Journal of South American Earth Sciences 91 (2019) 302–319

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Journal of South American Earth Sciences

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

The oldest known co-occurrence of dinosaurs and their closest relatives: A T

new lagerpetid from a Carnian (Upper Triassic) bed of Brazil with
implications for dinosauromorph biostratigraphy, early diversification and
biogeography
Maurício S. Garciaa,b,c,∗, Rodrigo T. Müllerd, Átila A.S. Da-Rosab, Sérgio Dias-da-Silvac
a
Centro de Ciências Naturais e Exatas, Universidade Federal de Santa Maria, Av. Roraima, 1000, Bairro Camobi, 97.105-900, Santa Maria, RS, Brazil
b
Laboratório de Estratigrafia e Paleobiologia, Departamento de Geociências, Universidade Federal de Santa Maria, Santa Maria, RS, 97.105-900, Brazil
c
Laboratório de Paleobiodiversidade Triássica, Departamento de Ecologia e Evolução, Universidade Federal de Santa Maria, Av. Roraima, 1000, Bairro Camobi, 97.105-
900, Santa Maria, RS, Brazil
d
Centro de Apoio à Pesquisa Paleontológica da Quarta Colônia, Universidade Federal de Santa Maria, Rua Maximiliano Vizzotto, 598, 97.230-000, São João do Polêsine,
RS, Brazil

A R T I C LE I N FO A B S T R A C T

Keywords: Late Triassic deposits from south Pangea (Argentina and Brazil) bear the oldest (so far) and most informative
Dinosauromorpha unequivocal records of dinosaurs worldwide. Herein we describe a new lagerpetid dinosauromorph from a
Lagerpetidae classic dinosaur-bearing locality (Cerro da Alemoa site) from the Candelária Sequence (Santa Maria Formation),
Dinosauria southern Brazil, and report the oldest co-occurrence of dinosaurs, lagerpetids and silesaurids, in the same layer of
Candelária sequence
a Triassic outcrop. Although tentatively regarded as a skeletally immature specimen, the new lagerpetid re-
South Pangea
presents a new morphotype due to the occurrence of the following features: (1) distal condyles transversely
wider than deep; (2) absence of flange in the craniomedial margin of the femur; (3) round shaped and medially
deflected medial condyle; (4) round shaped crista tibiofibularis. This suit of traits differentiates the new spe-
cimen from Ixalerpeton polesinensis, (which was so far the only lagerpetid found in Brazilian strata) and from
other known lagerpetids. As non-dinosaurian dinosauromorphs are still relatively scarce in Brazilian Triassic
rocks, recent records from this region are gradually providing new data regarding faunal turnovers and bios-
tratigraphy of south Pangean terrestrial deposits. Additionally, new insights on the oldest dinosauromorph-
bearing sites throughout the Triassic, as well as regarding taxonomic diversity and geographic distribution of
early dinosauromorphs reinforces previous hypotheses where dinosaurs and dinosaur-relatives consistently co-
existed (for at least 21 Ma) before their extinction.

1. Introduction inclusive clades (Lagerpetidae, Silesauridae and Dinosauria – Padian

1.1. Background of the Triassic dinosauromorphs
Dinosauromorpha includes the avian stem lineage, which is mainly
represented by some of the most emblematic vertebrates that ever lived,
the dinosaurs (Fig. 1A). It was firstly proposed by Benton (1985) and a
node-based definition was carried out by Sereno (1991) as including
“Lagerpeton chanarensis, Marasuchus lilloensis, Pseudolagosuchus major,
Dinosauria (including Aves), and all descendants of their most recent
common ancestor”. Dinosauromorphs comprise three major less
and May 1993a,b; Nesbitt et al., 2009a, Langer et al., 2010, but see
Cabreira et al., 2016), all coexisting throughout the Triassic Period
(Ezcurra, 2006; Irmis et al., 2007; Langer et al., 2013; Cabreira et al.,
2016). The basalmost branch of Dinosauromorpha comprises the en-
igmatic Lagerpetidae (Nesbitt et al., 2009a), a clade of cursorial and
relatively small-sized taxa (but see Martínez et al., 2016), in which most
specimens preserve only pelvic and hindlimb elements, obscuring their
anatomy (Novas, 1996; Cabreira et al., 2016; Nesbitt et al., 2017;
Müller et al., 2018a, see further below). However, novel discoveries
show that many of the features present in lagerpetids are actually

∗
Corresponding author. Centro de Ciências Naturais e Exatas, Universidade Federal de Santa Maria, Av. Roraima, 1000, Bairro Camobi, 97.105-900, Santa Maria,
RS, Brazil.
E-mail addresses: maurissauro@mail.ufsm.br (M.S. Garcia), rodrigotmuller@hotmail.com (R.T. Müller), atila@smail.ufsm.br (Á.A.S. Da-Rosa),
paleosp@gmail.com (S. Dias-da-Silva).

