis and
Postosuchus alisonae (Peyer et al. 2008; Gower and Schoch 2009;
Mastrantônio 2010; Raugust 2014). The coracoidal foramen serves
for the passage of the supracoracoid nerve as reported for
Postosuchus alisonae and Batrachotomus kupferzellensis (Peyer
et al. 2008; Gower and Schoch 2009). The coracoid forms most
part of the glenoid cavity as in UFRGS-0629-T, Postosuchus aliso-
nae, and Batrachotomus kupferzellensis (Peyer et al. 2008; Gower
and Schoch 2009; Mastrantônio 2010). In UFRGS-PV0629-T and
Batrachotomus kupferezellenis, the coracoid surface of glenoid
shows two discrete areas. However, these structures are not visible
in ULBRA-PVT-281. Moreover, a fracture crosses the coracoid
antero-posteriorly, dorsally to the coracoid foramen. Ventrally to
the coracoid foramen there is a crest that starts at the poster-
oventral margin and protrudes anterodorsally (Figure 7(c)) as in
many specimens of Prestosuchus chiniquensis (UFRGS-PV-0629-
T; UFRGS-PV-0152-T; BSPHG AS 1933L-12), (Mastrantônio
2010; Raugust 2014). According to Nesbitt (2011), this crest is
an autapomorphy of Prestosuchus chiniquensis. However, accord-
ing to Raugust (2014), the same structure is present in
Procerosuchus celer (BSPHG AS 1933L-34) and Batrachotomus
kupferzellensis (SMNS 80271). Unfortunately, the coracoid is not
preserved in Saurosuchus galilei.
Interclavicle. The interclavicle is a blade-like shaped element
placed between the coracoids, as in Postosuchus alisonae (Peyer
et al. 2008). The shaft is ventrally convex and its anterior end
shows a slight expansion that accommodates two articular facets
for the clavicles, as in other Prestosuchus chiniquensis specimens
(UFRGS-PV-0629-T, Mastrantônio 2010; BSPG1933L, Huene
1942) and in the aetosaur Polesinesuchus aurelioi (Roberto-da-
Silva et al. 2013). In ULBRA-PVT-281, the interclavicle is in
articulation with both clavicles. Distally, the body of interclavicle
is more expanded towards its distal end. Some striations are
visible along the body, as in Postosuchus alisonae (Peyer et al.
2008).
Clavicle. ULBRA-PVT-281 preserves both clavicles (Figure 7
(a)). They are rod-like shaped as in Postosuchus alisonae and
Prestosuchus chiniquensis (BSPG1933L, Huene 1942). The
right element is more visible, it is expanded, with both ends
almost the same proportion. There is a fracture across the
midline shaft, but not displaced from the original position.
The distal end contacts the anterodorsal margin of the sca-
pula close to its distal end. The left clavicle is less visible as its
distal end is covered by a cervical spine.
Ulna. Solely the proximal end of the ulna is preserved in
ULBRA-PVT-281 and this element is poorly visible as it is
covered by the skull (Figure 5(d)). Its articular surface to the
humerus is ‘comma-shaped’ and relatively slender than both
tibia and fibula from this specimen.
Femur. The right femur of ULBRA-PVT-281 is a strong bone
(1.72 times the tibial length) that presents a slightly sigmoidal
shape (Figure 8) as in UFRGS-PV0629-T and Batrachotomus
kupferzellensis (SMNS 52970; Gower and Schoch 2009;
14 L. ROBERTO-DA-SILVA ET AL.

Figure 7. Pectoral girdle of Prestosuchus chiniquensis (ULBRA-PVT-281). (a), right scapula and coracoid in lateral view. (b), right scapula and coracoid in posterior
view. (c), right coracoid in lateral view. Abbreviations: an, anterior notch; cf, coracoid foramen; cl, clavicle; co, coracoid; gf, glenoid; rdg, ridge; sc, scapule; sgl,
supraglenoid lip.

