Journal of South American Earth Sciences 120 (2022) 104039

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Journal of South American Earth Sciences

journal homepage: www.elsevier.com/locate/jsames

The oldest South American silesaurid: New remains from the Middle
Triassic (Pinheiros-Chiniquá Sequence, Dinodontosaurus Assemblage Zone)
increase the time range of silesaurid fossil record in southern Brazil
Flávio Augusto Pretto a, b, *, Rodrigo Temp Müller a, b, Debora Moro a, b, Maurício Silva Garcia a, b,
Voltaire Dutra Paes Neto c, Átila Augusto Stock Da Rosa b, d
a
Centro de Apoio à Pesquisa Paleontológica da Quarta Colônia, Universidade Federal de Santa Maria, São João do Polêsine, Brazil
b
Programa de Pós-Graduação Em Biodiversidade Animal, Universidade Federal de Santa Maria, Santa Maria, Brazil
c
Laboratório de Paleobiologia, Universidade Federal do Pampa, São Gabriel, Brazil
d
Laboratório de Estratigrafia e Paleobiologia, Universidade Federal de Santa Maria, Santa Maria, Brazil

A R T I C L E I N F O A B S T R A C T

Keywords: The Triassic deposits of South America are key to understand the early radiation of Dinosauromorpha. Though
Silesauridae the fossil record of the group is relatively abundant in Carnian- and Norian-aged strata from Argentina and
Dinosauromorpha Brazil, Middle Triassic dinosauromorphs are scarce, at least in the Brazilian record. In this contribution, we
Triassic
describe a set of fossils collected from a site biostratigraphically correlated to the Dinodontosaurus Assemblage
Paraná basin
Zone, which is regarded as Ladinian in age. Within the sample, we identified a single partial ilium with
anatomical features consistent with Silesauridae. As a result, we propose a new taxon, Gamatavus antiquus gen. et
sp. nov. as the first silesaurid from the Brazilian Middle Triassic. Biostratigraphic comparisons suggest the
Dinodontosaurus Assemblage Zone to be older than the Argentinean Massetognathus-Chanaresuchus AZ, rendering
the new specimen also the oldest South American silesaurid. This contribution adds to recent works that suggest
biostratigraphical similarity between the Brazilian Dinodontosaurus AZ and the African Lifua Member and Nta­
were Formation, from where the oldest dinosauromorph records are registered.

1. Introduction mostly restricted to a few specimens: the enigmatic Saltopus elginensis

The Triassic Period is a crucial moment in life history, and represents
a time in which most of modern key vertebrate groups originated
(Benton, 2016). Likewise, South American Triassic strata preserve an
important fossil record that provides insights about the evolution of
Triassic faunas (Bonaparte, 1982; Langer, 2005; Langer et al., 2007;
Martínez et al., 2013; Martinelli et al., 2017; Schultz et al., 2020). The
rise and early evolution of Dinosauromorpha was one of the major
events that occurred during the Triassic Period, and one that ultimately
shaped the ecosystems of the succeeding stages of the Mesozoic, with the
radiation of Dinosauria. In that sense, South American record is crucial
to understand this process (Brusatte et al., 2010; Langer et al., 2010;
Novas et al., 2021). Though the record of early dinosauromorphs (i.e.
non-dinosaurian Dinosauromorpha) is still relatively scarce, the fossils
recovered in the last two decades greatly increased the knowledge of this
radiation. In the last century, data on these ‘dinosaur forerunners’ were
(Huene, 1910) from Lossiemouth Sandstone (Late Triassic) of Scotland
(Benton and Walker, 2011) and some taxa from the Chañares Formation
of Argentina, namely Lagerpeton chanarensis (Romer, 1971), Lagosuchus
talampayensis (Romer, 1972a; Sereno and Arcucci, 1994; Agnolin and
Ezcurra, 2019) and Lewisuchus admixtus (Romer, 1972b; Arcucci, 1987;
Bittencourt et al., 2014; Ezcurra et al., 2019; Agnolín et al., 2021) that
collectively provided an understanding that the early stages of dino­
sauromorph evolution consisted mainly of small bipedal insectivores
(Novas, 1996). A dramatic change in this scenario came with the
description of Silesaurus opolensis (Dzik, 2003), and soon thereafter a
series of new discoveries (Table 1) allowed the establishment of an
entirely new clade of dinosauromorphs, the Silesauridae (Langer et al.,
2010; Nesbitt et al., 2010), generally recognized as semi-quadrupedal,
long-limbed forms, most with beak-tipped jaws possessing robust teeth
possibly related to an herbivorous feeding strategy (Langer et al., 2013),
but with an ancestral faunivorous condition (Ezcurra et al., 2019; Müller

* Corresponding author. Rua Maximiliano Vizzoto 598, São João do Polêsine, RS, 97230-000, Brazil.
E-mail address: flavio.pretto@ufsm.br (F.A. Pretto).

https://doi.org/10.1016/j.jsames.2022.104039
Received 28 April 2022; Received in revised form 13 September 2022; Accepted 13 September 2022
Available online 28 September 2022
0895-9811/© 2022 Elsevier Ltd. All rights reserved.
F.A. Pretto et al. Journal of South American Earth Sciences 120 (2022) 104039