https://doi.org/10.1016/j.jsames.2019.02.005
Received 18 December 2018; Received in revised form 6 February 2019; Accepted 7 February 2019
Available online 18 February 2019
0895-9811/ © 2019 Elsevier Ltd. All rights reserved.
M.S. Garcia, et al.
derived within Dinosauromorpha, as suggested by a recently recognized
clade of basal avemetatarsalians (i.e. Aphanosauria – Nesbitt et al.,
2017), instead of symplesiomorphic characters states for dinosaur-
omorphs. Aphanosaurs are bird-line archosaurs regarded as the sister
group of Ornithodira (Pterosauromorpha + Dinosauromorpha –
Gauthier, 1986), which provides clues regarding the ancestral body
plan of Dinosauromorpha and how it transitioned from what is ob-
served in early dinosauromorph taxa (Nesbitt et al., 2010, Nesbitt et al.,
2017). Although lagerpetids are considered the basalmost dinosaur-
omorphs, at this point, the earliest skeletal remains of Dinosaur-
omorpha are represented by members of Dinosauriformes (Novas,
1992) that are more related (or even belonging) to Silesauridae rather
than to Lagerpetidae or Dinosauria (Nesbitt et al., 2010; Peecook et al.,
2013, but see Martinelli et al., 2017b; Wynd et al., 2017). Hence, we
can argue that there is still a “lost chapter” in the evolutionary history
of dinosauromorphs, which is corroborated by ichnological findings
(Brusatte et al., 2010a,b). Silesauridae mainly includes quadrupedal
omnivorous/herbivorous dinosauriforms that existed throughout most
of the Triassic Period (Anisian – late Norian, see Discussion), nearly
achieving a global distribution (Dzik, 2003; Ferigolo and Langer, 2007;
Nesbitt et al., 2017). On the other hand, silesaurids are regarded by
some authors as members of the dinosaurian clade Ornithischia
(Ferigolo and Langer, 2007, Cabreira et al., 2016, see also Agnolin and
Rozadilla, 2017, Baron, 2017).
Besides the abovementioned major clades (Lagerpetidae and
Silesauridae), Dinosauria was undoubtedly the most successful group of
dinosauromorphs during the Mesozoic Era (Irmis, 2011), with a rapid
diversification and radiation soon after its “first breath” in the initial
Late Triassic (mid Carnian of south Pangea – Ezcurra, 2010; Müller
et al., 2018b), ultimately acquiring a plethora of body plans, in a way
that was only achieved by pseudosuchians, among terrestrial faunas
from the Triassic Period (Nesbitt et al., 2017). However, this
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Journal of South American Earth Sciences 91 (2019) 302–319
Fig. 1. A – Time calibrated phylogenetic affinities of
Archosauria focused in the avian line major groups.
Based in the topology of Nesbitt et al. (2017). B –
Diversity and disparity in dinosaurs (to scale) of the
initial Late Triassic (mid-Late Carnian to early
Norian). Large silhouettes (from left to right): Rio-
jasaurus incertus, Herrerasaurus ischigualastensis (PVSJ
53), Lessemsaurus sauropoides. Small silhouettes
(from left to right): Eoraptor lunensis, Pampadromaeus
barberenai, Eodromaeus murphi (see text for refer-
ences). Scale bar = 1 m.
morphological disparity was somehow already present in the latest
Carnian – Norian dinosaurian forms (Fig. 1B), ranging from tiny and
gracile (e.g. Eoraptor lunensis – Sereno et al., 1993) to large and robust
body plans (e.g. Riojasaurus incertus – Bonaparte, 1967), from small
primary/secondary consumers (e.g. Pampadromaeus barberenai –
Cabreira et al., 2011) to apex hypercarnivorous predators [e.g. Her-
rerasaurus ischigualastensis (PVSJ 53) – Reig 1963; Novas 1994; Langer
2004], from bipedal cursorial forms (e.g. Eodromaeus murphi – Martínez
et al., 2011) to quadrupedal with mediportal/graviportal locomotion
(e.g. Lessemsaurus sauropoides – Bonaparte, 1999; Apaldetti et al.,
2018), and so on. Moreover, dinosaurs such as sauropodomorphs and
ornithischians independently developed herbivorous feeding habits
several times in their evolutionary journey still in the beginning of the
Late Triassic (Barrett and Rayfield, 2006; Barrett et al., 2010; Nesbitt
et al., 2010; Cabreira et al., 2016). Accordingly, a remarkable aspect of
the evolutionary path of dinosauromorphs was their extreme ability of
adaptation and opportunism during environmental and faunal turn-
overs which, allied with their extensive geographical and evolutionary
radiation (primarily from Norian onward) (O’Donovan et al. 2018),
made this group the predominant faunal component in land ecosystems.
1.2. Context of the Lagerpetidae
Only recently, this noteworthy clade is receiving more attention, as
new materials increased the interest in early dinosauromorphs, whose
studies are shedding light in their initial evolution and dispersal (Irmis
et al., 2007; Brusatte et al., 2010a,b; Martínez et al., 2013, 2016;
Cabreira et al., 2016). Assumption of ghost-lineages of dinosaur-
omorphs and data from ichnofossils point that their origin possibly took
place still during the Early Triassic (Brusatte et al., 2010a,b; Benton
et al., 2014; Nesbitt et al., 2017), despite the fact that the earliest un-
equivocal records of dinosauromorphs arises in the Middle Triassic (e.g.
M.S. Garcia, et al.
Nesbitt et al., 2010; Peecook et al., 2013). Yet, lagerpetids have their
first skeletal remains unearthed in Carnian rocks from the Late Triassic
of South America (Sereno and Arcucci, 1994a; Martínez et al., 2013;
Cabreira et al., 2016). New lagerpetid specimens are shedding light on
their anatomy, diversification, and evolution throughout the Triassic,
although these findings are typically represented by relatively small
forms (but see Martínez et al., 2016) and pelvic/hindlimb elements,
hindering a more comprehensive knowledge of this group, and strongly
suggesting that its diversity is currently underestimated and mostly
unknown (Cabreira et al., 2016). Aside Lagerpeton chanarensis from Los
Chañares Formation, earliest Carnian of Argentina (Sereno and Arcucci,
1994a; Marsicano et al., 2016), prior knowledge of lagerpetids relied,
almost exclusively, on species from the Norian of North America: Dro-
momeron gregorii and Dromomeron romeri, Colorado City and Chinle
Formations, respectively (Irmis et al., 2007; Nesbitt et al., 2009a;
Sarigül, 2016, 2017; Marsh, 2018). Nowadays, Argentinean and Bra-
zilian strata allowed an increase in the lagerpetid record from both
Carnian and Norian horizons, with the recent addition of Dromomeron
gigas from Quebrada del Barro Formation, Ixalerpeton polesinensis from
the lower Candelária Sequence/upper Santa Maria Formation and an
unnamed lagerpetid (PVSJ 883) from Ischigualasto Formation
(Martínez et al., 2013, 2016; Cabreira et al., 2016). Even so, most of the
skeletal anatomy of lagerpetids is still unknown and poorly sampled
compared to other dinosauromorphs (Fig. 2) (Müller et al., 2018a).
Herein we describe a new lagerpetid specimen, which comprises a
partial left femur (distal end), from the lower to middle portion of the
Candelária Sequence/upper portion of the Santa Maria Formation of
southern Brazil (Hyperodapedon Assemblage Zone, mid Carnian –
Langer et al., 2018, see Geological Setting). Moreover, this stratigraphic
horizon is correlated to the site that yielded the lagerpetid Ixalerpeton
polesinensis (Cabreira et al., 2016). We also test the phylogenetic status
of the new specimen using three different morphological datasets
(Cabreira et al., 2016; Langer et al., 2017b; Nesbitt et al., 2017), and
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Journal of South American Earth Sciences 91 (2019) 302–319
discuss its significance regarding the co-occurrence of the three major
dinosauromorph clades in its outcrop and stratigraphic horizon (Cerro
da Alemoa site, see below). Nonetheless, we also present comments on
the biostratigraphical framework of the South American Triassic ter-
restrial Assemblage Zones and the early dinosauromorph diversification
and biogeographic distribution in the (Early to initial Late) Triassic.
1.3. Institutional abbreviations
CAPPA/UFSM, Centro de Apoio à Pesquisa Paleontológica da
Quarta Colônia, Universidade Federal de Santa Maria, São João do
Polêsine, Brazil; GR, Ruth Hall Museum of Paleontology, Ghost Ranch,
New Mexico; NMMNH, New Mexico Museum of Natural History and
Science, Albuquerque, New Mexico; PVL, Instituto Miguel lillo,
Tucumán, Argentina; PVSJ, Instituto y Museo de Ciencias Naturales,
San Juan, Argentina; TMM, Vertebrate Paleontology Laboratory, Texas
Natural Science Center, Austin, Texas; UFSM, Coleção de
Paleontologia, Laboratório de Estratigrafia e Paleobiologia,
Universidade Federal de Santa Maria, Brazil; ULBRA, Museu de
Ciências Naturais, Universidade Luterana do Brasil, Canoas, Brazil.
2. Geological Setting
The specimen was collected at Cerro da Alemoa site (Fig. 3A – D),
also known as Waldsanga or Sanga do Mato, located at the urban area
of Santa Maria city (29°41′51.0″S; 53°46′26.5″W), Rio Grande do Sul
state, southern Brazil (Da-Rosa, 2015). This site was radioisotopically
dated as 233.23 ± 0.73 Ma (Fig. 3E), mid Carnian in age (Langer et al.,
2018), which had so far yielded several noteworthy vertebrate fossil
specimens (Fig. 4), emphasizing the type-series of the sauropodomorph
dinosaur Saturnalia tupiniquim (Langer et al., 1999), the saurischian
dinosaur with putative theropod affinities Nhandumirim waldsangae
(Marsola et al., 2019), the proterochampsid archosauriform
Fig. 2. Skeletal reconstruction and silhouettes of known la-
gerpetids to scale. A – unnamed lagerpetid from Ischigualasto
Formation (PVSJ 883); B – Dromomeron romeri; C –
Lagerpeton chanarensis; D – Dromomeron gregorii; E –
Dromomeron gigas; F – The novel lagerpetid herein presented
(UFSM 11611); G – Ixalerpeton polesinensis.
M.S. Garcia, et al.
Cerritosaurus binsfeldi (Price, 1946), the probainognathian cynodont
Alemoatheruim huebneri (Martinelli et al., 2017a), and the traverso-
dontid cynodont Gomphodontosuchus brasiliensis (Huene, 1928). Nearby
outcrops belonging to the same complex yielded the type-specimens of
the pseudosuchian Rauisuchus tiradentes (Zahn Sanga/Sanga do Dente -
Lautenschlager and Rauhut, 2014), the proterocampsid archosauriform
Rhadinosuchus gracilis (Ezcurra et al., 2015), and the herrerasaurid
Fig. 4. A – Taxonomic diversity of the type-specimens of named taxa from Cerro da Alemoa. B – Photograph of the main outcrops of Cerro da Alemoa where most of
the specimens were sampled.
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Journal of South American Earth Sciences 91 (2019) 302–319
Fig. 3. Geographical and geological context of the
Cerro da Alemoa site within Rio Grande do Sul state,
southern Brazil. A – Geographical location of the
Cerro da Alemoa site. B – Satellite image of Cerro da
Alemoa in Santa Maria city, showing the remaining
portion of the site, and the formerly complex of
outcrops that yielded type-specimens of Rauisuchus
tirandentes and Staurikosaurus pricei. C – Photograph
of (partial) Cerro da Alemoa, showing the channel
and crevasse deposits (CH + CR) of the Caturrita
Formation, and the distal (FFd) and proximal (FFp)
floodplain deposits of the Santa Maria Formation,
and indicating the level of UFSM 11611.
Paleontologists (on the left side) for scale. D –
Stratigraphic profile of both Caturrita and Santa
Maria Formations at Cerro da Alemoa, indicating the
level of UFSM 11611. E – Geologic time table of the
Triassic Period, emphasizing the recent radio-
isotopical dating of Cerro da Alemoa.
dinosaur Staurikosaurus pricei (Sanga Grande/Sanga de Baixo - Colbert,
1970), although nowadays most of the original complex has been
buried due to urban expansion, isolating or even destroying some
outcrops (Da-Rosa, 2018).
Additionally, specimens commonly collected in this site correspond
to indeterminate archosauriforms, cynodonts, dinosaurs, and rhyncho-
saurs (e.g. Da-Rosa et al., 2016). The latter is the most abundant faunal
M.S. Garcia, et al.
Fig. 5. Photographs at Cerro da Alemoa, exhibiting the upper, intermediate,
and lower levels of the site. A – Photograph at the upper level of Cerro da
Alemoa, interpreted as a proximal floodplain (FFp). In the background, inter-
mediate and lower levels of Cerro da Alemoa, both interpreted as distal
floodplains (FFd). B – Photograph at the lower level of Cerro da Alemoa,
showing the abundance of carbonatic concretions. Humans for scale.
element, represented by the presence of the genus Hyperodapedon
(Langer et al., 2007), allowing, as stated above, the assignment of this
site to the Hyperodapedon Assemblage Zone (AZ), corresponding to the
lower to middle portion of the Candelária Sequence (Horn et al., 2014),
which includes the upper portion of the Santa Maria Formation, in
which the Cerro da Alemoa is type-locality (Andreis et al., 1980; Da-
Rosa, 2015).
The outcrops of the Cerro da Alemoa complex, as several other sites
of the Hyperodapedon AZ, are lithologically composed by ravines of
reddish massive to laminated mudstones and fine to conglomeratic
sandstones, interpreted as floodplain deposits (Da-Rosa, 2015). These
are subdivided into upper, intermediate and, lower levels, in which the
upper level contacts the yellowish stratified sandstone typical of the
upper portion of the Candelária Sequence, which includes the Caturrita
Formation outcrops (Da-Rosa, 2015). The upper level represents a
proximal floodplain (Fig. 5A), whereas the intermediate and lower le-
vels correspond to distal floodplain deposits (Fig. 5B) (Holz and Souto-
Ribeiro, 2000; Zerfass et al., 2003; Da-Rosa, 2015). Sedimentological
and post-depositional features, along with paleoecological inferences
suggest that these alluvial sediments were deposited by meandering
fluvial systems, characterizing a semi-arid climate, with presence of
seasonal rainfalls that produced the floodplains comprising these strata
(Holz and Souto-Ribeiro, 2000; Zerfass et al., 2003; Horn et al., 2014;
Da Rosa, 2015).
Comparison with the lower portion of the Argentinean Ischigualasto
Formation (i.e. Scaphonyx-Exaeretodon-Herrerasaurus AZ), dated as
231.4 ± 0.3 Ma (Martínez et al., 2011, 2013), indicates that the faunal
content of both units is similar. Nevertheless, based upon radioisotopic
dating, the Brazilian Cerro da Alemoa site and its correlated deposits
from the Hyperodapedon AZ seem to be slightly older and better asso-
ciated to the upper portion of the Los Chañares Formation (i.e. Masse-
tognathus-Chanaresuchus AZ), which is earliest-mid Carnian in age, and
dated as 233.7 ± 0.4 Ma (Ezcurra et al., 2017). However, the faunistic
components of both strata (Hyperodapedon AZ and Massetognathus-
Chanaresuchus AZ) are quite different, showing that there is a faunistic
discrepancy in the assemblages of southern Brazil and western
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Journal of South American Earth Sciences 91 (2019) 302–319
Argentina, respectively (see Discussion).
3. Material and methods
3.1. Material
UFSM 11611 is a partial left femur with about 22 mm in preserved
length and 15 mm in distal transverse width. It preserves only the distal
end and lacks part of the crista tibiofibularis. It is housed at the pa-
leovertebrates collection of Laboratório de Estratigrafia e Paleobiologia
of the Universidade Federal de Santa Maria (UFSM).
3.2. Locality and horizon
The specimen comes from the Cerro da Alemoa site (29°41′51.0″S;
53°46′26.5″W), Santa Maria, Rio Grande do Sul state, Brazil; Alemoa
Member (Da-Rosa, 2015), Santa Maria Formation (Andreis et al., 1980);
Candelária Sequence (Horn et al., 2014); Hyperodapedon Assemblage
Zone (Langer et al., 2007), mid Carnian (233.23 ± 0.73 Ma - Langer
et al., 2018), Late Triassic.
3.3. Datasets and procedure
UFSM 11611 was scored in three different morphological datasets
from: Cabreira et al. (2016), Langer et al. (2017b), and Nesbitt et al.
(2017). In addition, we followed the modifications by Müller et al.
(2018a) on these datasets. Although each of them reflects distinct
evolutionary hypotheses, they are based almost upon the same sample
of Triassic dinosauromorph taxa. The aforementioned datasets were
filled with additional femoral characters that vary among lagerpetids,
and included all lagerpetid OTUs known to date: Dromomeron gigas,
Dromomeron gregorii, Dromomeron romeri, Ixalerpeton polesinensis, La-
gerpeton chanarensis, and the unnamed lagerpetid of the Ischigualasto
Formation (PVSJ 883). Modified versions of the datasets of Cabreira
et al. (2016), Langer et al. (2017b) and Nesbitt et al. (2017) including
UFSM 11611 totalized respectively 260 characters and 46 OTUs, 462
characters and 85 OTUs, 422 characters and 89 OTUs.
We removed Agnosphitys cromhallensis from the analysis of the da-
taset of Langer et al. (2017b) due to its fragmentary condition and
floating phylogenetic position (e.g. Martínez et al., 2013; Langer et al.,
2017b). The datasets were processed with the software TNT v1.5 (Go-
loboff et al. 2008). Additive (i.e. ordered) characters were treated as in
the original datasets, and all (ordered and unordered) received the
same weight. Euparkeria capensis was constrained to root the most
parsimonious trees (MPTs) in the analyses of the datasets of Cabreira
et al. (2016) and Langer et al. (2017b), and Mesosuchus browni was
constrained to root the MPTs in the analysis of the dataset of Nesbitt
et al. (2017). All MPTs were recovered via “traditional search” [random
addition sequence + tree bisection reconnection (TBR)], with 5000
replicates of Wagner trees (random seed = 0), TBR and branch swap-
ping, saving 20 trees save per replicate.
Additionally, we performed tree constrained analyses with the da-
taset of Cabreira et al. (2016), using the same parameters, in order to
access the required number of steps to recover UFSM 11611 as sister
group of other Carnian lagerpetids due to geographical and strati-
graphic closeness: (1) UFSM 11611 + Ixalerpeton polesinensis; (2) UFSM
11611 + Lagerpeton chanarensis; (3) UFSM 11611 + PVSJ 883.
4. Systematic paleontology
Archosauria Cope, 1869
Ornithodira Gauthier, 1986
Dinosauromorpha Benton, 1985
Lagerpetidae Arcucci, 1986
M.S. Garcia, et al. Journal of South American Earth Sciences 91 (2019) 302–319