Mastrantônio 2010). The distal end is slightly downturned. The
femoral head is not prominent and slightly displaced medially in
order to connect to the acetabulum, and there is no sign of
a neck connecting to the shaft as occurs in Postosuchus kirkpa-
tricki and Arganasuchus dutuiti (Jalil and Peyer 2007;
Weinbaum 2013). In contrast, in Argentinian Fasolasuchus
tenax and Saurosuchus galilei, a neck is present right below the
femoral head (Bonaparte 1981; Jalil and Peyer 2007; Weinbaum
2013). Moreover, ULBRA-PVT-281 possesses the femoral shaft
columnar, differently than observed in Fasolasuchus tenax, in
which the femoral shaft is slender in comparison. Laterally, the
great trochanter is present. A small and rounded anteromedial
tuber is distally placed from the femoral head in ULBRA-PVT
-281, as in UFRGS-PV0152T (Nesbitt 2011; Raugust 2014). In
the posterior face, a well-marked tuber is present as in UFRGS-
PV0629T (Mastrantônio 2010). The fourth trochanter in
ULBRA-PVT-281 is little prominent, being a slight protuber-
ance dorsoventrally oriented, resembling Batrachotomus kupfer-
zellensis and UFRGS-PV0152T (Gower and Schoch 2009;
Raugust 2014). In contrast, it is a low, elongate, knob-shaped
structure in Postosuchus kirkpatricki, Saurosuchus galilei, and
Hesperosuchus agilis (Weinbaum 2013). The fourth trochanter
of Fasolasuchus tenax is more pronounced than in ULBRA-PVT
-281. This trochanter is mentioned as the insertion area for the
caudofemoralis muscle (Jalil and Peyer 2007). Additionally, the
femur of Fasolasuchus tenax is comparatively less robust and

Figure 8. Right femur of Prestosuchus chiniquensis (ULBRA-PVT-281) in (a), anterior (b), posterior, (c), lateral, (d), medial (e), proximal, and (f), distal views.
Abbreviations: 4t, fourth trochanter; ctf, crista tibiofibular; fc, fibular condyle; g, groove; gt, great trochanter; lc, lateral condyle; mc, medial condyle; pf, popliteal
fossa; pmt, posteromedial tuber.
sigmoid. The distal end bears three condyles. There is a popliteal
fossa between the medial condile and the crista tibiofibularis.
Tibia. It is a robust element (Figure 9) in which the
proximal end is more expanded than the distal one, as
in Postosuchus kirkpatricki and Batrachotomus kupferzel-
lensis (Gower and Schoch 2009; Weinbaum 2013). The
tibial shaft is a columnar and transversally cylindrical
structure, resembling that from other specimens of
Prestosuchus chiniquensis (UFRGS-PV-0629-T; BSPHG
AS 1933L-12) and Batrachotomus kupferzellensis, (Huene
1942; Gower and Schoch 2009; Mastrantônio 2010). The
proximal end shows two concave areas for articulation
with the femur. Close to the proximal end, medially,
there is a deep excavation (pit) perhaps for insertion of
the puboischiotibialis muscle, as occurs in UFRGS-PV
-0629-T and Batrachotomus kupferzellensis (Gower and
Schoch 2009; Mastrantônio 2010; Raugust 2014). The dis-
tal end possesses a dorsoventrally orientated notch
between the articulation from astragalus and the ventro-
medial margin as in UFRGS-PV-0152-T (Raugust 2014).
Fibula. The fibula is articulated to the tibia in ULBRA-
PVT-281, and both have the same length (Figure 9). This is
an elongate and slightly sigmoidal bone as in UFRGS-PV
-0629-T and Postosuchus alisonae (Peyer et al. 2008;
Mastrantônio 2010). Its proximal end poorly visible, as it
is partially covered by the tibia. The shaft is not expanded
as that from Batrachotomus kupferzellensis (Gower and
Schoch 2009). Its articulation with the femur is concave
HISTORICAL BIOLOGY 15
and anteroposteriorly elongate as in UFRGS-PV-0629-T
and Batrachotomus kupferzellensis (Gower and Schoch
2009; Mastrantônio 2010). Unfortunately, the region of
trochanter for the iliofibularis muscle is not visible in
ULBRA-PVT-281.
Pes. The right posterior autopodium of Prestosuchus chini-
quensis (ULBRA-PVT-281) is partially preserved and articu-
lated (Figure 9). The pes is plantigrade, almost symmetrical
and possesses a prominent calcaneal tuber as in Postosuchus
alisonae (Peyer et al. 2008). Four metatarsals are observed in
ULBRA-PVT-281 and only the metatarsal V is not preserved.
These elements are very similar to that of another specimen
of Prestosuchus chiniquensis (BSPHG AS 1933L 11) being the
metatarsal IV more robust as in Postosuchus kirkpatricki
(Chatterjee 1985; Weinbaum 2013). The element I is more
gracile compared to IV, whereas I and II show intermediary
condition. The metatarsal III, together with IV are more
elongated. In all elements both proximal and distal ends are
very expanded and turn inward towards the fourth distal
tarsal, resembling Postosuchus kirkpatricki and Postosuchus
alisonae (Chatterjee 1985; Peyer et al. 2008; Weinbaum
2013). A separate description of both astragalus and calca-
neum is provided below.