Table 1 silesaurid Sacisaurus agudoensis from overlaying Norian strata (Ferigolo

List of named lagerpetid and non-dinosaurian dinosauromorph taxa. Abrev.: and Langer, 2007) of the Riograndia AZ. Lagerpetids (which may alter­
MTr, Middle Triassic; LTr, Late Triassic. natively be recovered as dinosauromorphs or pterosauromorphs) are
Taxon Age Provenance Reference represented by Ixalerpeton polesinensis, as well as isolated materials, from
1. Gamatavus MTr (Ladinian) Brazil This contribution
late Carnian strata (Cabreira et al., 2016; Garcia et al., 2019a) of the
antiquus Hyperodapedon AZ. Furthermore, there is an indeterminate dinosaur­
2. Asilisaurus M/LTr Tanzania (Nesbitt et al., 2010, 2020) omorph from the Middle Triassic beds of the Dinodontosaurus AZ (Müller
kongwe (Ladinian/ and Garcia, 2022). The new specimens here described come from the
Carnian)
Picada do Gama site, which belongs to the Dinodontosaurus AZ. Histor­
3. Lutungutali M/LTr Zambia (Peecook et al., 2013,
sitwensis (Ladinian/ 2017) ically, there are no records of silesaurids referred to this AZ. This renders
Carnian) the material presented here the first and only silesaurid specimen re­
4. Lagerpeton LTr (Carnian) Argentina (Romer, 1971, 1972a) ported for this unit, representing the oldest records of the clade, both in
chanarensis Brazil and in South America.
5. Lagosuchus LTr (Carnian) Argentina (Romer, 1971, 1972a;
talampayensis Agnolin and Ezcurra,
2019) 1.1. Geological settings
6. Lewisuchus LTr (Carnian) Argentina (Romer, 1972b; Ezcurra
admixtus et al., 2019) Specimens were collected in the Picada do Gama site (Fig. 2) which
7. Kongonaphon LTr (Carnian) Madagascar Kammerer et al. (2020)
crops-out at the northern side of the BR 158 federal road
kely
8. Ixalerpeton LTr (Carnian) Brazil Cabreira et al. (2016) (29◦ 46′ 37.60′′ S; 54◦ 06′ 00.42′′ W), Dilermando de Aguiar Municipality,
polesinensis circa 33 Km southwest of Santa Maria. The outcrop is formed by a
9. Ignotosaurus LTr (Carnian) Argentina Martínez et al. (2013) subvertical fault that juxtaposes medium sandstones and reddish
fragilis massive mudstones (Da-Rosa et al., 2005). Based on surrounding out­
10. Diodorus LTr (Carnian/ Morocco Kammerer et al. (2012)
scytobrachion Norian)
crops, the sandstones are the basalmost unit. The Picada do Gama site
11. Silesaurus LTr (Carnian/ Poland Dzik (2003) belongs to the lower portion of Santa Maria Formation (Andreis et al.,
opolensis Norian) 1980), corresponding to the Pinheiros-Chiniquá Sequence (Horn et al.,
12. Dromomeron LTr (Norian) Argentina Martínez et al. (2016) 2014) of the Santa Maria Supersequence (Zerfass et al., 2003).
gigas
Metric fining-upward cycles compose the medium to fine sandstones
13. Sacisaurus LTr (Norian) Brazil Ferigolo and Langer
agudoensis (2007) at the base, being the coarser lithologies embedded in channel-like
14. Dromomeron LTr (Norian) USA Irmis et al. (2007) structures. Trough cross bedding is common in medium sandstones, as
gregorii well as silicified rhyzoliths and carbonate nodules. The massive reddish
15. Dromomeron LTr (Norian) USA Nesbitt et al. (2009a) mudstones intercalate with decimetric intraformational conglomerates,
romeri
16. Soumyasaurus LTr (Norian) USA Sarigül et al. (2018)
composed by carbonatic nodules. Vertebrate fossils were collected in
aenigmaticus both the sandstones and mudstones, although are more common in the
17. Technosaurus LTr (Norian) USA (Chatterjee, 1984; Sarigül latter. The sandstones can be interpreted as fluvial sand bars, later
smalli et al., 2018) colonized by vegetation. The mudstones were deposited in a floodplain
18. Eucoelophysis LTr (Norian) USA (Sullivan and Lucas, 1999;
context, so both lithologies can be associated as lateral portions of the
baldwini Ezcurra, 2006; Nesbitt
et al., 2007) same fluvial environment.
19. Kwanasaurus LTr (Norian) USA Martz and Small (2019) Archosaur, traversodontid cynodont (cf. Massetognathus) and kan­
williamparkeri nemyeriid dicynodont (cf. Dinodontosaurus) remains are common at the
outcrop, which led Da-Rosa et al. (2005) to correlate this fossil assem­
blage to former Therapsida Cenozone sensu (Barberena, 1977) and
and Garcia, 2020). Recently, reinterpretations placed Lagerpetidae as
(Scherer et al., 1995), equivalent to the Dinodontosaurus Assemblage
early diverging members of Pterosauromorpha (i.e. the sister-clade to
Zone (Fig. 3) sensu (Barberena et al., 1985). Although no absolute dating
Dinosauromorpha), rather than as early dinosauromorphs (Ezcurra
is yet available for the Dinodontosaurus AZ, the existing 237 ± 1.5 Ma for
et al., 2020). In that sense, the earliest named dinosauromorph would be
the overlying Santacruzodon AZ (Philipp et al., 2018) suggests the former
Lagosuchus talampayensis, which depending on the phylogenetic inter­
to be placed at the end of Ladinian, or at least at Ladinian – Carnian
pretation would be a sister-taxon to either Dracohors (i.e. Silesauridae +
boundary (Fig. 3). Data from the Tarjadia AZ from the Chanãres For­
Dinosauria), or to Dinosauria (if silesaurids are considered ornithis­
mation of Argentina corroborate this age assignation (Ezcurra et al.,
chians). Though the phylogenetic positioning of Silesauridae is still a
2017; Novas et al., 2021; Müller and Garcia, 2022). Though the presence
matter of debate (see Müller and Garcia (2020) and references therein),
of Dinodontosaurus and Massetognathus alone is not enough a biostrati­
the group is well diverse and widely dispersed over Pangea (Fig. 1).
graphic marker to distinguish between the Santacruzodon AZ and the
At the same time as new discoveries bolstered the known diversity of
Dinodontosaurus AZ (Horn et al., 2014), identified that the outcrops of
early dinosauromorph records, a refinement of the chronostrati­
the former are confined by tectonic fracturing, and therefore the San­
graphical framework was carried out for several relevant fossil-bearing
tacruzodon AZ has a limited geographic distribution. The Picada do
units (Nesbitt et al., 2009b; Irmis et al., 2011; Martinez et al., 2011;
Gama site is at least 180 km west of the westernmost occurrence of the
Marsicano et al., 2016; Langer et al., 2018; Philipp et al., 2018). In that
Santacruzodon AZ, and following the tectonic framework proposed by
sense, it is recognized that non-dinosaurian dinosauromorphs diverged
Horn et al. (2014), would be situated in the outcropping area of the
prior to the oldest unequivocal dinosaurs, but coexisted with those for at
Dinodontosaurus AZ (but see also Martinelli et al., 2017 for a critical
least 10 million years (Irmis et al., 2007; Nesbitt et al., 2010; Garcia
account on this Assemblage Zone).
et al., 2019a).
In this paper we report new silesaurid materials from southern Brazil.
2. Results
Along the text, for nomenclatural purposes, we consider silesaurids to be
non-dinosaurian dinosauromorphs, except when stated otherwise. The
2.1. Systematic paleontology
Brazilian non-dinosaurian dinosauromorph record is currently repre­
sented by unnamed silesaurid materials (Mestriner et al., 2021) from
Archosauria Cope, 1869 (sensu Gauthier and Padian, 1985).
Carnian layers of the Hyperodapedon Assemblage Zone (AZ), plus the
Ornithodira Gauthier, 1986 (sensu Sereno, 1991).

2
F.A. Pretto et al.
Fig. 1. Paleogeographic map of the world during the Triassic, showing major paleoclimatic belts and the distribution of lagerpetid and non-dinosaurian dino­
sauromorph occurrences. Light beige color on the map represents arid areas, green colors represent temperate zones (Benton, 2016). Silhouette colors refer to the age
of occurrences: purple, Middle Triassic (Anisian and Ladinian); magenta, Late Triassic (Carnian); pink, Late Triassic (Norian). See Table 1 for taxon names. Sil­
houettes not to scale, redrawn from different sources. Based on paleogeographic map generated on Fossilworks (fossilworks.org). Silhouettes redrawn based on
Langer et al. (2013).
Dinosauromorpha Benton, 1985 (sensu Ezcurra et al., 2020).
Silesauridae Langer et al. (2010), Nesbitt et al. (2010).
Gamatavus antiquus gen. et sp. nov.
Etimology – Generic name is an alliteration of the particles “Gama”
(alluding to the Picada do Gama site, from where the fossil was
collected) and “atavus” (Latin, great-grandfather). Specific name “anti­
quus” (Latin, ancient). Both specific and generic names make allusion to
the fact that the materials were collected from strata older than the
usually dinosauromorph-yielding strata from Brazil.
Holotype. UFSM 11348a, partial right ilium. The specimen was
collected in association with a partial left femur (UFSM 11348 b) and
four incomplete vertebrae, but the assignation of the specimens to a
single individual is doubtful (see Discussion, below).
Type Horizon and Locality – ‘Picada do Gama’ site, composed by
mudstones exposed by a roadcut (BR-158 federal road) on the munici­
pality of Dilermando de Aguiar, 33 Km of Santa Maria (Fig. 2). It belongs
to the lower Santa Maria Formation (Andreis et al., 1980),
Pinheiros-Chiniquá Sequence (Horn et al., 2014). The outcrop was
correlated by Da-Rosa et al. (2005) to the Dinodontosaurus AZ, which is
regarded as Ladinian/early Carnian in age (Schultz et al., 2020).
Diagnosis – Gamatavus antiquus is the only silesaurid known for the
Dinodontosaurus AZ. It is also diagnosed by the following unique com­
bination of iliac features: high dorsal iliac blade; supracetabular crest
gently rounded; well-developed brevis fossa, only partially visible in
lateral view and with a well-developed ventral portion limited laterally
by a brevis shelf; no triangular process contiguous to the brevis fossa;
and articular facet for the pubis directed cranioventrally.
Ontogenetic Assessment – The specimen appears to have reached
some advanced degree of skeletal maturity based on the presence of a
raised muscle scar on the postacetabular process of the ilium (Garcia
et al., 2019b).
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Journal of South American Earth Sciences 120 (2022) 104039
3. Description
UFSM 11348a includes an almost complete right ilium, which lacks
only part of the dorsal iliac lamina and the preacetabular process
(Fig. 4). The ilium is craniocaudally longer than dorsoventrally high.
The dorsal iliac lamina is higher than the maximum acetabular height
(in an order of 1.16:1), but it could possibly be slightly lower in its
missing portion. Nonetheless, it is unlikely that it reached the low profile
seen in Silesaurus opolensis (Piechowski et al., 2014). In lateral view, the
dorsal margin of the postacetabular process is almost straight (Fig. 4A
and B) and cranioventrally sloped. In dorsal view, the dorsal iliac lamina
shows a strong lateral concavity, with its deepest point located over the
caudal half of the acetabulum (Fig. 4E). This maximum concavity is
cranially limited by a strong preacetabular ridge which in lateral view is
seen rising craniodorsally from the midpoint of the supracetabular crest.
Most of the dorsal portion of the preacetabular ridge is missing,
including its continuation that forms part of the preacetabular process in
closely related forms.
In cranial view, the preacetabular ridge shows a wide lateromedial
development, forming an almost vertical wall, with its preserved medial
margin dorsomedially inclined. The preserved portion of the pre­
acetabular face is entirely flat, and there is no preacetabular fossa be­
tween the preacetabular ridge and the dorsal edge of the pubic peduncle.
The preserved lateral surface of the dorsal iliac blade is mostly smooth,
except for its caudalmost portion, over the postacetabular process.
There, a rough scar rises, possibly marking the attachment of m. flexor
tibialis externus and part of m. flexor tibialis internus (Hutchinson, 2001).
The ventral portion of the postacetabular process is marked by a series of
longitudinal ridges. Starting at the caudal margin of the ischial
peduncle, a ventral ridge extends towards the midpoint of the post­
acetabular process. There, it forms a triangular longitudinal fossa,
bounded by two further ridges. The medial one is sharp and oblique to
the horizontal plane, being congruent with a ridge that occupies much of
the medial side of the ilium. The lateral ridge starts close to the caudal
F.A. Pretto et al. Journal of South American Earth Sciences 120 (2022) 104039