Fig. 6. Left femur of the unnamed lagerpetid from Cerro da Alemoa (UFSM 11611). A – Medial view; B – Caudal view; C – Lateral view; D – Cranial view; E – Distal
view; F – Emphasis in the parallel striations in the bone surface.

4.1. Description and comparison
In the comparative description, if only Dromomeron is mentioned,
the cited condition or feature is present in the three species of
Dromomeron (D. gigas, D. gregorii, and D. romeri), unless commented
otherwise.
The femoral morphology resembles that of lagerpetid dinosaur-
omorphs (Figs. 6 and 7), mostly Lagerpeton chanarensis, Ixalerpeton
polesinensis, and PVSJ 883, and differs from other non-lagerpetid
dinosauromorphs mainly regarding the presence of an expanded crista
tibiofibularis (fibular condyle for some authors) and a deep inter-
condylar groove. As in all lagerpetids, the cranial margin of the distal
end of the femur is concave in UFSM 11611 (Figs. 6E and 7). Moreover,
the distal condyles are transversely wide relative to their maximum
craniocaudal depth, similar to the condition found in Ixalerpeton pole-
sinensis, Dromomeron romeri, and small specimens of Dromomeron gre-
gorii (e.g. TMM-31100-764) (Figs. 6E and 7), which contrasts with other
dinosauromorphs, such as Marasuchus lilloensis, herrerasaurids,

Fig. 7. Line drawings of select dinosauromorph left (or reversed right) femora in distal view. A – unnamed lagerpetid from Cerro da Alemoa (UFSM 11611); B –
Ixalerpeton polesinensis (ULBRA-PVT 059); C – Lagerpeton chanarensis (PVL 4619); D – Dromomeron gregorii (TMM-31100-764) reversed; E – Dromomeron gregorii
(TMM-31100-1306) reversed; F – Dromomeron romeri (GR 218); G – unnamed lagerpetid from Ischigualasto Formation (PVSJ 883); H – Dromomeron gigas (PVSJ 898);
I – Marasuchus lilloensis (PVL 3871); J – Silesaurus opolensis (ZPAL Ab III/361/23 – Dzik, 2003); K – Tawa hallae (GR 244); L – Coelophysis bauri (NMMNH P-29046 –
Spielmann et al., 2007) reversed; M – Buriolestes schultzi (CAPPA/UFSM 0035) reversed; N – Staurikosaurus pricei (MCZ 1669); O – Herrerasaurus ischigualastensis
(PVSJ 373 – Novas, 1994). Abbreviations: ctf – crista tibiofibularis, f – flange, lc – lateral condyle, mc – medial condyle, pf – popliteal fossa.