Astragalus. Visible in posteroventral view (Figure 9), the
right astragalus is preserved in close association with the
calcaneum, but not articulated. Their articulation surface is
a typical crocodile normal (sensu Chatterjee 1982), found in
pseudosuchians, in which the astragalus projects a lateral
16 L. ROBERTO-DA-SILVA ET AL.
Figure 9. Right Tibia, Fibula and pes of Prestosuchus chiniquensis (ULBRA-PVT-281) in medial view. Abbreviations: as, astragalus; cal, calcaneum; f, fibula; ph,
phalanges; t, tibia; mt, metatarsal.
peg which fits into the socket of the calcaneum so that there
is a rotating ankle joint between these two elements
(Weinbaum 2013). Also, the astragalus is robust and bears
two articulation facets (fibial and tibial), as in Postosuchus
kirkpatricki (Weinbaum 2013). In ULBRA-PVT-281 the size
of the astragalus matches that of the calcaneum. In contrast,
the astragalus of the Fasolasuchus tenax is smaller than the
calcaneum (Bonaparte 1981). The tibial facet is visible pos-
teriorly and its surface is slightly concave. In contrast,
Effigia okeefeae shows a well-marked concave tibial facet
(Nesbitt 2007). This facet is divided into two articular
diverging surfaces separated by a ridge, resembling that of
Saurosuchus galilei and Fasolasuchus tenax (Lecuona and
Desojo 2011), the so-called screw joint articulation to the
tibia, typical for Pseudosuchia (Lecuona and Desojo 2011).
In anterior view, the astragalus shows a smooth and con-
cave surface of articulation to metatarsals I and II (roller
sensu Nesbitt 2011) as in Fasolasuchus tenax (Bonaparte
1981).
Calcaneum. This element shows a typical ventral articulation
with the astragalus (Figure 9), constituted of a deep socket
supported from a dorsally concave lip (Martz 2002). Overall,
the calcaneum of Prestosuchus chiniquensis (Huene 1942;
Raugust 2014; ULBRA-PVT-281) resembles that from
Batrachotomus kupferzellensis (Gower and Schoch 2009).
Many details are not visible due to the presence of a layer of
concretion covering this element, as also occurs with the astra-
galus. In posterodistal view, the calcaneum shows a proximal
rolling with two articular facets to fibula and to the fourth
tarsal distal as in Postosuchus kirkpatricki (Chatterjee 1985;
Weinbaum 2013). Medially to the calcaneal condyle, there is
a deep calcaneal socket inside a concave surface of the distal
astragalar facet, as in the aetosaur Polesinesuchus aurelioi
(Roberto-da-Silva et al. 2014).
Results of the phylogenetic analysis
The phylogenetic analysis recovered 360 MPTs of 1395 steps
(Consistency Index = 0.353; Retention Index = 0.761). In all
MPTs, the new Prestosuchus chiniquensis lies as an early diver-
ging member of Loricata (Figure 10). However, distinct from
other studies (e.g. Nesbitt 2011; Roberto-da-Silva et al. 2016),
Prestosuchus chiniquensis does not nest as the basalmost mem-
ber of the group in any MPT. Actually, Ticinosuchus ferox
occupies the basalmost branch of Loricata in the strict consensus
tree. This clade is supported by three synapomorphies: (i) nearly
pointed anterior margin of the antorbital fenestra [character 30
(state 1)]; (ii) centra of the cervical vertebrae 3 to 5 shorter or
with the same length of the mid-dorsal [181(0)]; and (iii) outline
of the distal end of the ischium rounded or elliptical [293(1)].
Despite the position of Ticinosuchus ferox as the basalmost
member of Loricata in the strict consensus tree (Figure 9),
Mandasuchus tanyauchen occupies this position in some of the
MPTs. However, the same taxon also lies as the sister taxon of
Paracrocodylomoprha in several MPTs. In contrast, with the
results by Nesbitt and Deosjo (2017), our analysis does not
recover a South American clade composed of Luperasuchus
fractus, Prestosuchus chiniquensis, and Saurosuchus galilei.