Fig. 2. Geographical location of the Picada do Gama outcrop. In A and B, simplified geological chart of the southern Brazilian successions on the Paraná Basin
(yellow-shaded area in A). In C, satellite image identifying the location of the outcrop. D, vertical profile of the lithological succession of the outcrop, including the
levels where the dinosauromorph fossils were identified. E, original exposure of the outcrop in 2003, including the point where dinosauromorph fossils
were identified.

portion of the supracetabular crest, it is sharp and laterally projected,
corresponding to the brevis shelf. As it extends caudodorsally towards
the caudalmost portion of the postacetabular process, its lateral surface
loses its sharpness, becoming more rounded. The longitudinal fossa
bounded by these two ridges corresponds to the brevis fossa, proposed as
the area of origin of m. caudofemoralis brevis (Hutchinson, 2001). Given
the slight ventral inclination of the medial shelf that bounds it, the
caudal portion of the brevis fossa is partially visible in lateral view. In
ventral view, the fossa and the two ridges that bound it end at the same
point, giving the caudalmost tip of the postacetabular process a straight
margin.
The ventral margin of the acetabulum is convex, indicating that it
was fully closed. The thin acetabular wall is possibly broken at its ven­
tralmost portion. Therefore, though the ventral margin of the acetabular
wall apparently described an angle, such as seen in many early orni­
thodirans and non-ornithodiran archosaurs, the clear observation of this
feature may be hampered in Gamatavus antiquus. Both the lateral iliac
acetabular wall and the roof of the acetabulum are markedly concave.
The mid portion of the iliac lateral acetabular wall shows a depression
that resulted in an extremely thin wall, convergent to the point where
the ventral margin is broken. This depression creates a rounded
depression at the center of the acetabulum, similar to that recognized in
Saturnalia tupiniquim by (Langer, 2003). This putatively corresponded to
one of the main articular surfaces that received the femoral head, which
among dinosaurs is commonly perforated, as seen for example in her­
rerasaurids (Novas, 1993). The area of the antitrochanter is smooth and
indistinct, with no particular elevation. The articulation margin that
received the pubis describes a convex curvature and is subequal in
length to the margin that contacted the ischium, which is straighter,
apart from a faint notch that marks the ventral margin of the ilium,
cranial to the main articular surface of the ischial peduncle.
The pubic peduncle is cranioventrally inclined when observed in
lateral view. In cranial view, it is lateromedially expanded, and the
dorsal portion of the peduncle is slightly tilted medially. The flat dorsal
margin is smooth, and the vicinity of the articular surface that received
the pubis is composed of a bone rougher than in the remainder of the
structure. The ischial peduncle is more vertically oriented than the pubic
peduncle. The vicinity of the articular surface that received the ischium
is lightly bulged laterally, giving the whole articular surface a teardrop
shape in ventral view. As with the pubic peduncle, this region of the

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F.A. Pretto et al. Journal of South American Earth Sciences 120 (2022) 104039

Fig. 3. Chronostratigraphic scheme of the Brazilian Triassic (excluding the Mata Sequence). Modified from (Zerfass et al., 2003; Horn et al., 2014) and Philipp et al.
(2018). Radiometric ages (yellow pentagons) from Langer et al. (2018) and Philipp et al. (2018). Silhouettes (not to scale, redrawn from different sources) represent
selected taxa from each of the four Assemblage Zones (Schultz et al., 2020) from Middle and Late Triassic.

ischial peduncle is composed of porous bone.
The supracetabular crest roofs the acetabulum, covering it dorsally
and also forming some of its cranial margin. It overhangs slightly the
acetabular area, in such a way that the actual roof of the acetabulum is
concave. In dorsal view, the supracetabular crest reaches its highest
lateral expansion just over the ovoid depression in the acetabular wall.
Cranial to that, the supracetabular crest gently merges to the pubic
peduncle, but does not reach its extremity. The caudal portion of the
supracetabular crest that extends towards the ischial peduncle ends
much more abruptly, and reaches its greatest dorsoventral thickness
close to its caudalmost extent, from where the brevis shelf rises.
Medially, the ilium shows a flat surface of compact bone at the area
that corresponds to the acetabulum in lateral view. A depression ac­
companies the main length of the pubic peduncle, but this seems exag­
gerated by taphonomic deformation. Dorsal to the flat area, some bone
ridges mark the articulation surfaces of the two primordial sacral
vertebrae. The cranialmost ridge has the shape of a crescent and rises
from the medial expansion that forms the preacetabular process,
marking the articular surface of the lateral process of the first primordial
sacral vertebra (S1). It forms a dorsally concave curvature at a point at
the level where the acetabular roof is located in the lateral side of the
ilium, and from its lowest point it splits in two. The dorsal ridge rises
dorsocaudally towards the ventral margin of the postacetabular process,
forming the roof of the brevis fossa. The ventral ridge extends almost
horizontally from the lowest point of the articular surface that received
S1, extending caudally towards the caudal margin of the ischial
peduncle. This forms a slightly triangular surface between the two
ridges, which received the lateral process of the second primordial
caudal vertebra (S2). This area does not expand caudally, to form the
medial wall, or triangular process, seen in several taxa (see discussion
below). Gamatavus antiquus shows an additional ridge at the medial
surface of the ilium, rising dorsal to the articular facet for S1, and