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M.S. Garcia, et al.
Fig. 8. Phylogenetic affinities of UFSM 11611. A, B1, and C1 – Strict consensus tree of the analysis employing the datasets of Cabreira et al. (2016), Langer et al.
(2017b), and Nesbitt et al. (2017), respectively. B2–B4 and C2–C7 – Alternative hypotheses for the inner relations of Lagerpetidae based on the results of the analyses
employing the datasets of Langer et al. (2017b) and Nesbitt et al. (2017), respectively.
silesaurids and dinosaurs, where the distal condyles are craniocaudally
deeper than transversely wide (e.g. Sereno and Arcucci, 1994a; Novas,
1994, Griffin and Nesbitt, 2016b; Pinheiro, 2016). In medial and lateral
views, the preserved portion of the femoral shaft is mostly slender
(Fig. 6A, C). Indeed, UFSM 11611 present a sharp medial surface
(Fig. 6A), which may represent a craniocaudally oriented sedimentary
compression, though it did not distort the morphology of the distal
condyles.
The crista tibiofibularis is partially fragmented (Fig. 6B–C, E).
Nevertheless, it is noticeable that the crista tibiofibularis of UFSM
11611 is transversely expanded relative to its maximum craniocaudal
depth, as typical of lagerpetids, especially similar to the rounded shape
seen in PVSJ 883, differing from Dromomeron, in which the crista ti-
biofibularis is subretangular, and from Lagerpeton chanarensis, in which
it is subquadrangular. An intercondylar groove excavates the femur
proximodistally, and separates the crista tibiofibularis from the lateral
condyle. Due to the fragmentation of the crista tibiofibularis it is not
possible to determine the exact form of the groove in distal view. Still,
its excavation on the lateral condyle appears to be comparable to those
of Dromomeron romeri and Dromomeron gigas.
In distal view, the medial (or tibial) condyle occupies about half of
the transverse width of the femur (Fig. 6E). Similar to PVSJ 883 and
Lagerpeton chanarensis, the medial condyle is rounded, differently from
the ovoid and subquadrate shape of Ixalerpeton polesinensis and Dro-
momeron gregorii, respectively. Moreover, in UFSM 11611 the medial
condyle is deflected medially, as in Ixalerpeton polesinensis and PVSJ
883. As in Lagerpeton chanarensis and PVSJ 883, the popliteal fossa
between the medial condyle and the crista tibiofibularis, in the caudal
margin of the femur, is faint and shallow. In addition, the fossa differs
from the condition observed in several silesaurids (e.g. Asilisaurus
kongwe, Sacisaurus agudoensis, Silesaurus opolensis), where it extends
further proximally along the length of the bone (Nesbitt et al., 2010).
UFSM 11611 does not present a protruded shape (i.e. flange) in the
craniomedial margin of the femur (Irmis et al., 2007), a condition
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Journal of South American Earth Sciences 91 (2019) 302–319
shared with Lagerpeton chanarensis, Ixalerpeton polesinensis, and some
specimens of Dromomeron gregorii among lagerpetids (Figs. 6E and 7).
This corner (craniomedial margin) of the distal end of the femur is
roughly rounded, forming a more acute angle as in Ixalerpeton pole-
sinensis. The lateral condyle of UFSM 11611 seems to be smaller than
the crista tibiofibularis, and is more laterally oriented than that of
Dromomeron romeri. Distally, UFSM 11611 is straight, bearing a quite
shallow distal groove, contrasting with PVSJ 883.
Regarding taphonomic and additional remarks, although frag-
mented in some areas, the external bone surface of UFSM 11611 is well
preserved and the specimen does not present significant diagenetic
morphological alterations (Fig. 6). Similar to an immature specimen of
Pampadromaeus barberenai (CAPPA/UFSM 0028 - Müller et al., 2017b),
an early-diverging sauropodomorph dinosaur, the external bone surface
of UFSM 11611 is remarkably marked by longitudinal parallel striations
(Fig. 6F), which seems to be unrelated to typical muscle scars (e.g.
rugosities or tuberosities), being otherwise an indicative of low bone
maturation, resembling the condition observed in the bone surface of
immature ornithodirans (Bennett 1993, 2015; Tumarkin-Deratzian
et al., 2006; Holliday et al., 2010; Rauhut et al., 2012; Watanabe and
Matsuoka, 2013). A similar patch of striations is also observable on the
homologous surface of the femora of Ixalerpeton polesinensis.
The femoral shaft of UFSM 11611 is hollow and ellipsoid in cross-
section, but different of PVSJ 883 and similar to Dromomeron gigas, the
medullary cavity occupies most of the volume of the shaft, resulting in
thin walls. Martínez et al. (2016) measured the cortex thickness/
minimum shaft diameter ratio of PVSJ 883 and Dromomeron romeri in
0.2, Dromomeron gigas presenting a ratio of 0.14, and UFSM 11611 a
much thinner condition, with a ratio of 0.08.
5. Results
All phylogenetic analyses recovered UFSM 11611 as a lagerpetid
dinosauromorph (Fig. 8). However, the inner affinities of Lagerpetidae
M.S. Garcia, et al.
and the phylogenetic position of UFSM 11611 was not consensual re-
garding all three datasets, although most topologies are similar to the
original studies and to the results of Müller et al. (2018a).
The analysis with the dataset provided by Cabreira et al. (2016),
modified by Müller et al. (2018a), recovered 54 MPTs of 851 steps each.
The inclusion of UFSM 11611 in this dataset did not dissolve the
stepwise relationship of lagerpetids (Fig. 8A), and aside the ingroup
relations of Lagerpetidae, the topology of the strict consensus tree re-
mained identical to those of Cabreira et al. (2016) and Müller et al.
(2018a). Our specimen was recovered basal to all other lagerpetids,
except for Lagerpeton chanarensis.
In the constrained analyses performed with the dataset of Cabreira
et al. (2016), all three resulted in one extra step to recover UFSM 11611
as sister taxon of Ixalerpeton polesinensis, Lagerpeton chanarensis and
PVSJ 883, respectively.
Afterwards, the phylogenetic analysis with the dataset provided by
Langer et al. (2017b), modified by Müller et al. (2018a), recovered
60480 MPCs of 1930 steps each. The resultant strict consensus tree
(Fig. 8B1) is equal to those recovered by Müller et al. (2018a), excepting
for the inclusion of our specimen. UFSM 11611 and Ixalerpeton pole-
sinensis form a trichotomy (see Fig. 8B2-4 for possible topologies) with a
stepwise clade composed of PVSJ 883 and Dromomeron, and the ex-
clusion of Lagerpeton chanarensis, which is recovered as the basal most
member of Lagerpetidae.
Finally, the phylogenetic analysis with the dataset provided by
Nesbitt et al. (2017), modified by Müller et al. (2018a), recovered
1400 MPCs of 1406 steps each. The resultant topology (Fig. 8C1) has
UFSM 11611 forming a polytomy (see Fig. 8C2-7 for possible topologies)
with Ixalerpeton polesinensis, Lagerpeton chanarensis and a stepwise clade
formed by PVSJ 883 and Dromomeron, similar to our analysis with the
dataset of Langer et al. (2017b). As in the analyses with the datasets of
Cabreira et al. (2016), Langer et al. (2017b), and in the analyses ori-
ginally performed by Nesbitt et al. (2017) and Müller et al. (2018a)
using this dataset, the overall topologies of the strict consensus tree
remained unchanged with the inclusion of UFSM 11611.
6. Discussion
6.1. Taxonomic and ontogenetic status of UFSM 11611
The taxonomic assignation of UFSM 11611 within Lagerpetidae
relies on the presence of a remarkably expanded crista tibiofibularis on
the distal end of the femur, a trait systematically recognized as diag-
nostic for lagerpetids (Irmis et al., 2007; Nesbitt, 2011; Müller et al.,
2018a). Similar to Müller et al. (2018a), our results were not consensual
regarding the basalmost lagerpetid, though Lagerpeton chanarensis, Ix-
alerpeton polesinensis, and UFSM 11611 are the most plausible candi-
dates.
Based upon the proportions of known lagerpetids, we estimated
about 70 mm of total femoral length to UFSM 11611, which matches
the femoral size of some of the smallest lagerpetids such as Lagerpeton
chanarensis with around 78 mm (Martínez et al., 2016) and Ixalerpeton
polesinensis with 60 mm. Indeed, relative size is not necessarily and/or
directly linked to skeletal maturity in dinosauromorphs (Piechowski
et al., 2014, Griffin and Nesbitt, 2016b; Peecook et al., 2018, Garcia
et al. in press). However, it is possible to associate some characteristics
present in UFSM 11611 to an immature ontogenetic stage, based in
growth series present in other close-related dinosauromorphs, such as
Dromomeron gregorii (Nesbitt et al., 2009a). In addition, as in juvenile
ornithodirans (Bennett, 1993; Piechowski et al., 2014; Griffin and
Nesbitt, 2016b; Müller et al., 2017b), UFSM 11611 presents a series of
parallel longitudinal striations on the femoral external surface that in-
dicate low bone maturation, in which the internal canals of the bone
would remain exposed due to a fast growth in the earlier ontogenetic
stages (Bennett 1993, 2015; Tumarkin-Deratzian et al., 2006; Rauhut
et al., 2012; Watanabe and Matsuoka, 2013).
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Journal of South American Earth Sciences 91 (2019) 302–319
In spite of the suggested immaturity of UFSM 11611, known onto-
genetic series for the lagerpetid Dromomeron gregorii (Nesbitt et al.,
2009a) and other dinosauromorphs (Piechowski et al., 2014; Griffin
and Nesbitt, 2016b; Müller, 2017; Müller et al., 2017b; Peecook et al.,
2018) provide evidence that some features in which UFSM 11611 dif-
fers from other lagerpetids are not essentially controlled by ontogeny
(e.g. absence of the craniomedial flange; medial condyle medially de-
flected; shallow popliteal fossa). Hence, it is more likely that our spe-
cimen does not represent a skeletally immature individual of any Car-
nian lagerpetid taxa known to date. Therefore, UFSM 11611 is
considered as a new dinosauromorph morphotype, actually belonging
to a still unknown lagerpetid taxon of the Candelária Sequence, Late
Triassic of southern Brazil.
However, both sites (Buriol and Cerro da Alemoa) that yielded
Brazilian lagerpetids (Ixalerpeton polesinensis and UFSM 11611) share
the presence of the rhynchosaur Hyperodapedon (see Müller et al.,
2017a), and are considered coeval, though regarding other tetrapods
there are little overlapping faunal content. Furthermore, the Cerro da
Alemoa is more taxonomically diverse (see Geological Setting) than the
Buriol site, which yielded isolated archosauriform teeth, the aetosaur
Polesinesuchus aurelioi (Roberto-Da-Silva et al., 2014), as well as the
sauropodomorph Buriolestes schultzi (see further below).
6.2. Diversity of dinosauromorphs in the Cerro da Alemoa site
Historically, the Cerro da Alemoa site and its complex of outcrops
(e.g. Sanga Grande/Sanga de Baixo, type-locality of Staurikosaurus
pricei) has been of great importance to investigate the Triassic biota of
southern Brazil, and consequently south Pangea, as it yields type-spe-
cimens of several Triassic vertebrates such as non-dinosauromorph
archosauriforms and cynodonts (Huene, 1928, Price, 1946,
Lautenschlager and Rauhut, 2014; Martinelli et al., 2017a). Moreover,
this site provided an extensive sample of rhynchosaurs and a wide
variety of dinosauromorphs (Fig. 9), mainly represented by saurischian
dinosaurs (Colbert, 1970; Langer et al., 1999). Even though these out-
crops provided type-specimens of many dinosaurs such as Nhandumirim
waldsangae (Marsola et al., 2019), Saturnalia tupiniquim and Staur-
ikosaurus pricei, latest intensive fieldwork in this locality and reassess-
ment of already collected material revealed a hidden diversity of di-
nosauromorph groups (dinosaurs, silesaurids and lagerpetids) that is
only comparable to faunal assemblages in (younger) horizons from the
Ischigualasto Formation of Argentina and from the Hayden Quarry of
southwestern USA (see Irmis et al., 2007; Martínez et al., 2013).
Overall, recent expeditions in Upper Triassic beds from southern Brazil
are unearthing an outstanding diversity of dinosauromorphs, that is still
Fig. 9. Quantitative sampling of dinosauromorphs collected at Cerro da
Alemoa. A – Overall proportion of the major clades of dinosauromorphs
(Lagerpetidae, Silesauridae, Dinosauria). B – Quantity of specimens per major
clade [Herrerasauridae, Theropoda (putative affinity), and Sauropodomorpha]
within Dinosauria.
M.S. Garcia, et al.
relatively scarce in this area, in comparison to other vertebrate groups
such as cynodonts and rhynchosaurs, with exceptionally well-preserved
and reasonably complete specimens (Cabreira et al., 2011, 2016; Müller
et al., 2018b, 2018d; Pretto et al., 2018). New dinosauromorph speci-
mens of Cerro da Alemoa are mostly composed of material still under
study and laboratorial preparation (e.g. Garcia et al., 2018), but pre-
liminary assessments show a wide range of saurischian dinosaurs,
mainly including sauropodomorphs and herrerasaurids. As mentioned
above, fieldwork at Cerro da Alemoa site and revaluation of its speci-
mens are shedding light in a much rich paleofauna of dinosauromorphs
for this site, as it now bears not only saurischian dinosaurs, but also
lagerpetids (UFSM 11611) and silesaurids (Langer et al., 2017a).
New sauropodomorphs from the Cerro da Alemoa range from spe-
cimens smaller and larger (e.g. UFSM 11612) than the average size of
the type-series of Saturnalia tupiniquim, together with several other
specimens currently under study and to be presented elsewhere. UFSM
11612 is the most fragmentary of the specimens, comprising a partial
left ilium (Müller et al., 2018c) and two articulated caudal vertebrae,
which is indistinguishable from Saturnalia tupiniquim, and preliminary
classified as cf. Saturnalia. Additionally, two herrerasaurids from this
site comprise partial skeletons larger than the holotype of Staur-
ikosaurus pricei (e.g. Garcia et al., 2018). For instance, the largest spe-
cimen (UFSM 11330, under study) is about 40% larger than the holo-
type of Staurikosaurus pricei (Colbert, 1970; Bittencourt and Kellner,