Instead, these three taxa are in a pectinate arrangement, where
Prestosuchus chiniquensis is basal to a clade supporting
Saurosuchus galilei as the sister taxon of Luperasuchus fractus
plus more derived loricatans. Nevertheless, a constrained analy-
sis forcing the monophyly of these three taxa demands just one
extra step (MPTs = 1260; Consistency Index = 0.352; Retention
Index = 0.761). There is no consensus regarding the inner

Figure 10. Reduced strict consensus tree of the phylogenetic analysis. Numbers represent Bremer support values higher than one.
affinities of Luperasuchus fractus, Prestosuchus chiniquensis, and
Saurosuchus galilei. However, none of the trees supports
Luperasuchus fractus and Prestosuchus chiniquensis as sister
taxa. In this analysis, Mandasuchus tanyauchen is the basalmost
member of Loricata in all the MPTs, whereas Ticinosuchus ferox
is the sister taxon of Paracrocodylomorpha and Nundasuchus
songeaensis is the sister taxon of Archosauria. This arrangement
agrees with those by Butler et al. (2017), except by the position of
Nundasuchus songeaensis, which nests within Pseudosuchia.
Discussion and conclusions
As already pointed out, the incomplete and fragmentary record
of ‘Rauisuchia’ is usually regarded as the main reason for its
poorly resolved phylogenetic framework (Gower 2000; França et
al. 2011, 2013; Lautenschlager and Desojo 2011; Lacerda et al.
HISTORICAL BIOLOGY 17
2015, 2016). However, a significant number of comprehensive
phylogenetic studies has been published (e.g. Brusatte et al. 2010;
Nesbitt 2011, Ezcurra 2016). Yet, topology, inner relationships,
and phylogenetic status of ‘rausuchians’ remain largely
unsolved. Two most comprehensive high-level phylogenies of
Archosauria, provided by Brusatte et al. (2010) and Nesbitt
(2011), furnished different and somehow conflicting results.
Brusatte et al. (2010), recovered ‘Rauisuchian’ as a monophyletic
group divided into two major subclades, Poposauroidea and
Rauisuchoidea. This latter comprises Prestosuchidae and
Rauisuchidae plus an additional clade including Arganasuchus,
Fasolasuchus, Stagonosuchus, and Ticinosuchus. Altogheter,
‘rauisuchian’ taxa are represented by 20 species in Brusatte et
al. (2010). On the other hand, according to Nesbitt (2011),
‘Rauisuchia’ would be a paraphyletic branch group regarding
Crocodylomorpha. Poposauroidea is recovered on this analysis
18 L. ROBERTO-DA-SILVA ET AL.
as a monophyletic clade in a sister group relationship with
Loricata, both forming the large Paracrocodylomorpha.
Ticinosuchus, was recovered as sister group of Paracrocody-
lomorpha. Prestosuchus, Saurosuchus, Batrachotomus, and
Fasolasuchus are considerated as an early paraphyletic branch
regarding Rauisuchidae plus Crocodylomorpha. Butler et al.
(2014) proposed a more extensive analysis of pseudosuchians,
including new taxa in the dataset of Nesbitt (2011). The inclu-
sion of Erpetosuchidae and Gracilisuchidae colappsed
Archosauria into a polytomic clade, composed of Ornithodira,
Erpetosuchidae, Ornithosuchidae, Gracilisuchidae, Phytosauria,
Revueltosaurus plus Aetosauria, and Paracrocodylomorpha.
This latter one recovered as identical to the original analysis of
Nesbitt (2011). In this sense, the paraphyletic nature of ‘rauisu-
chians’ was maintained with the inclusion of new data from the
dataset of Nesbitt (2011), although the inner relationships of
Archosauria were poorly resolved. Indeed, in a recent study
focusing on the phylogenetic affinities of the problematic
Erpetosuchidae, Lacerda et al. (2018) suggest two hypothesis of
relationships for this clade: (i) as an early branch of pseudosu-
chians, being a sister group of Ornithosuchidae; or (ii) in a closer
relationship with the clade composed by Gracilisuchidae and
Paracrodylomorpha with respect to Orntithosuchidae and
Aetosauria. Therefore, progress has been made along the last
years, but the inner affinities of less inclusive groups are still
unstable. For instance, Nesbitt and Deosjo (2017) recovered a
clade composed by Luperasuchus fractus, Prestosuchus chini-
quensis, and Saurosuchus galilei. As already pointed out, our
new analysis fails to recover this clade. Instead, these taxa are
distributed in a pectinate pattern in the MPTs. However, we
found that just one extra step is necessary to support the
hypothesis by Nesbitt and Deosjo (2017). Therefore, both
scenarios are plausible and agree with the placement of
Prestosuchus chiniquensis within Loricata, whereas Ticin-
osuchus ferox nests in Loricata only in the analysis with a single
less step.