5
F.A. Pretto et al.
Fig. 4. Gamatavus antiquus, holotype UFSM 11348a, a partial right ilium in lateral view (A), medial view (B), cranial view (C), ventral view (D), dorsal view (E) and
caudal view (F). Dashed lines reconstruct the hypothetic outline of missing portions. Scalebar: 2 cm. Abbrev, amr, additional medial ridge (see text); bs, brevis shelf;
bfo, brevis fossa; fto, scar marking the origin site of the flexor tibialis musculature (see text); ip, ischial peduncle; mr, medial ridge; par, preacetabular ridge; pp, pubic
peduncle; sac, supracetabular crest.
extending subparallel to the ridge that limits the articular facet for S2. Its
shape is partially obscured by concretions, which hampers a precise
evaluation of its morphology. It extends dorsocaudally towards the
dorsal margin of the dorsal iliac lamina but fails to reach its dorsal
margin. Assuming this ridge to be an anatomical feature, and not a
taphonomic artifact, this might represent an autapomorphic feature for
Gamatavus antiquus. This however must be considered in light of the
uncertain nature of the feature, added to the fact that few early dino­
sauromorph ilia are clearly exposed in medial view. A rough scar marks
the caudodorsal portion of the medial iliac wall, extending parallel to
the dorsal margin of the iliac blade.
Associated specimen. UFSM 11348 b, partial left femur collected in
association with the holotype.
A proximal fragment of a left femur was found associated to the ilium
and is broken at the distal portion of the fourth trochanter (Fig. 5). As
preserved, the shaft has a thick compact bone wall, with a hollowed
core, and the preserved fragment was compressed during fossildia­
genesis, collapsing lateromedially.
As preserved, the proximal surface of the femur is slender and
elongated, but this was exaggerated by taphonomic compression. A deep
groove extends longitudinally across this surface, dividing it in two. A
groove at the proximal surface of the femur is found in most silesaurids
except in Lewisuchus admixtus (Ezcurra et al., 2019), but the condition of
UFSM 11348 b is more similar to that of Silesaurus opolensis (Dzik, 2003),
differing from the less prominent and anteroposteriorly displaced
groove of Eucoelophysis baldwini (Ezcurra, 2006). An angled, but small
bump marks the craniolateral corner of the proximal surface represents
the craniolateral tuber. A marked bulge on the midpoint of the caudo­
medial margin marks the proximal view of the caudomedial tuber.
Caudal to this, the facies articularis antitrochanterica is deeply marked,
but this depth seems to have been increased in part by the collapse of the
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Journal of South American Earth Sciences 120 (2022) 104039
bone wall. The craniomedial tuber is not visible in proximal view,
though this area may be damaged in the specimen, like the transition
between the femoral head and main shaft, which is poorly preserved.
The fourth trochanter can be observed in the medial surface. It appears
as a proximodistally oriented crest, but does not assume the aliform
morphology seen in dinosaurs, rather being just a low elongated ridge.
The surface immediately cranial to the fourth trochanter is smooth, with
no depressions such as the ovoid pit commonly seen in dinosaur­
omorphs. A marked crest extends distally from the craniolateral tuber,
seen in lateral view. Distal to that, a conspicuous bone knob marks the
position of the cranial trochanter. It is not particularly raised and is fully
integrated to the femoral shaft. Differing from several dinosauriforms
that show a trochanteric shelf contiguous to the cranial trochanter, the
bone wall in the vicinity of the cranial trochanter is smooth. Indeed,
even the linea intermuscularis cranialis, which commonly occurs as a very
pronounced scar in dinosauromorphs, is very subtle in the specimen, if
present at all. The area where the dorsolateral trochanter is observed in
dinosauromorphs shows no particular scarring in the specimen, and a
very faint bump on that surface remains the single indicative of a
dorsolateral trochanter. The area between this region and the ridge that
extends ventrally from the cranial trochanter shows no particular scar­
ring indicative of muscle attachment. Though most of femoral features
related to muscle attachment may be strongly affected by ontogeny
(Nesbitt et al., 2009a; Müller et al., 2019), the thickness of the bone wall
of the specimen is suggestive of maturity.
Associated specimens. UFSM 11348c, four partial vertebrae,
collected in association with the holotype.
Four partial vertebrae were collected in association with the other
fragments. These are composed of two isolated centra (missing the
neural arches, Fig. 6A and B), an isolated neural arch (Fig. 6C), and a
badly worn caudal vertebra (Fig. 6D). There is no evidence that these
F.A. Pretto et al.
Fig. 5. UFSM 11348 b, partial left femur collected with the holotype of Gamatavus antiquus in caudomedial view (A), craniolateral view (B) and proximal view (C).
Abbrev, clt, craniolateral tuber; cmt, caudomedial tuber; ctr, cranial trochanter; dlt, dorsolateral trochanter; faa, facies articularis antitrochanterica; ftr, fourth
trochanter; g, groove.
pertained to the same animal. The isolated centra are similar in shape,
but vary in size, one of them being notably larger. This could indicate
that they pertain to different regions of the axial skeleton, or to two
distinct individuals. Centra are spool-shaped and constricted at the
midpoint. The intervertebral articular surfaces are acoelous. The
contribution of the centrum to the neural channel is well-marked in both
vertebrae and, at least in the larger specimen, it forms a deep pit in its
midportion, where it excavates the centrum. The isolated neural arch is
compatible in size with the larger isolated centrum, but it is not possible
to ascertain if they form a single vertebral unit. It preserves part of the
left neural arch peduncle, but all relevant anatomical features were
eroded away. The bases of the postzygapophyses remain relatively intact
and suggest that the vertebra lacked a postzygodiapophyseal lamina.
The neural spine is complete and robust. It has a distal expansion that
forms a rough spine table. There are also two parallel laminae on its
caudal margin that are suggestive of the spinopostzygapophyseal
laminae, but these are badly preserved. The caudal vertebra, although
more complete than the other elements, is the most damaged element. It
was partially altered by the formation of hard concretions, and severely
worn by overpreparation. The centrum is spool shaped, and the distal
intervertebral articulation is displaced ventrally, compared to its prox­
imal counterpart. The proximal articular surface is very lightly concave,
whereas the distal articular surface is apparently flat. The transverse
processes are elongated and almost parallel to the horizontal plane.
Their surface is severely worn. The zygapophyses were not preserved.
7
Journal of South American Earth Sciences 120 (2022) 104039
The preserved portion of the neural spine is craniocaudally short and
dorsally tall, being comparatively higher than the centrum. The distal­
most portion is missing, and its entire surface is badly preserved,
hampering the visualization of any structures.
4. Phylogenetic analysis
The holotype (UFSM 11348a) of Gamatavus antiquus was submitted
to two phylogenetic analyses employing different datasets (see supple­
mentary files) in order to assess its relationships among archosaurs. The
first round of analyses involved the scoring of the holotype in the dataset
of (Ezcurra, 2016), as modified by (Ezcurra et al., 2020). The analysis
was conducted following the same procedures employed by (Ezcurra
et al., 2020). Given the positive phylogenetic correlation of Gamatavus
antiquus with Silesauridae, a second round of analyses involved the
scoring of the holotype in an updated version of the dataset of (Müller
and Garcia, 2020), which includes a comprehensive sample of sile­
saurids in a single phylogenetic analysis. The analysis was performed
according the former study.
The analysis employing the dataset of (Ezcurra et al., 2020) resulted
in 280 most parsimonious trees of 5002 steps (CI = 0.214, RI = 0.669).
Gamatavus antiquus is recovered in the strict consensus tree as a member
of Silesauridae, grouped with Asilisaurus kongwe (Fig. 7 A, Fig. S1). It
shares with Ornithodira the presence of a of a lateral crest dorsal to the
supracetabular rim (462: 0 → 2). Gamatavus antiquus and Asilisaurus
F.A. Pretto et al.
Fig. 6. UFSM 11348c, incomplete vertebrae collected in association with the holotype of Gamatavus antiquus. A, vertebral centrum in dorsal (top) and lateral
(bottom) views. B, vertebral centrum in dorsal (top) and lateral (bottom) views. C, incomplete neural arch in lateral (left) and caudal (right) views. D, caudal vertebra
in lateral (left) and cranial (right) views. Abbrev, poz, postzygapophysis; prz, prezygapophysis; spt, spine table; tp, transverse process. All elements are to scale.
kongwe share the caudofemoralis brevis muscle origin being laterally
rimed by a brevis shelf and with a ventrally facing brevis fossa (character
465: 2 → 3, but see discussion below). The second analysis (i.e.
employing the dataset of (Müller and Garcia, 2020) resulted in 396 most
parsimonious trees of 983 steps (CI = 0.320, RI = 0.667). The results
recovered ‘silesaurids’ related to Ornithischia as in the original analysis
(Müller and Garcia, 2020). Although UFSM 11348a groups with these
‘silesaurids’ in the strict consensus, the inclusion of the specimen col­
lapses most of the early-diverging branches that lead to core Ornithi­
schia (Fig. 7B, Fig. S2).
5. Discussion
5.1. Fossil assemblage
Despite the fact that all elements presented in this study are reported
as having been collected in association, and most even show compatible
sizes, we opted to refer to them as isolated entities. Although they may
represent a single individual, there is no strong evidence other than the
topotypical argument to support this idea. The poor preservation of the
vertebrae hampers any assertive comparative analysis. Additionally, the
fragmentary condition of the femur and the deformation, especially on
the region of the femoral head, casts doubt on the condition of several
critical anatomical features (most notably the proximal tuberosities of
the bone). Therefore, caution demands only the ilium to be referred to
the taxon here proposed.
5.2. Comparative anatomy of the ilium
Gamatavus antiquus differs from all other silesaurids from which iliac
elements are known by a unique combination of features. Regarding the
8
Journal of South American Earth Sciences 120 (2022) 104039
relative height of the ilium (and particularly of the dorsal iliac blade),
based on the preserved portion of the blade of Gamatavus antiquus, it
does not reach the maximum height seen in Asilisaurus kongwe and
Lutungutali sitwensis (Peecook et al., 2013; Nesbitt et al., 2020), but also
differs from the low iliac blades seen in Silesaurus opolensis and Kwa­
nasaurus williamparkeri. The ventral portion of the cranial surface of the
preacetabular process of the ilium of Gamatavus antiquus is smooth,
whereas in Sacisaurus agudoensis there is an excavation. The supra­
cetabular crest is gently curved in lateral view, differing from the more
straight supracetabular crest of Ignotosaurus fragilis and Kwanasaurus
williamparkeri (Martínez et al., 2013; Martz and Small, 2019). The
ventral margin of the acetabulum is convex, and probably formed a
marked, angular ventral flange, such as that of Asilisaurus kongwe,
Lutungutali sitwensis and Ignotosaurus fragilis, though this structure is
partially fractured in UFSM 11348a. This markedly angular profile is
similar to that of Lagerpetidae, Euparkeria capensis, and most other
non-ornithodiran archosaurs (Ewer, 1965; Nesbitt, 2011; Cabreira et al.,
2016; Ezcurra et al., 2020). Other silesaurids may show a more rounded,
straight or even concave ventral surface (Dzik, 2003; Nesbitt et al., 2010;
Langer et al., 2013; Piechowski et al., 2014; Martz and Small, 2019).
Nonetheless, in some silesaurids this region of the acetabulum is badly
preserved, most likely due to its overall fragility (Ferigolo and Langer,
2007; Piechowski et al., 2014; Ezcurra et al., 2019). In fact, this region
may have a marked reduction in wall thickness, such as shown in
Gamatavus antiquus, which may contribute to the poor preservation of
this feature.
The articulation with the pubis is cranioventrally oriented, similar to
that of Asilisaurus kongwe, Lutungutali sitwensis and Ignotosaurus fragilis.
This differs from the more ventrally oriented profile of the iliopubic
articular facet seen in Lewisuchus admixtus (Ezcurra et al., 2019; Agnolín
et al., 2021). Like most silesaurids except for Asilisaurus kongwe, the area
F.A. Pretto et al. Journal of South American Earth Sciences 120 (2022) 104039