2009), although the ingroup affinities of this specimen are still under
assessment. Regarding silesaurids, the Cerro da Alemoa yields frag-
mentary remains of a single specimen (UFSM 11529) also under study,
comprising many small fragments, including a tooth-bearing bone and a
left femur, which allows the putative assignment of UFSM 11529 to
Silesauridae (Langer et al., 2017a). The presence of a lagerpetid (UFSM
11611) in the Cerro da Alemoa site, which was recently radio-
isotopically dated by Langer et al. (2018) renders it as the oldest di-
nosaur-bearing fossiliferous locality, that also bears the oldest local
clustering of dinosaurs, silesaurids and lagerpetids. This unique co-oc-
currence of three major dinosauromorph clades in the same layer of
outcrops, was never recorded in any other Triassic locality elsewhere
before, especially considering that the specimens were unearthed in a
few square meters of distance, although in different excavations. Pre-
viously, Cabreira et al. (2016) reported a co-occurrence of lagerpetids
and sauropodomorphs in a unique Carnian assemblage from southern
Brazil, but no silesaurids where excavated from this outcrop. Indeed,
excepting UFSM 11529, so far, the only putative record of a silesaurid
from the Hyperodapedon AZ in Brazil comprises a heavily fragmentary
femur (UNIPAMPA-0632) from the Janner site (Müller et al., 2014a, see
further below). Therefore, the faunal composition of the Cerro da
Alemoa site supports that all these clades shared the same land eco-
systems during the rise of dinosaurs and untill the demise of non-di-
nosaur dinosauromorphs (e.g. Irmis et al., 2007), a time interval of
approximately 21 Ma (Langer et al., 2018). During this time interval,
lagerpetids reached a wide geographical distribution (Müller et al.,
2018a) and increased their body size (Martínez et al., 2016), while
dinosaurs adopted distinct feeding strategies and evolved into the three
major clades (Cabreira et al., 2016).
6.3. Brazilian dinosauromorph-bearing Late Triassic beds:
biostratigraphical framework
Biostratigraphic correlations of the Hyperodapedon AZ with the
Scaphonyx-Exaeretodon-Herrerasaurus AZ (Ischigualasto Formation)
from Argentina so far suggested that the lower portion of the Candelária
Sequence (upper portion of the Santa Maria Formation), was latest
Carnian in age, around 231 Ma (Langer et al., 2007; Martínez et al.,
2011, 2013; Da-Rosa, 2015). Recently, Langer et al. (2018) U-Pb age
constraints recovered an older age to the Cerro da Alemoa, of about
233 Ma, extended to the whole coeval portion of the Candelária Se-
quence (Fig. 10). This age is comparable to the Argentinean
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Journal of South American Earth Sciences 91 (2019) 302–319
Massetognathus-Chanaresuchus AZ (upper portion of the Los Chañares
Formation), which yielded many dinosauromorphs, and is considered
earliest-mid Carnian (Ezcurra et al., 2017). It is noteworthy that Langer
et al. (2018) dated the upper level of the Cerro da Alemoa site, and
considered that the lower portion of this unit on the Hyperodapedon AZ
(which corresponds to the level that yielded Staurikosaurus pricei)
should be even older, when taking into account that the Hyperodapedon
AZ is preceded by the Santacruzodon AZ (Santa Cruz Sequence), which
was dated with a maximum age of deposition of 237 Ma (Philipp et al.,
2018). However, the faunal content of the Santacruzodon AZ are cor-
related to the upper portion of the Los Chañares Formation (i.e. Mas-
setognathus-Chanaresuchus AZ), and so far, has not yielded any dino-
sauromorphs. Moreover, the Dinodontosaurus AZ (Pinheiros-Chiniquá
Sequence) correlates to both lower and upper portions of the Los
Chanãres Formation (i.e. Tarjadia and Massetognathus-Chanaresuchus
AZs respectively), and present evidence that it is subdivided into two
distinct faunal assemblages, possibly ranging from Anisian to earliest
Carnian in age, mainly because of the presence of the rhynchosaur
Brasinorhynchus mariantensis, and the cynodonts Luangwa sudamericana
and Aleodon cromptoni (see Abdala and Teixeira, 2004; Langer et al.,
2007; Schultz et al., 2016; Ezcurra et al., 2017; Martinelli et al., 2017b).
These biostratigraphical discrepancies could be the result of specific
issues (e.g. lack of absolute dating in several Brazilian Triassic beds,
such as those of the Dinodontosaurus AZ; intensive fieldwork in beds
belonging to AZs such as Hyperodapedon, in detriment of less prospected
ones like Santacruzodon), as well as other causes, such as methodolo-
gical biases or perhaps the Brazilian AZs are, in fact, slightly older than
those from Argentina. The Massetognathus-Chanaresuchus AZ yielded
many dinosauromorphs (see below), and conversely, the Santacruzodon
AZ does not present any record of dinosauromorphs, although this AZ is
not as intensively prospected as the Hyperodapedon AZ (Philipp et al.,
2018). Yet, the Santacruzodon AZ is an attention-grabbing zone for the
discovery of both early dinosaurs and/or non-dinosaurian dinosaur-
omorphs (idem for the Dinodontosaurus AZ), once that the correlated
Argentinean beds provide such records (e.g. Lagerpeton chanarensis,
Lewisuchus admixtus) and younger Brazilian strata (Hyperodapedon and
Riograndia AZs) present a reasonable dinosauromorph diversity.
On the other hand, while the Hyperodapedon AZ is correlated to the
Scaphonyx-Exaeretodon-Herrerasaurus AZ (Martínez et al., 2013), one of
the most conspicuous and better sampled sites of the Candelária Se-
quence, the Janner site (= Várzea do Agudo), has a high incidence of
the traversodontid cynodont Exaeretodon riograndensis, although it also
bears hyperodapedontin rhynchosaurs, but with a lesser abundance
(Abdala et al., 2002; Da-Rosa, 2015). The vast presence of Exaeretodon
in the Janner site may indicate that this site is younger than other sites
from the Hyperodapedon AZ (e.g. Cerro da Alemoa), and is therefore
correlated to the Exaeretodon AZ from the Ischigualasto Formation of
Argentina, which overlaps the Scaphonyx-Exaeretodon-Herrerasaurus AZ,
in which Exaeretodon is also abundant in detriment of rhynchosaurs
(Langer et al. 2010, Martínez et al., 2013; Da-Rosa, 2015; Müller et al.,
2015b). Notably, while Exaeretodon is significantly abundant in the
Janner site, other traversodontid cynodonts as a whole are scarce in
other localities of the Hyperodapedon AZ (but see Pavanatto et al.,
2018), although their records are quite recurrent in older strata of both
Dinodontosaurus and Santacruzodon AZs (Abdala et al., 2001, Reichel
et al. 2009). This particular inconsistency does not seem to result of
sampling biases, once several localities of the Hyperodapedon AZ, as
comparatively, both Buriol and Cerro da Alemoa (including Janner)
sites, were extensively prospected in the last decade. Regarding dino-
sauromorphs, the Janner site has provided an interesting diversity,
though archosauriforms are not typically as common as specimens of
Exaeretodon riograndensis, and so far, this site yielded specimens of
Pampadromaeus barberenai, Bagualosaurus agudoensis, a putative sile-
saurid (UNIPAMPA-0632), and other fragmentary specimens (Cabreira
et al., 2011; Müller et al., 2014a, 2015a; 2015b; Pretto et al., 2015,
2018). The sauropodomorph content present at Janner Site is consistent
M.S. Garcia, et al.
with a younger age (regarding the rest of the Hyperdodapedon AZ), since
these are more advanced than other forms such as Buriolestes schultzi
and Saturnalia tupiniquim (see Müller et al., 2018b, 2018d; Pretto et al.,
2018) (Fig. 10). For instance, while Buriolestes schultzi and Saturnalia
tupiniquim both have a ziphodont dentition (Cabreira et al., 2016;
Bronzati et al., 2017), compatible with a faunivorous diet (see further
below), Bagualosaurus agudoensis and Pampadromaeus barberenai have a
lanceolate dentition, which is regarded to an omnivorous/herbivorous
diet. Moreover, Bagualosaurus agudoensis already present a small head
in relation to the femur (less than 2/3 of the femoral length), a trait
usually associated with younger sauropodomorphs (see Müller et al.,
2018d; Pretto et al., 2018). Nonetheless, Bagualosaurus agudoensis is, so
far, the largest known Carnian sauropodomorph, reaching a femoral
length of ca. 215 mm, while other sauropodomorphs of the Hyper-
odapedon AZ range from ca. 110–150 mm (e.g. Langer, 2003, Müller
et al. 2018). Among Carnian dinosauromorphs, the size of Bagual-
osaurus agudoensis is comparable to that of herrerasaurids (e.g. Novas,
1994; Bittencourt and Kellner, 2009; Pacheco et al., 2014). Further-
more, a younger age for the Janner site would not only elucidate why
traversodontid cynodonts (Exaeretodon) are largely abundant in this
site, but mostly absent in other sites of the Hyperodapedon AZ. More-
over, such younger age helps filling and tracing the morphological
evolution of the Brazilian Triassic sauropodomorphs, from typically
Carnian (i.e. “Eoraptor-like”) forms in other sites of the Hyperodapedon
AZ (e.g. Buriol, Cerro da Alemoa, Piche– Langer et al., 1999; Cabreira
et al., 2016; Müller et al., 2018b, Garcia et al. in press), followed by
“intermediate taxa” from the Janner (Cabreira et al., 2011; Müller et al.,
2014a; Pretto et al., 2018), and finally to more advanced forms from the
Riograndia AZ (Leal et al., 2004; Müller et al., 2014b, 2016, 2018d).
Indeed, the presence and extensive geographical distribution of
advanced sauropodomorphs in many localities of the Riograndia AZ
could indicate that these might also be used as index fossils for the
recognition of this AZ within the stratigraphic horizon of the Caturrita
Formation, upper portion of the Candelária Sequence (Bittencourt et al.,
2012a, 2012b; Müller et al., 2014b, 2016, 2018d). In any case, the
abundant record of Exaeretodon in detriment of rhynchosaurs (espe-
cially Hyperodapedon) suggests that the former replaced the latter as a
primary consumer on the terrestrial ecosystems, and that it may have
had an extended temporal occurrence (Ribeiro et al., 2011) than pre-
viously thought, which is reflected in the common dominance of Ex-
aeretodon in the Argentinean Exaeretodon AZ (whereas rhynchosaurs are
completely absent – Martínez et al., 2013), and in the Brazilian Janner
site. Following this, it is increasingly evident that the Hyperodapedon AZ
could be subdivided into two faunal assemblages (Fig. 10). The older
(lower) bearing Hyperodapedon as index fossil (i.e. Hyperodapedon Acme
Zone of Langer et al., 2007) and the younger (upper) bearing
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Journal of South American Earth Sciences 91 (2019) 302–319
Fig. 10. Chronostratigraphy of vertebrate assem-
blage zones of the Santa Maria Supersequence based
in maximum ages from radioisotopic dating. Ages in
orange indicate radioisotopic dating of Brazilian
strata (see text for references). AZ – Assemblage
Zone. (For interpretation of the references to colour
in this figure legend, the reader is referred to the
Web version of this article.)
Exaeretodon + Hyperodapedon as index fossils, similar to the Ischi-
gualastian AZs, though providing further assessments in this matter is
beyond the scope of this paper. It is worth reminding that this sub-
division is based on the relative abundance of Hyperodapedon and on
the presence/absence of Exaeretodon, since this cynodont is absent in
the “lower Hyperodapedon AZ”. Additionally, a newly reported locality
(nearby the Janner site) with a remarkable presence of the traverso-
dontid cynodont Siriusgnathus niemeyerorum launches more questions
about the biostratigraphy of the Candelária Sequence (Pavanatto et al.,
2018). Despite the fact that Niemeyer site has been heavily sampled in
the last few years, it so far did not provide any index fossil typical of the
already well-established Triassic AZs of the Candelária Sequence (Hy-
perodapedon, Riograndia), although its lithology is compatible with beds
of the Hyperodapedon AZ. At this point, the Niemeyer site has provided
a single fragmentary dinosauromorph specimen, which is regarded as a
new morphotype. Furthermore, the lack of index fossils prevents a more
robust biostratigraphical framework for this locality (Pavanatto et al.,
2018).
6.4. South American Triassic diversity of dinosauromorphs
Investigation of Triassic land ecosystems of South America is vital
for a comprehensive understanding of the dawn of the dinosaurs, a key
event in the evolutionary history of the bird line archosaurs (Langer
et al. 2010; Nesbitt et al., 2017). Hence, south Pangea is regarded as the
birthplace of this conspicuous clade (Dinosauria – Lee et al., 2018;
Marsola et al., 2018), since basalmost unequivocal dinosaurs have been
unearthed in continental tetrapod-bearing localities from both Argen-
tina and Brazil, dating back to the Carnian (roughly 230 Ma) epoch
(Langer et al., 2007; Martínez et al., 2013; Cabreira et al., 2016). De-
spite the fact that dinosauromorphs are not the most common faunal
component during the Middle (Ladinian) – initial Late Triassic (Car-
nian) of South America, they are relatively well sampled in the Can-
delária Sequence (Fig. 11), and both Los Chañares and Ischigualasto
Formations (Langer, 2014). In fact, dinosauromorphs as a whole were
not as diverse and abundant in the Ladinian/Carnian – early Norian as
they were in the late Norian onwards (Langer et al. 2010; Ezcurra,
2012; Langer, 2014). As in the lower portion of the Ischigualasto For-
mation (Carnian, Scaphonyx-Exaeretodon-Herrerasaurus and Exaeretodon
AZs), the dinosauromorph fauna from the lower portion of the Cande-
lária Sequence (Carnian, Hyperodapedon AZ) is mainly composed of
early-diverging sauropodomorphs and herrerasaurids, whereas la-
gerpetids, silesaurids, and putative theropods and ornithischians also
have been recorded, but in a lesser extent (Langer et al., 2007; Martínez
et al., 2013).
Lagerpetids so far comprise Ixalerpeton polesinensis (Cabreira et al.,
M.S. Garcia, et al. Journal of South American Earth Sciences 91 (2019) 302–319