In addition to the taxonomic difficulties surrounding
‘Rauisuchia’, since the first reports by Von Huene (1942),
research involving Prestosuchus chiniquensis usually casts
doubts and generate controversies regarding the taxonomic
status of this taxon. Two species of Prestosuchus were
described, both from early Carnian strata of the Santa Maria
Supersequence (Pinheiros-Chiniquá sequence sensu Horn et
al. 2014; Marsicano 2016): P. chiniquensis and P. loricatus
(Huene 1942), both later considered synonyms, but the para-
lectotype can be attributed to P. chiniquensis and the lecto-
type to other potential new taxon (Krebs 1976; Barberena
1978; Desojo & Rauhut 2009; Lacerda et al. 2016). Kischlat
(2000) discussed this issue and proposed the new combina-
tion ‘Abaporu loricatus’. However, its description was some-
how unclear and did not observe the International Code of
Zoological Nomenclature (ICZN). Therefore, so far, only
Prestosuchus chiniquensis is considered a valid taxon
(Lacerda et al. 2016). The specimen UFRGS-PV-0156-T,
which comprises a huge skull and some postcranial remains
(Barberena 1978; Azevedo 1991, 1995a, 1995b), was ascribed
to P. chiniquensis. However, many authors proposed that this
material, together with the paralectotype of P. chiniquensis
(BSPG1933/L), would actually be a new species (Parrish 1987;
Sereno 1991). Kischlat and Barberena (1999) regarded this
alleged new taxon as ‘Crurotarsi indeterminata’. Again, unfol-
lowing the ICZN and thus considered nomen nudum
(Lacerda et al. 2016). This designation was based upon the
presence of a weak mandibular symphysis, a narrow calca-
neum with an elongate tuber, and a rectilinear femur. Kischlat
(2000) also proposed the name ‘Karamuru vorax’ to UFRGS-
PV-0156-T, UFRGS-PV-0152-T, paralectotype of P. chini-
quensis (BSPHG 1933L/7) and parts of P. loricatus (BSPHG
1933L/24).
At this point, three autapomorphies are acknowledged for
Prestosuchus chiniquensis: (i) an anterior notch between the
scapula and coracoid; (ii) a longitudinal ridge on the dorsal
surface of the ilium, and (iii) the presence of a sharp ridge
leading from the glenoid to anteroventral corner of the
coracoid (Desojo and Rauhut 2008; Nesbitt 2011). Two of
these traits are present in ULBRA-PVT-281, as the specimen
does not preserve the ilium. Moreover, Raugust (2014)
recognized the presence of a similar sharp ridge in the
coracoid of Batrachotomus kupferzellensis. This reinforces
the necessity of a redescription and taxonomic review for
Prestosuchus, as well as discovery and description of new
specimens as the one herein presented. We observed a few
differences among the specimens ascribed to Prestosuchus
chiniquensis, including ULBRA-PVT-281 (morphological,
measurements and even new structures present in this new
specimen). Nevertheless, the morphology of the new speci-
men is consistent with those of UFRGS-PV-0156-T and
UFRGS-PV-0152-T, reinforcing the combination of these
specimens as a unique operational taxonomic unit in phy-
logenetic studies. Regarding the morphological variation
that surrounds the specimens ascribed to Prestosuchus, alter-
native and equally possible hypothesys can be drawn in
order to explain them: sexual dimorphism, ontogenetic fea-
tures, interspecific polymorphism, artefact of preservation,
and even taxonomic misidentification. Unfortunately, at this
point, studies addressing to any of these issues are scarce.
Actually, Lacerda et al. (2016) recognized some ontogenetic
trends regarding the rostrum of Prestosuchus chiniquensis,
which pinpoints the beginning of studies focusing on biolo-
gical aspects of this top predator. Therefore, a continuous
sample effort may produce new information regarding this
issue in the next years.
Acknowledgments
The authors are greatly indebted to Pablo Ortiz, Ricardo Martínez and
Oscar Alcober for allowing the access to the specimens under their care.
We also extend our gratitude to the two reviewers for their contribution
to the improvement of this manuscript. We are also grateful to the Willi
Hennig Society for the gratuity of TNT software.
Disclosure statement
No potential conflict of interest was reported by the authors.

Funding
We thank the Fundação de Apoio à Tecnologia e Ciência (FATEC -
process [3.01.0046]) for the financial support to LRS; the Conselho
Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for the
research grant to SDS (process [301801/2012-6]; and the Fundação de
Amparo à Ciência e Tecnologia do Estado de Pernambuco (FACEP)
(process [APQ-0165-2.04/14]) for the financial support to MAGF.
ORCID
Rodrigo Temp Müller http://orcid.org/0000-0001-8894-9875
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