Fig. 7. Strict consensus of the phylogenetic analyses (some clades were collapsed). A, analysis based on the Ezcurra et al. (2020) dataset (280 MPTs, length = 5002,
CI = 0.214, RI = 0.669); B, analysis based on Müller and Garcia (2020) dataset (396 MPTs, length = 983, CI = 0.320, RI = 0.667). Colors highlight the corresponding
clades: red, Dinosauromorpha (excluding Lagerpetidae); blue, ‘Silesauridae’; yellow, core Ornithischia; green, Saurischia. Bremer Support values (>1) are indicated
in the nodes.

of attachment of m. caudofemoralis brevis of Gamatavus antiquus is m. caudofemoralis brevis faces laterally for most of its extent except for a
observable in lateral view. However, it does not form a distinct medial very small portion of the caudal extremity of the postacetabular process.
wall (Ezcurra et al., 2019), or a triangular process (Martz and Small,
2019), differing from those of Lewisuchus admixtus, Kwanasaurus wil­
5.3. Biostratigraphic implications
liamparkeri, Ignotosaurus fragilis and some specimens of Silesaurus opo­
lensis. This condition also seems to be absent in Asilisaurus kongwe and
There is an established biostratigraphic correlation between the
Lutungutali sitwensis, and possibly in Sacisaurus agudoensis, though it may
Brazilian Pinheiros-Chiniquá Sequence and the Argentinean Chañares
be affected by fractures in the known specimens of the latter two taxa.
Formation (Schultz et al., 2020), the latter with a conspicuous dino­
This region seems to be related with the articulation of the second pri­
sauromorph fossil record, most remarkably represented by Lewisuchus
mordial sacral vertebra medially, effectively increasing the area of
admixtus and Lagosuchus talampayensis (Romer, 1972b; Ezcurra et al.,
contact. It was possibly closely related to the attachment area for m.
2019; Agnolín et al., 2021). The Chañares Formation encompasses two
caudofemoralis brevis, as also suggested for other dinosauromorphs,
distinct Assemblage Zones (AZs): the Massetognathus-Chanaresuchus AZ,
though the muscle may not have directly attached to this expanded area
which is approximately 236 Ma in age (Marsicano et al., 2016; Ezcurra
(Hutchinson, 2001; Langer, 2003; Piechowski, 2020). Indeed, a large,
et al., 2017) and the underlying Tarjadia AZ, which lacks radioisotopi­
ventrally concave area along the ventral margin of the postacetabular
cally inferred ages (Ezcurra et al., 2017). Nevertheless, following strat­
area (Fig. 8) is recognized by these authors as the main attachment area
igraphic data, the latter is usually placed within the Ladinian – Carnian
for the origin of that muscle. Gamatavus antiquus shows a relatively wide
boundary (Ezcurra et al., 2017). The Brazilian Dinodontosaurus AZ has
and well-developed subtriangular concavity in such region, which is
been recently correlated in part with the Tarjadia AZ given the presence
partially visible in lateral view and most evident in lateroventral view.
of closely related stenaulorhynchine rhynchosaurs, erpetosuchids, and
In some taxa, such as Asilisaurus kongwe and several dinosaurs, this
chiniquodontid cynodonts (Ezcurra et al., 2017; Garcia et al., 2019a;
concavity is almost completely obscured from lateral view by a ventrally
Schultz et al., 2020; Novas et al., 2021). Thereby, the Dinodontosaurus
deflected lateral ridge. In others, like Lutungutali sitwensis, the origin of
AZ is putatively older than the Massetognathus-Chanaresuchus AZ.