Fig. 11. Geographical distribution of Dinosauromorph specimens in the central region of Rio Grande do Sul state, showing the overall location of dinosauromorph-
bearing outcrops/sites, of the Candelária Sequence (upper Santa Maria and Caturrita Formations). Silhouettes represent the main dinosauromorph specimens
recorded for each location (see Discussion). Small circles of some of the localities were suppressed due to scaling, with dashed lines connecting to the geographically
closest site. Large circles of outcrops of the same site complex are connected by full lines.

2016) and UFSM 11611 (see above). While Ixalerpeton polesinensis is, so
far, the most complete specimen of a lagerpetid dinosauromorph
worldwide, UFSM 11611 is very fragmentary (comprised only by an
incomplete left femur). However, it has a considerable relevance re-
garding the diversity of dinosauromorphs in the Cerro da Alemoa site
and in the Hyperodapedon AZ of the Candelária Sequence as a whole,
showing that lagerpetids were even more diverse in the Brazilian as-
semblages than previously known. In addition to the diversity of non-
dinosaurian dinosauromorphs from the Hyperodapedon AZ, the sile-
saurids are, so far, limited to two fragmentary specimens (Müller et al.,
2014a; Langer et al., 2017b). This situation is similar to the records of
both lagerpetids and silesaurids in Carnian beds from Argentina (both
Los Chañares and Ischigualasto Formations), where lagerpetids (La-
gerpeton chanarensis and PVSJ 883) and silesaurids (Ignotosaurus fragilis
and Pisanosaurus mertii – see Agnolin and Rozadilla, 2017) are more
scarce than dinosaurian dinosauromorphs, such as sauropodomorphs
and herrerasaurids (Sereno and Arcucci, 1994a; Martínez et al., 2013).
In addition, there is a vast taxonomic abundance of dinosaur-
omorphs in the upper Los Chañares Formation (earliest Carnian –
Massetognathus-Chanaresuchus AZ), mainly represented by dinosauri-
forms. Besides Lagerpeton chanarensis, dinosauromorphs of these
Argentinean beds include: Marasuchus lilloensis, usually regarded as the
basalmost dinosauriform (Sereno and Arcucci, 1994b); Lagosuchus ta-
lampayensis, a poorly preserved dinosauromorph skeleton, which hin-
ders any further considerations (Romer, 1971, Sereno and Arcucci,
1994a); Lewisuchus admixtus, a dinosauriform possibly related to Sile-
sauridae (Romer, 1972; Nesbitt, 2011; Bittencourt et al. 2015; Cabreira
et al., 2016); and Pseudolagosuchus major, a problematic dinosauriform,
that has been considered a synonym of Lewisuchus admixtus, however,
more material is necessary to test this proposition, as both taxa share a
small amount of overlapping material (Arcucci, 1987; Nesbitt et al.,
2010; Baron et al., 2017; Langer et al., 2017b). Even if Lewisuchus ad-
mixtus and Pseudolagosuchus major represent the same taxon, it is re-
markable that the Los Chañares Formation yielded so many dinosaur-
omorphs, especially considering that the chronocorrelated
Dinodontosaurus and Santacruzodon AZs of southern Brazil, so far, have
no records of unequivocal dinosauromorphs. Indeed, Spondylosoma
absconditum from the Dinodontosaurus AZ was regarded as either a
saurischian dinosaur or an indeterminate dinosauriform (Huene, 1942;
Langer, 2004), though Nesbitt et al. (2017) recovered this taxon as the
youngest representative of the Aphanosauria, a group of basal aveme-
tatarsalians.
Differently from Argentinean Late Carnian beds (Scaphonyx-
Exaeretodon-Herrerasaurus and Exaeretodon AZs), where herrerasaurids
are the most abundant faunal component (regarding dinosauromorphs),
in the Brazilian Middle – Late Carnian beds (Hyperodapedon AZ),
sauropodomorphs are substantially more abundant and quantitatively
more distributed than any other dinosauromorph group, as they are by
far the most common dinosauromorphs in tetrapod-bearing localities of
the Hyperodapedon AZ (Martínez et al., 2013, Garcia et al, in Press). Not
long ago, Brazilian herrerasaurids were known mainly by the holotype
of Staurikosaurus pricei and fragmentary material (Silva et al. 2017), but
recently, novel and better-preserved specimens unearthed have the
potential to provide more data regarding the anatomy and diversifica-
tion of the Brazilian herrerasaurids (Pacheco et al., 2014; Garcia et al.,
2018). In southern Brazil, as well as everywhere else, ornithischians are
completely absent of Triassic dinosauromorph-bearing rocks hitherto,
with a single taxon described from Argentina (Pisanosaurus mertii –
Casamiquela, 1967), that historically had been regarded as the basal-
most ornithischian and later assigned to Silesauridae (Casamiquela,