9
F.A. Pretto et al.
Although the Tarjadia AZ is correlated with the Dinodontosaurus AZ,
there is no record of dinosauromorphs at that unit (Martinelli et al.,
2017; Ezcurra et al., 2017; Novas et al., 2021; Müller and Garcia, 2022).
The fossil record of dinosauromorphs from the Chañares Formation is
restricted to the Massetognathus-Chanaresuchus AZ (Novas et al., 2021).
Therefore, the dinosauromorphs from the Dinodontosaurus AZ are pu­
tatively older than those of the Chañares Formation.
There are also biostratigraphical markers that suggest correlations of
the Dinodontosaurus AZ with early dinosauromorph bearing units in
Africa. The Lifua Member of the Manda Beds (Tanzania), from which
Asilisaurus kongwe was collected, shares several taxa with the Brazilian
Pinheiros-Chiniquá Sequence, including the cynodonts Aleodon and
Scalenodon, as well as the loricatan pseudosuchian Prestosuchus (Marti­
nelli et al., 2017; Desojo et al., 2020). The Ntawere Formation in
Zambia, from which the remains of Lutungutali sitwensis were collected,
shares with the Brazilian unit the presence of the cynodont Luangwa,
also present in the Omingode Formation of Namibia, from which dino­
sauromorphs are still undocumented. In Namibia, the stahleckerid
dicynodont Stahleckeria potens was also described for the base of the
Omingonde Formation, also suggesting biostratigraphic affinities with
the Brazilian Dinodontosaurus AZ (Abdala et al., 2013). Though the
dinosauromorph yielding beds of the Lifua Member and Ntawere
10
Journal of South American Earth Sciences 120 (2022) 104039
Fig. 8. Comparative morphology of selected
dinosauromorph ilia. A, Gamatavus antiquus
(UFSM11348a) in lateral (A1) and lateroventral
(A2) views. B, Lutungutali sitwensis (NHCC LB32)
in lateral (B1) and lateroventral (B2) views. C,
Asilisaurus kongwe (NMT RB159) in lateral (C1)
and lateroventral (C2) views. D, sauropodomorph
dinosaur (CAPPA/UFSM 0200) in lateral (D1) and
lateroventral (D2) views. Abbrev, cfb, caudal
portion of the attachment area of m. caudofe­
moralis brevis.
Formation are usually recognized as Middle Triassic (Anisian), no
radiometric dates are available (Peecook et al., 2017; Novas et al.,
2021). The same is true for the Pinheiros-Chiniquá Sequence, though the
overlaying Santa Cruz Sequence has an Early Carnian datum (Philipp
et al., 2018). The lack of precise radioisotopic dates, added of complex
biostratigraphic scenarios (see Martinelli et al., 2017, Peecook et al.,
2017 and Novas et al., 2021 for a discussion), preclude a precise tem­
poral calibration of many early dinosauromorph records. Nonetheless,
there is an increasing similarity on the faunal content of these units,
traditionally regarded as Middle Triassic.
The fossil record of dinosauromorphs from the Dinodontosaurus AZ is
scarce. Historically, fossils of the elusive Spondylosoma absconditum were
discussed by some authors as representing a saurischian dinosaur
(Huene, 1942; Galton, 2000; Langer, 2004), though the taxon was
recently reassessed as a member of Aphanosauria (Nesbitt et al., 2017).
França et al. (2013) suggested possible dinosauromorph affinities to the
archosauriform Barberenasuchus brasiliensis, which was previously
considered an early crocodylomorph (Mattar, 1989). However, these
were not further investigated. Finally, Müller and Garcia (2022)
described an isolated femur assigned to Dinosauromorpha, but its
fragmentary condition precludes a less inclusive assignation within the
group. Therefore, the new record presented here expands the fossil
F.A. Pretto et al.
record of Dinosauromorpha from the Dinodontosaurus AZ and represents
the oldest silesaurid from South America.
6. Conclusions
Gamatavus antiquus increases the diversity of the Dinodontosaurus
Assemblage Zone and extends the Brazilian record of silesaurids to the
Middle Triassic. However, though the assemblage of bones that
comprise UFSM 11348 includes several elements of the skeleton, only
the ilium can be confidently assigned to Silesauridae, and thus the femur
and vertebrae are not included in the hypodigm of the taxon. Finally, we
agree with previous workers that the increasing faunal similarity among
the Dinodontosaurus AZ (Brazil), the Tarjadia AZ (Argentina) and the
dinosauromorph bearing units of the Lifua Member (Tanzania) and the
Ntawere Formation (Zambia) may indicate that these units share in part
the same temporal range.
CRediT authorship contribution statement
Flávio Augusto Pretto: Writing – review & editing, Writing – orig­
inal draft, Visualization, Validation, Supervision, Software, Resources,
Project administration, Methodology, Investigation, Funding acquisi­
tion, Formal analysis, Data curation, Conceptualization. Rodrigo Temp
Müller: Writing – review & editing, Writing – original draft, Software,
Methodology, Investigation, Funding acquisition, Formal analysis, Data
curation, Conceptualization. Debora Moro: Writing – review & editing,
Software, Methodology, Formal analysis, Data curation. Maurício Silva
Garcia: Writing – review & editing, Software, Investigation, Formal
analysis, Conceptualization. Voltaire Dutra Paes Neto: Writing – re­
view & editing, Methodology, Investigation, Formal analysis. Átila
Augusto Stock da Rosa: Writing – review & editing, Investigation,
Formal analysis, Data curation.
Declaration of competing interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Data availability
Data will be made available on request.
Acknowledgements
We thank Prof. Leopoldo Witeck Neto (UFSM) for collecting most of
the materials from the Picada do Gama Site, as well as Dr. Max Cardoso
Langer, Dr. Brandon Peecook and an anonymous reviewer for sugges­
tions that greatly improved the manuscript.
This work was supported by Fundação de Amparo à Pesquisa do
Estado do Rio Grande do Sul (FAPERGS process 21/2551-0000619-6 to
FAP and 21/2551-0000680-3 to RTM); Conselho Nacional de Desen­
volvimento Científico e Tecnológico (CNPq processes 313494/2018–5
and 303972/2021–1 to AASDR; 404095/2021–6 to RTM); and Coor­
denação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES
grants 88887.644439/2021–00 to DM and 88887.608076/2021–00 to
MSG).
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.jsames.2022.104039.
11
Journal of South American Earth Sciences 120 (2022) 104039
References
Abdala, F., Marsicano, C.A., Smith, R.M.H., Swart, R., 2013. Strengthening western
gondwanan correlations: a Brazilian dicynodont (synapsida, anomodontia) in the
Middle triassic of Namibia. Gondwana Res. 23, 1151–1162. https://doi.org/
10.1016/j.gr.2012.07.011.
Agnolin, F.L., Ezcurra, M.D., 2019. The validity of Lagosuchus talampayensis romer, 1971
(archosauria, dinosauriformes), from the late triassic of Argentina. Breviora 565, 21.
https://doi.org/10.3099/0006-9698-565.1.1.
Agnolín, F., Brissón Egli, F., Ezcurra, M.D., Langer, M.C., Novas, F., 2021. New specimens
provide insights into the anatomy of the dinosauriform Lewisuchus admixtus Romer,
1972 from the upper Triassic levels of the Chañares Formation, NW Argentina. Anat.
Rec. 24731 https://doi.org/10.1002/ar.24731.
Andreis, R.R., Bossi, G.E., Montardo, D.K., 1980. O grupo rosario do Sul (triassico) no Rio
Grande do Sul. In: Anais Do XXXI Congresso Brasileiro de Geologia, pp. 659–673.
Camboriú.
Arcucci, A.B., 1987. Un nuevo Lagosuchidae (Thecodontia-Pseudosuchia) de la fauna de
Los Chañares (Edad Reptil Chanarense, Triasico Medio), La Rioja, Argentina.
Ameghiniana 24, 89–94.
Barberena, M.C., 1977. Bioestratigrafia preliminar da Formação Santa Maria. Pesquisas
7, 111–129.
Barberena, M.C., Araújo-Barberena, D.C., Lavina, E.L., 1985. Late permian and triassic
tetrapods of southern Brazil. Natl. Geogr. Res. 1, 5–20.
Benton, M.J., 1985. Classification and phylogeny of the diapsid reptiles. Zool. J. Linn.
Soc. 84, 97–164.
Benton, M.J., Walker, A.D., 2011. Saltopus, a dinosauriform from the upper triassic of
Scotland. Earth Environ Sci Trans R Soc Edinb 101, 285–299. https://doi.org/
10.1017/S1755691011020081.
Benton, M.J., 2016. The triassic. Curr. Biol. 26, R1205–R1225. https://doi.org/10.1016/
j.cub.2016.10.060.
Bittencourt, J. de S., Arcucci, A.B., Marsicano, C.A., Langer, M.C., 2014. Osteology of the
Middle Triassic archosaur Lewisuchus admixtus Romer (Chañares Formation,
Argentina), its inclusivity, and relationships amongst early dinosauromorphs
dinosauromorphs. J. Syst. Palaeontol. 13, 189–219. https://doi.org/10.1080/
14772019.2013.878758.
Bonaparte, J.F., 1982. Faunal replacement in the triassic of South America. J. Vertebr.
Paleontol. 2, 362–371.
Brusatte, S.L., Nesbitt, S.J., Irmis, R.B., Butler, R.J., Benton, M.J., Norell, M.A., 2010. The
origin and early radiation of dinosaurs. Earth Sci. Rev. 101, 68–100. https://doi.org/
10.1016/j.earscirev.2010.04.001.
Cabreira, S.F., Kellner, A.W.A., Dias-da-Silva, S., Silva, L.R., Bronzati, M., Marsola, J.C.
de A., Müller, R.T., Bittencourt, J. de S., Batista, B.J., Raugust, T., Carrilho, R.,
Brodt, A., Langer, M.C., 2016. A unique Late Triassic dinosauromorph assemblage
reveals dinosaur ancestral anatomy and diet. Curr. Biol. 26, 3090–3095.
Chatterjee, S., 1984. A new ornithischian dinosaur from the triassic of north America.
Naturwissenschaften 71, 630–631.
Cope, E.D., 1869. Synopsis of the extinct batrachia, reptilia and aves of north America.
Trans. Am. Phil. Soc. 14, 1–252. https://doi.org/10.2307/1005355.hdl:2027/
nyp.33433090912423.
Da-Rosa, Á.A.S., Schwanke, C., Witeck Neto, L., Aurélio, P.L.P., 2005. Sítio Picada do
Gama”: a new fossiliferous locality for the Middle Triassic of southern Brazil. In:
I Congresso Latino-Americano de Paleontologia de Vertebrados. Boletim de
Resumos, Rio de Janeiro, pp. 91–92.
Desojo, J., von Baczko, M., Rauhut, O., 2020. Anatomy, taxonomy and phylogenetic
relationships of Prestosuchus chiniquensis (Archosauria: pseudosuchia) from the
original collection of von Huene, Middle-Late Triassic of southern Brazil. Palaeontol.
Electron. https://doi.org/10.26879/1026.
Dzik, J., 2003. A beaked herbivorous archosaur with dinosaur affinities from the Early
Late Triassic of Poland. J. Vertebr. Paleontol. 23, 556–574.
Ewer, RosalieF., 1965. The anatomy of the thecodont reptile Euparkeria capensis broom.
Phil. Trans. Biol. Sci. 248, 379–435. https://doi.org/10.1098/rstb.1965.0003.
Ezcurra, M.D., 2006. A review of the systematic position of the dinosauriform archosaur
Eucoelophysis baldwini Sullivan & Lucas, 1999 from the upper Triassic of New
Mexico, USA. Geodiversitas 28, 649–684.
Ezcurra, M.D., 2016. The phylogenetic relationships of basal archosauromorphs, with an
emphasis on the systematics of proterosuchian archosauriforms. PeerJ 4, e1778.
https://doi.org/10.7717/peerj.1778.
Ezcurra, M.D., Fiorelli, L.E., Martinelli, A.G., Rocher, S., von Baczko, M.B., Ezpeleta, M.,
Taborda, J.R.A., Hechenleitner, E.M., Trotteyn, M.J., Desojo, J.B., 2017. Deep
faunistic turnovers preceded the rise of dinosaurs in southwestern Pangaea. Nat Ecol
Evol 1, 1477–1483. https://doi.org/10.1038/s41559-017-0305-5.
Ezcurra, M.D., Nesbitt, S.J., Fiorelli, L.E., Desojo, J.B., 2019. New specimen sheds light
on the anatomy and taxonomy of the early late triassic dinosauriforms from the
Chañares Formation, NW Argentina. Anat. Rec. 24243 https://doi.org/10.1002/
ar.24243.
Ezcurra, M.D., Nesbitt, S.J., Bronzati, M., Dalla Vecchia, F.M., Agnolin, F.L., Benson, R.B.
J., Brissón Egli, F., Cabreira, S.F., Evers, S.W., Gentil, A.R., Irmis, R.B., Martinelli, A.
G., Novas, F.E., Roberto da Silva, L., Smith, N.D., Stocker, M.R., Turner, A.H.,
Langer, M.C., 2020. Enigmatic dinosaur precursors bridge the gap to the origin of
Pterosauria. Nature 588, 445–449. https://doi.org/10.1038/s41586-020-3011-4.
Ferigolo, J., Langer, M.C., 2007. A Late Triassic dinosauriform from south Brazil and the
origin of the ornithischian predentary bone. Hist. Biol. 19, 23–33. https://doi.org/
10.1080/08912960600845767.
França, M.A.G., Bittencourt, J. de S., Langer, M.C., 2013. Reavaliação taxonômica de
Barberenasuchus brasiliensis (Archosauriformes), Ladiniano do Rio Grande do Sul
(Zona-Assembléia de Dinodontosaurus). Paleontologia em Destaque 230.
F.A. Pretto et al.
Galton, P.M., 2000. Are spondylosoma and staurikosaurus (Santa Maria Formation,
middle-upper triassic, Brazil) the oldest saurischian dinosaurs? Paläontol. Z. 74,
393–423.
Garcia, M.S., Müller, R.T., Da-Rosa, Á.A.S., Dias-da-Silva, S., 2019a. The oldest known
co-occurrence of dinosaurs and their closest relatives: a new lagerpetid from a
Carnian (Upper Triassic) bed of Brazil with implications for dinosauromorph
biostratigraphy, early diversification and biogeography. J South Am Earth Sci 91,
302–319. https://doi.org/10.1016/j.jsames.2019.02.005.
Garcia, M.S., Pretto, F.A., Dias-Da-silva, S., Müller, R.T., 2019b. A dinosaur ilium from
the Late Triassic of Brazil with comments on key-character supporting Saturnaliinae.
An. Acad. Bras. Cienc. 91 https://doi.org/10.1590/0001-3765201920180614.
Gauthier, J., 1986. Saurischian monophyly and the origin of birds. In: Padian, K. (Ed.),
The Origin of Birds and the Evolution of Flight. Memoir of the California Academy of
Science, vol. 8. California Academy of Sciences, pp. 1–55.
Gauthier, J., Padian, K., 1985. Phylogenetic, functional, and aerodynamic analyses of the
origin of birds and their flight. In: Hecht, M.K., Ostrom, J.H., Viohl, G.,
Wellnhofer, P. (Eds.), The Beginnings of Birds, Freude des Jura-Museums,
pp. 185–197. Eichstätt, Germany.
Horn, B.L.D., Melo, T.M., Schultz, C.L., Philipp, R.P., Kloss, H.P., Goldberg, K., 2014.
A new third-order sequence stratigraphic framework applied to the Triassic of the
Paraná Basin, Rio Grande do Sul, Brazil, based on structural, stratigraphic and
paleontological data. J South Am Earth Sci 55, 123–132. https://doi.org/10.1016/j.
jsames.2014.07.007.
Huene, F. von, 1942. Die fossilen Reptilien des sudamerikanischen Gondwanalandes. C.
H. Beck, Munich.
Huene, F. von, 1910. Ein primitiver Dinosaurier aus der mittleren Trias von Elgin.
Geologie und Paläontologie Abhandlungen 8, 317–322.
Hutchinson, J., 2001. The evolution of pelvic osteology and soft tissues on the line to
extant birds (Neornithes). Zool. J. Linn. Soc. 131, 123–168. https://doi.org/
10.1006/zjls.2000.0254.
Irmis, R.B., Nesbitt, S.J., Padian, K., Smith, N.D., Turner, A.H., Woody, D., Downs, A.,
2007. A Late Triassic dinosauromorph assemblage from New Mexico and the rise of
dinosaurs. Science 317, 358–361. https://doi.org/10.1126/science.1143325, 1979.
Irmis, R.B., Mundil, R., Martz, J.W., Parker, W.G., 2011. High-resolution U – Pb ages
from the Upper Triassic Chinle Formation (New Mexico, USA) support a diachronous
rise of dinosaurs. Earth Planet Sci. Lett. 309, 258–267. https://doi.org/10.1016/j.
epsl.2011.07.015.
Kammerer, C.F., Nesbitt, S.J., Shubin, N.H., 2012. The first silesaurid dinosauriform from
the Late Triassic of Morocco. Acta Palaeontol. Pol. 57, 277–284.
Kammerer, C.F., Nesbitt, S.J., Flynn, J.J., Ranivoharimanana, L., Wyss, A.R., 2020. A tiny
ornithodiran archosaur from the Triassic of Madagascar and the role of
miniaturization in dinosaur and pterosaur ancestry. Proc. Natl. Acad. Sci. USA 117,
17932–17936. https://doi.org/10.1073/pnas.1916631117.
Langer, M.C., 2003. The pelvic and hind limb anatomy of the stem-sauropodomorph
Saturnalia tupiniquim (Late Triassic, Brazil). PaleoBios 23, 1–40.
Langer, M.C., 2004. Basal saurischia. In: Weishampel, D.B., Dodson, P., Osmólska, H.
(Eds.), The Dinosauria. University of California Press, Berkeley, pp. 25–46.
Langer, M.C., 2005. Studies on continental Late Triassic tetrapod biochronology. II. The
Ischigualastian and a Carnian global correlation. J South Am Earth Sci 19, 219–239.
https://doi.org/10.1016/j.jsames.2005.04.002.
Langer, M.C., Ribeiro, A.M., Schultz, C.L., Ferigolo, J., 2007. The continental tetrapod-
bearing Triassic of South Brazil. In: Lucas, S.G., Spielmann, J.A. (Eds.), The Global
Triassic, New Mexico Museum of Natural History and Science Bulletin, vol. 41,
pp. 201–218.
Langer, M.C., Ezcurra, M.D., Bittencourt, J.S., Novas, F.E., 2010. The origin and early
evolution of dinosaurs. Biol. Rev. 85, 55–110. https://doi.org/10.1111/j.1469-
185X.2009.00094.x.
Langer, M.C., Nesbitt, S.J., Bittencourt, J. de S., Irmis, R.B., 2013. Non-dinosaurian
Dinosauromorpha. Geological Society, London, Special Publications. https://doi.
org/10.1144/SP379.9.
Langer, M.C., Ramezani, J., Da Rosa, Á.A.S., 2018. U-Pb age constraints on dinosaur rise
from south Brazil. Gondwana Res. 57, 133–140. https://doi.org/10.1016/j.
gr.2018.01.005.
Marsicano, C.a., Irmis, R.B., Mancuso, A.C., Mundil, R., Chemale, F., 2016. The precise
temporal calibration of dinosaur origins. Proc. Natl. Acad. Sci. USA 113, 509–513.