312
M.S. Garcia, et al.
Fig. 12. Putative dinosauromorph dispersal through the Early-initial Late (Carnian) Triassic Period. A – Record of dinosauromorphs represented by African taxa, and
so far, restricted to south Pangea. B – Record of dinosauromorphs represented by Argentinean taxa, and so far, restricted to south Pangea. C – Dinosauromorphs are
virtually cosmopolitans in Pangaea, presenting body fossils in Laurasia, and with high taxonomic diversification.
1967; Agnolin and Rozadilla, 2017; Baron, 2017). Other studies re-
cently did recover Silesauridae within Dinosauria, as Triassic basal re-
presentatives of the clade Ornithischia (Ferigolo and Langer, 2007;
Cabreira et al., 2016).
Regarding theropods, Brazilian specimens from Carnian are so far
restricted to Nhandumirim waldsangae (Marsola et al., 2019), which is
putatively associated to this clade. Similar to silesaurids/ornithischians,
Herrerasauridae is often regarded as a basal branch of Theropoda. Yet,
the phylogenetic position of Herrerasauridae is widely debated as they
were also considered non-dinosaurian dinosauriforms, basal saurischian
dinosaurs outside the dichotomic eusaurischian branch of Theropoda
plus Sauropodomorpha, and even as the sister taxon of Saur-
opodomorpha (Sues et al., 2011; Cabreira et al., 2016; Baron et al.,
2017; Langer et al., 2017b). Finally, the Scaphonyx-Exaeretodon-Her-
rerasaurus AZ of the Ischigualasto Formation provided specimens of a
saurischian dinosaur, usually considered a non-neotheropod theropod,
Eodromaeus murphi (Martínez et al., 2011). However, recent studies
recovered this taxon as a saurischian basal to the Theropoda/Saur-
opodomorpha dichotomy (Cabreira et al., 2016; Müller et al., 2018b).
Therefore, there are no unequivocal record of theropods in these Car-
nian beds.
Norian dinosauromorphs from the Candelária Sequence (Riograndia
AZ, 225.42 ± 0.37 Ma, according to Langer et al., 2018) are mainly
represented by sauropodomorphs more derived and more abundant
than those from the Hyperodapedon AZ (Leal et al., 2004; Bittencourt
et al., 2012a, 2012b; Müller et al., 2014b, 2016; 2018d; Pretto et al.,
2016). Among them, Unaysaurus tolentinoi was usually regarded as a
plateosaurid sauropodomorph, close related to Plateosaurus engelhardti
of Germany than to sauropodomorphs present in Norian strata from
Argentina (Leal et al., 2004; Bittencourt et al. 2015; Müller et al., 2016;
313
Journal of South American Earth Sciences 91 (2019) 302–319
Baron et al., 2017). However, a newly reported sauropodomorph spe-
cies (Macrocollum itaquii) from the Riograndia AZ provides new data
about the early evolution of sauropodomorphs in the Candelária Se-
quence, especially regarding the Norian forms (Müller et al., 2018d).
Macrocollum itaquii is recovered as close related to Unaysaurus tolentinoi,
together with Jaklapallisaurus asymmetrica comprising Unaysauridae, a
newly erected clade of south Pangean non-plateosaurian saur-
opodomorphs and thus, reinforcing abovementioned biostratigraphic
insights regarding post-Carnian sauropodomorphs from Brazilian
Norian beds. Nonetheless, Argentinean Norian deposits yielded a con-
siderable diversity of medium to large-bodied advanced saur-
opodomorphs (e.g. Coloradisaurus brevis – Bonaparte, 1978, Ingentia
prima – Apaldetti et al., 2018, Lessemsaurus sauropoides – Bonaparte,
1999, Riojasaurus incertus – Bonaparte, 1967), although they are more
advanced than Brazilian forms (Leal et al., 2004; Müller et al., 2014b,
2016, 2018d), and both Los Colorados (Early – Middle Norian) and
Quebrada del Barro (Late Norian – Rhaetian) Formations are probably
younger than the Riograndia AZ (Early Norian) of the Caturrita For-
mation/upper portion of the Candelária Sequence (Colombi et al.,
2014, 2015; Kent et al., 2014; Langer et al., 2018). The record of la-
gerpetids and ornithischians is, so far, absent in the Brazilian Norian
beds, whereas Dromomeron gigas, from the Quebrada del Barro Forma-
tion, represent lagerpetids in Argentina (Martínez et al., 2016). On the
other hand, silesaurids from the Riograndia AZ are represented by
several specimens of Sacisaurus agudoensis (Ferigolo and Langer, 2007;
Langer and Ferigolo, 2013). Curiously, no silesaurid remains are re-
corded in Argentinean beds from the Norian epoch (see further below).
In contrast, Theropod records occur in a much more fragmentary way,
with a single specimen recorded, a distal end of a femur (Pinheiro,
2016). Although some authors attribute theropod affinities to
M.S. Garcia, et al.
Guaibasaurus candelariensis, this taxon has a floating phylogenetic po-
sition, being also recovered as either a sauropodomorph or a non-eu-
sarischian saurischian dinosaur (Bonaparte et al. 1999, 2007; Langer
et al., 2011, 2017b; Cabreira et al., 2016; Baron et al., 2017). Theropods
are present in both Los Colorados and Quebrada del Barro Formations,
well-represented by coelophysoid neotheropods (Arcucci and Coria,
2003; Ezcurra and Novas, 2007; Ezcurra, 2017; Martínez and Apaldetti,
2017).
6.5. Early dispersal of Triassic dinosauromorphs and their evolutionary
trends
Together, the geographic dispersal of the earliest bird-line arch-
osaurs (i.e. Aphanosauria – Nesbitt et al., 2017), ghost-lineage as-
sumption, and report of ichnofossils pull back the early members of
Dinosauromorpha to the Early Triassic (Brusatte et al., 2010a,b; Benton
et al., 2014; Nesbitt et al., 2017). However, the oldest unequivocal
skeletal remains of dinosauromorphs known to date come from the
Anisian epoch (Middle Triassic) from eastern/southeastern Pangea
(Tanzania and Zambia) (Fig. 12A), which nowadays corresponds to the
African continent (Nesbitt et al., 2010, 2017; Peecook et al., 2013,
2018, but see Martinelli et al., 2017b; Wynd et al., 2017). Although
being the oldest dinosauromorph skeletal remains worldwide, these
African forms are not considered the basalmost taxa. Actually, Asili-
saurus kongwe and Lutungutali sitwensis are usually regarded as dino-
sauriform dinosauromorphs members of Sileauridae (Nesbitt et al.,
2010, 2017; Peecook et al., 2013; Langer et al., 2017b, see also Cabreira
et al., 2016). Nevertheless, taxa regarded as the basalmost dinosaur-
omorphs come from initial Late Triassic beds of Argentina and Brazil
(e.g. Ixalerpeton polesinensis, Lagerpeton chanarensis, UFSM 11611),
which suggests that, more likely, the rise and likely parallel early ra-
diation of dinosauromorphs originated in this region of Pangaea (i.e. a
south Pangean origin for the clade – Ezcurra, 2012, Langer et al. 2010,
2013, 2017b, contra Baron et al., 2017). A conspicuous dinosauriform
of the Manda Formation of Tanzania (Nyasasaurus parringtoni) may be
the oldest dinosaurian-dinosauromorph, although its fragmentary con-
dition prevents further taxonomic assignations (Nesbitt et al., 2013, but
see Agnolin, 2016). In consequence, unequivocal records of dinosaur-
omorphs in the Ladinian epoch are absent worldwide, because dating of
the Los Chañares Formation recovered an earliest-mid Carnian age for
this unit (Marsicano et al., 2016; Ezcurra et al., 2017). Although a
portion of Triassic Brazilian strata is considered to be at least late La-
dinian in age (Pinheiros-Chiniquá Sequence, Dinodontosaurus AZ), it has
no records of unequivocal dinosauromorphs hitherto, and the lack of
absolute dating precludes refining biostratigraphical correlations
(Abdala and Teixeira, 2004; Langer et al., 2007; Schultz et al., 2016;
Ezcurra et al., 2017; Martinelli et al., 2017b).
If Early – Middle Triassic beds almost do not provide data regarding
the taxonomic diversity of dinosauromorphs, soon after the Middle-Late
Triassic boundary there is a dramatic increase in these records, not only
quantitatively, but also regarding morphological disparity (Langer et al.
2010; 2013) (Fig. 12B). While Middle Triassic dinosauromorphs were
mostly pictured by dinosauriforms related to Silesauridae (Nesbitt
et al., 2010), dinosauromorphs from initial Late Triassic, earliest Car-
nian (herein regarded by forms present in the upper Los Chañares
Formation, Massetognathus-Chanaresuchus AZ) are composed of basal
representatives of both non-dinosauriform dinosauromorph and non-
dinosaurian dinosauriform groups (Lagerpeton chanarensis, Lewisuchus
admixtus, Marasuchus lilloensis), as well as other dinosauromorphs of
uncertain affinities (Lagosuchus talampayensis, Pseudolagosuchus major),
possibly indicating that the initial geographic dispersal of early dino-
sauromorphs (latest Anisian – earliest Carnian) was restricted to the
southern portion of Pangaea (Sereno and Arcucci, 1994a,b; Ezcurra,
2012; Langer et al., 2013; Ezcurra et al., 2017). However, since Middle
Triassic localities with abundant terrestrial vertebrates are scarce, this
hypothesis is vulnerable to sampling and preservational biases.
314
Journal of South American Earth Sciences 91 (2019) 302–319
Although, terrestrial faunas from the Los Chañares Formation show that
early diversification of dinosaurs throughout the Carnian epoch was
preceded by rapid faunal turnovers at southwestern Pangea (Ezcurra
et al., 2017), that along to other well recognized events (see below)
may have paved the way for the “dinosaurian empire” established
during the Late Norian – Rhaetian, lasting approximately 150 Ma