https://doi.org/10.1073/pnas.1512541112.
Martinelli, A.G., Kammerer, C.F., Melo, T.P., Paes Neto, V.D., Ribeiro, A.M., Da-Rosa, Á.
A.S., Schultz, C.L., Soares, M.B., 2017. The african cynodont Aleodon (cynodontia,
probainognathia) in the triassic of southern Brazil and its biostratigraphic
significance. PLoS One 12, e0177948. https://doi.org/10.1371/journal.
pone.0177948.
Martinez, R.N., Sereno, P.C., Alcober, O.A., Colombi, C.E., Renne, P.R., Montañez, I.P.,
Currie, B.S., 2011. A basal dinosaur from the dawn of the dinosaur era in
southwestern pangaea. Science 331, 206–210, 1979.
Martínez, R.N., Apaldetti, C., Alcober, O.A., Colombi, C.E., Sereno, P.C., Fernandez, E.,
Malnis, P.S., Correa, G.A., Abelin, D., 2013. Vertebrate succession in the
ischigualasto formation. J. Vertebr. Paleontol. 32, 10–30.
Martínez, R.N., Apaldetti, C., Correa, G.A., Abelín, D., 2016. A norian lagerpetid
dinosauromorph from the quebrada del barro formation, northwestern Argentina.
Ameghiniana 53, 1–13. https://doi.org/10.5710/AMGH.21.06.2015.2894.
Martz, J.W., Small, B.J., 2019. Non-dinosaurian dinosauromorphs from the chinle
formation (upper triassic) of the eagle basin, northern Colorado: dromomeron
romeri (Lagerpetidae) and a new taxon, Kwanasaurus williamparkeri (Silesauridae).
PeerJ 7, e7551. https://doi.org/10.7717/peerj.7551.
12
Journal of South American Earth Sciences 120 (2022) 104039
Mattar, L.C.B., 1989. Descrição osteólogica do crânio e segunda vértebra cervical de
Barberenasuchus brasiliensis Mattar, 1987 (Reptilia, Thecodontia) do Mesotriássico do
Rio Grande do Sul, Brasil. An. Acad. Bras. Cienc. 61, 319–333.
Mestriner, G., LeBlanc, A., Nesbitt, S.J., Marsola, J.C.A., Irmis, R.B., Da-Rosa, Á.A.S.,
Ribeiro, A.M., Ferigolo, J., Langer, M., 2021. Histological analysis of
ankylothecodonty in Silesauridae (Archosauria: dinosauriformes) and its
implications for the evolution of dinosaur tooth attachment. Anat. Rec. 24679
https://doi.org/10.1002/ar.24679.
Müller, R.T., Langer, M.C., Pacheco, C.P., Dias-da-Silva, S., 2019. The role of ontogeny on
character polarization in early dinosaurs: a new specimen from the Late Triassic of
southern Brazil and its implications. Hist. Biol. 31, 794–805. https://doi.org/
10.1080/08912963.2017.1395421.
Müller, R.T., Garcia, M.S., 2020. A paraphyletic ‘Silesauridae’’ as an alternative
hypothesis for the initial radiation of ornithischian dinosaurs. Biol. Lett. 16,
20200417 https://doi.org/10.1098/rsbl.2020.0417.
Müller, R.T., Garcia, M.S., 2022. Oldest dinosauromorph from South America and the
early radiation of dinosaur precursors in Gondwana. Gondwana Res. 107, 42–48.
https://doi.org/10.1016/j.gr.2022.02.010.
Nesbitt, S.J., Irmis, R.B., Parker, W.G., 2007. A critical re-evaluation of the Late Triassic
dinosaur taxa of North America. J. Syst. Palaeontol. 5, 209–243. https://doi.org/
10.1017/S1477201907002040.
Nesbitt, S.J., Irmis, R.B., Parker, W.G., Smith, N.D., Turner, A.H., Rowe, T., 2009a.
Hindlimb osteology and distribution of basal dinosauromorphs from the Late Triassic
of North America. J. Vertebr. Paleontol. 29, 498–516.
Nesbitt, S.J., Smith, N.D., Irmis, R.B., Turner, A.H., Downs, A., Norell, M.A., 2009b.
A complete skeleton of a late triassic saurischian and the early evolution of
dinosaurs. Science 326, 1530–1533, 1979.
Nesbitt, S.J., Sidor, C.A., Irmis, R.B., Angielczyk, K.D., Smith, R.M.H., Tsuji, L.A., 2010.
Ecologically distinct dinosaurian sister group shows early diversification of
Ornithodira. Nature 464, 95–98. https://doi.org/10.1038/nature08718.
Nesbitt, S.J., 2011. The early evolution of archosaurs: relationships and the origin of
major clades. Bull. Am. Mus. Nat. Hist. 1–292.
Nesbitt, S.J., Butler, R.J., Ezcurra, M.D., Barrett, P.M., Stocker, M.R., Angielczyk, K.D.,
Smith, R.M.H., Sidor, C.A., Niedźwiedzki, G., Sennikov, A.G., Charig, A.J., 2017. The
earliest bird-line archosaurs and the assembly of the dinosaur body plan. Nature 544,
484–487. https://doi.org/10.1038/nature22037.
Nesbitt, S.J., Langer, M.C., Ezcurra, M.D., 2020. The anatomy of Asilisaurus kongwe, a
dinosauriform from the Lifua member of the Manda beds (~Middle triassic) of africa.
Anat. Rec. 303, 813–873. https://doi.org/10.1002/ar.24287.
Novas, F.E., 1993. New information on the systematics and postcranial skeleton of
Herrerasaurus ischigualastensis (theropoda: herrerasauridae) from the ischigualasto
formation (upper triassic) of Argentina. J. Vertebr. Paleontol. 13, 400–423.
Novas, F.E., 1996. Dinosaur monophyly. J. Vertebr. Paleontol. 16, 723–741. https://doi.
org/10.1080/02724634.1996.10011361.
Novas, F.E., Agnolin, F.L., Ezcurra, M.D., Temp Müller, R., Martinelli, A.G., Langer, M.C.,
2021. Review of the fossil record of early dinosaurs from South America, and its
phylogenetic implications. J South Am Earth Sci 110, 103341. https://doi.org/
10.1016/J.JSAMES.2021.103341.
Peecook, B.R., Sidor, C.A., Nesbitt, S.J., Smith, R.M.H., Steyer, J.S., Angielczyk, K.D.,
2013. New silesaurid from the upper Ntawere Formation of Zambia (Middle triassic)
demonstrates the rapid diversification of Silesauridae (avemetatarsalia,
dinosauriformes). J. Vertebr. Paleontol. 33, 1127–1137.
Peecook, B.R., Steyer, J.S., Tabor, N.J., Smith, R.M.H., 2017. Updated geology and
vertebrate paleontology of the Triassic Ntawere Formation of northeastern Zambia,
with special emphasis on the archosauromorphs. J. Vertebr. Paleontol. 37, 8–38.
https://10.1080/02724634.2017.1410484.
Philipp, R.P., Schultz, C.L., Kloss, H.P., Horn, B.L.D., Soares, M.B., Basei, M.A.S., 2018.
Middle Triassic SW Gondwana paleogeography and sedimentary dispersal revealed
by integration of stratigraphy and U-Pb zircon analysis: the Santa Cruz Sequence,
Paraná Basin, Brazil. J South Am Earth Sci 88, 216–237. https://doi.org/10.1016/J.
JSAMES.2018.08.018.
Piechowski, R., Tałanda, M., Dzik, J., 2014. Skeletal variation and ontogeny of the late
triassic dinosauriform Silesaurus opolensis. J. Vertebr. Paleontol. 34, 1383–1393.
https://doi.org/10.1080/02724634.2014.873045.
Piechowski, R., 2020. The Locomotor Musculature and Posture of the Early
Dinosauriform Silesaurus Opolensis Provides a New Look into the Evolution of
Dinosauromorpha 1044–1100. https://doi.org/10.1111/joa.13155.
Romer, A.S., 1971. The Chañares (Argentina) Triassic Reptile Fauna. X. Two new but
incompletely known long-limbed pseudosuchians. Breviora 1–10.
Romer, A.S., 1972a. The Chañares (Argentina) triassic reptile fauna. XV. Further remains
of the thecodonts Lagerpeton and Lagosuchus. Breviora 1–7.
Romer, A.S., 1972b. The Chañares (Argentina) Triassic Reptile Fauna. XIV. Lewisuchus
admixtus, gen. et sp. nov., a further thecodont from the Chañares beds. Breviora 1–7.
Sarigül, V., Agnolín, F.L., Chatterjee, S., 2018. Description of a multitaxic bone
assemblage from the Upper Triassic Post Quarry of Texas (Dockum Group), including
a new small basal dinosauriform taxon. Hist. Nat. (Corr.) 8, 5–24.
Scherer, C.M.S., Faccini, U.F., Barberena, M.C., Schultz, C.L., Lavina, E.L., 1995.
Bioestratigrafia da Formação Santa Maria: utilização das cenozonas como horizontes
de correlação. Comunicações do Museu de Tecnologia da PUCRS (Série Ciências da
Terra) 1, 43–50.
Schultz, C.L., Martinelli, A.G., Soares, M.B., Pinheiro, F.L., Kerber, L., Horn, B.L.D.,
Pretto, F.A., Müller, R.T., Melo, T.P., 2020. Triassic faunal successions of the Paraná
Basin, southern Brazil. J South Am Earth Sci 104, 102846. https://doi.org/10.1016/
j.jsames.2020.102846.
Sereno, P.C., 1991. Basal archosaurs: phylogenetic relationships and functional
implications. J. Vertebr. Paleontol. 11, 1–53.
F.A. Pretto et al.
Sereno, P.C., Arcucci, A.B., 1994. Dinosaurian precursors from the Middle Triassic of
Argentina: Marasuchus lilloensis, gen. nov. J. Vertebr. Paleontol. 14, 53–73.
Sullivan, R.M., Lucas, S.G., 1999. Eucoelophysis baldwini, a new theropod dinosaur from
the Upper Triassic of New Mexico, and the status of the original types of Coelophysis.
J. Vertebr. Paleontol. 19, 81–90.
13
Journal of South American Earth Sciences 120 (2022) 104039
Zerfass, H., Lavina, E.L., Schultz, C.L., Garcia, A.J.V., Faccini, U.F., Chemale, F., 2003.
Sequence stratigraphy of continental Triassic strata of Southernmost Brazil: a
contribution to Southwestern Gondwana palaeogeography and palaeoclimate.
Sediment. Geol. 161, 85–105. https://doi.org/10.1016/S0037-0738(02)00397-4.