(Archibald and Fastovsky, 2004; Brusatte et al., 2010a,b; Irmis, 2011;
Müller et al., 2016).
Still in the Carnian epoch, the so-called dawn of the dinosaur took
place, rapidly establishing some of the most important dinosauromorph
clades regarding land ecosystems, such as Herrerasauridae and
Sauropodomorpha (Novas, 1994, Langer et al., 1999; Langer et al.
2010; Cabreira et al., 2016). Despite the fact that non-archosaurian
archosauromorphs (e.g. rhynchosaurs and Tanystropheidae – Pritchard
et al., 2015; Ezcurra, 2016; Oliveira et al., 2018), pseudosuchians (e.g.
Erpetosuchidae, Aetosauria, “rauisuchians” – Nesbitt, 2011; Nesbitt
et al., 2013, Roberto-Da-Silva et al., 2014; Ezcurra et al., 2017, Biacchi-
Brust et al. 2018; Lacerda et al., 2018), and synapsids (cynodonts and
dicynodonts – Martínez et al., 2013; Martinelli et al., 2017a; Pavanatto
et al., 2018; Stefanello et al., 2018; Ugalde et al., 2018) played im-
portant roles in Triassic land ecosystems since the Permian-Triassic
extinction (Brusatte et al. 2008a,b; Ezcurra and Butler, 2018), Late
Triassic events (such as the well-known “Carnian Pluvial Event” –
Simms and Ruffell, 1989, Mueller et al., 2016, Benton et al., 2018,
Bernardi et al., 2018) and their aftermaths “offered” the opportunity for
the explosion in diversity seem in dinosauromorphs (mostly dinosaurs)
along the initial Late Triassic (Ezcurra, 2010; Ezcurra et al., 2017;
Müller et al., 2018b). While older dinosauromorphs such as Asilisaurus
kongwe may be regarded as omnivorous/generalists (Nesbitt et al.,
2010; Cabreira et al., 2016), the bauplan (and in some cases the den-
tition) of most early dinosauromorphs (e.g. Marasuchus lilloensis, Lewi-
suchus admixtus), at the exception of silesaurids, provide clues towards a
putative faunivorous diet, even though the ancestral diet of Dinosaur-
omorpha and Dinosauria is still a matter of debate (Barrett and
Rayfield, 2006, Barrett et al., 2010, Bittencourt et al. 2015; Cabreira
et al., 2016, Bronzati et al., 2017). In any case, the environmental shifts
occurred during the Carnian (and early Norian) may triggered a wide
spectrum of morphological disparity that changed the course of the
evolution of Dinosauromorpha during the Mesozoic Era, turning this
clade into the core faunal component of later land Norian ecosystems
(Langer, 2014). Carnian dinosauromorphs (mainly latest Carnian) are
fairly distributed throughout Pangea (Fig. 12C), although still con-
centrated in south Pangea (Brazil and Argentina – Martínez et al., 2013,
Garcia et al. in Press, see above), ranging from small hypercursorial
lagerpetids (e.g. UFSM 11611, Ixalerpeton polesinensis – Cabreira et al.,
2016; Müller et al., 2018a) and herbivorous/omnivorous silesaurids
(e.g. Silesaurus opolensis from Poland – Dzik, 2003, Martínez et al.,
2013, see also Agnolin and Rozadilla, 2017) to large predatory her-
rerasaurids (e.g. UFSM 11330, Herrerasaurus ischigualastensis – Novas,
1994; see above), and small-sized predatory/generalist saur-
opodomorphs and theropods (e.g. Buriolestes schultzi, Eodromaeus
murphi, Panphagia protos, Saturnalia tupiniquim – Martínez and Alcober,
2009; Martínez et al., 2011; Bronzati et al., 2017; Müller et al., 2018b),
filling a wide gamut of “slots” in the terrestrial trophic webs (Cabreira
et al., 2016). While already well-established in South American de-
posits, the records of Carnian dinosauromorphs are also reasonably
distributed in south Pangea (Ezcurra, 2012), with additional (frag-
mentary) specimens found in: lower Maleri Formation of India (Al-
walkeria maleriensis – Chatterjee, 1987, but see Langer, 2004, Ezcurra,
2012), Timezgadiouine Formation of Morocco (Diodorus scytobrachion –
Kammerer et al., 2012), Pebbly Arkose Formation of Zimbabwe (a pu-
tative saurischian fragmentary femur). In this temporal extent, dino-
sauromorphs may also have reached the Laurasian portion of Pangaea
as well, mostly represented by dinosauriforms of uncertain affinities
and with ambiguous chronostratigraphical constraints: Saltopus elgi-
nensis present in the Lossiemouth Sandstone of Scotland (Benton and
M.S. Garcia, et al.
Walker, 2011; Langer et al., 2013; Baron and Williams, 2018), Case-
osaurus crosbyensis from the North American Tecovas Member of the
Dockum Group (Hunt et al., 1998, but see Langer, 2004; Baron and
Williams, 2018), and Chindesaurus bryansmalli (Hunt et al., 1998, but
see Langer, 2004; Nesbitt et al., 2007; Baron and Williams, 2018) and
Eucoelophysis baldwini (a silesaurid dinosauriform – Ezcurra, 2006) both
from the Petrified Forest Member of the Chinle Formation.
Finally, in the Norian, dinosauromorphs became virtually cosmo-
politan, spreading along most of the Pangaea, and predominantly re-
presented by sauropodomorphs (Müller et al., 2016, see above).
Meanwhile, theropods (Nesbitt et al., 2009b; Ezcurra, 2017; Martínez
and Apaldetti, 2017), lagerpetids (Irmis et al., 2007; Nesbitt et al.,
2009a; Martínez et al., 2016; Marsh, 2018) and silesaurids (Ezcurra,
2006, Ferigolo and Langer, 2007) are also present, but to a lesser ex-
tent. The absence of unequivocal herrerasaurids in the Norian epoch
points toward the extinction of this group in the Carnian – Norian
transition (see Baron and Williams, 2018), while ornithischians also do
not have any unambiguous Triassic record (see Agnolin and Rozadilla,
2017). The rise of theropods in detriment of other predatory archosaurs
(Bonaparte, 1982), such as “rauisuchians” was probably influenced by
the faunal turnover occurred across the Carnian – Norian boundary,
with larger hypercarnivores (e.g. herrerasaurids and “rauisuchians”)
struggling with the decline of rhynchosaurs and dicynodonts (i.e. easy-
to-catch prey), followed by their replacement by more agile herbi-
vorous dinosauromorphs (e.g. cursorial silesaurids and subcursorial
sauropodomorphs), which, associated with the modifications in the
Triassic landscape across the Norian (Baranyi et al., 2017), may have
been decisive for the extinction of “rauisuchians” and the prevalence of
theropod dinosaurs in the gateway for the Jurassic Period (Chatterjee
and Majumdar, 1987). This context supports the idea that the major
dinosauromorph clades (Lagerpetidae, Silesauridae and Dinosauria)
shared the same ecosystems for at least 21 Ma (e.g. Irmis et al., 2007;
Langer et al., 2018), starting at least in the initial Late Triassic, as
evidenced by the findings in the Cerro da Alemoa site, mid Carnian of
southern Brazil. This scenario also corroborates ideas in which the three
major dinosauromorph clades were fairly widespread in Pangaea long
before the complete extinction of non-dinosaurian dinosauromorphs at
the end of the Triassic Period along other land tetrapod clades (Langer
et al., 2013), that resulted in the opportunistic domination of the Early
Jurassic land ecosystems by the major groups of non-avian dinosaurs
315
Journal of South American Earth Sciences 91 (2019) 302–319
(Sauropodomorpha, Theropoda and Ornithischia – Benton, 2004,
Langer et al. 2010), which lasted unto the end of the Cretaceous Period,
culminating into their ultimate mass killing ∼66 Ma (Alvarez et al.,
1980; Hildebrand et al., 1991).
7. Conclusions
UFSM 11611 expands the scarce record and distribution of la-
gerpetids to the earliest Carnian of southern Brazil, so far comprised
only by Ixalerpeton polesinensis. Actually, the morphology of this new
specimen differs from the latter, suggesting a hidden diversity of la-
gerpetids in those beds. Moreover, the new specimen associated with a
new precise time-calibration for Brazilian Triassic beds (Langer et al.,
2018) helps to understand the ecological relationships between dino-
saurs and their major relatives joined existence throughout the Triassic
Period. Indeed, the co-occurrence of UFSM 1161, a silesaurid, and
distinct dinosaur clades (i.e. sauropodomorphs, herrerasaurids, and a
putative theropod) brings support that they all shared the same land
ecosystems from the rise of dinosaurs until the demise of non-dinosaur
dinosauromorphs (e.g. Irmis et al., 2007), a time interval of approxi-
mately for 21 Ma (Langer et al., 2018). Such impressive diversity of
dinosauromorphs living in the same time range also suggest that they
were ecologically diverse, with their members playing distinct roles in
these land ecosystems during the rise of dinosaurs.
Acknowledgements
We thank Gabriel Boeira (UFSM) and Gabrielle Santos (UFSM) for
helping in the screening and early preparation of the material, and Max
Langer (Laboratório de Paleontologia de Ribeirão Preto, Universidade
de São Paulo) for useful suggestions on an earlier version of this project.
We also are indebted to an anonymous reviewer, as well as to Federico
Agnolin and Mateusz Talanda for fine corrections and suggestions that
greatly improved the quality of this manuscript. This work was sup-
ported by the Fundação de Amparo à Pesquisa do Estado do Rio Grande
do Sul (FAPERGS) scholarship to MSG and by the Conselho Nacional de
Desenvolvimento Científico e Tecnológico [CNPq; research grant to
SDS, process number 306352/2016-8]. We extend our gratitude to the
Willi Hennig Society, for the free use of TNT software.
M.S. Garcia, et al.
Appendix 1
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