Journal of South American Earth Sciences 104 (2020) 102846

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Journal of South American Earth Sciences

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

Triassic faunal successions of the Paraná Basin, southern Brazil

Cesar L. Schultz a, *, Agustín G. Martinelli b, Marina B. Soares c, Felipe L. Pinheiro d,
Leonardo Kerber e, Bruno L.D. Horn f, Flávio A. Pretto e, Rodrigo T. Müller e, Tomaz P. Melo a
a
PPGGeo-Universidade Federal do Rio Grande do Sul, Av. Bento Gonçalves, 9500, 91540-000, Porto Alegre, RS, Brazil
b
CONICET-Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Av. Ángel Gallardo 470, C1405 DJR, Buenos Aires, Argentina
c
Departamento de Geologia e Paleontologia, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista s/nº, CEP 20940-040, Rio de Janeiro, Brazil
d
Laboratório de Paleobiologia, Universidade Federal do Pampa, Av. Antônio Trilha, 1847, São Gabriel, Brazil
e
Centro de Apoio à Pesquisa Paleontológica da Quarta Colônia (CAPPA), Universidade Federal de Santa Maria, São João do Polêsine, Rio Grande do Sul, Brazil
f
Serviço Geológico do Brasil/CPRM, Superintendência Regional de Porto Alegre, Rua Banco da Província, 105, CEP90840-030, Porto Alegre, Brazil

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

Keywords: The Paraná Basin was filled by a sedimentary package deposited in successive sedimentation episodes related to
Middle-Late Triassic the tectonic events that hit the SW portion of the Gondwana. The Triassic portion of this package, known
Faunal associations worldwide for its continental tetrapod fauna, occurs only in the southern portion of the basin and is represented
South America
by 2 s-order sequences: the Sanga do Cabral Supersequence (SCS - Early Triassic) and the Santa Maria Super­
Fossil record
Biostratigraphy
sequence (SMS - Middle-Late Triassic). The SCS fauna, including temnospondyls, parareptiles (mainly Procolo­
phon), archosauromorphs, putative synapsids, and a number of indeterminate specimens, is traditionally
considered Early Triassic and corresponds to the “Procolophon abundant zone” of the Karoo Basin (the upper
levels of the Lystrosaurus AZ), in the upper Katberg Formation, which is Induan to early Olenekian in age. The
sedimentary environment of the SCS is thought to be a wide alluvial plain, in which small and shallow channels
spread northwards into a vast semiarid environment. By its turn, the Middle-Upper Triassic Santa Maria
Supersequence is divided into four third-order sequences, from base to top: Pinheiros-Chiniquá, Santa Cruz,
Candelária and Mata. Each of these sequences begins with fluvial deposition (low sinuosity rivers) that is overlain
by transgressive shallow lacustrine deposits. The first three of these sequences present a very rich record of fossil
tetrapods, including four successive faunal associations: Dinodontosaurus Assemblage Zone (Ladinian, within the
Pinheiros-Chiniquá Sequence), Santacruzodon AZ (Ladinian/Carnian, in the Santa Cruz Sequence), Hyper­
odapedon AZ (Carnian) and the Riograndia AZ (early Norian), the latter two respectively at the base and top of the
Candelária Sequence. In general, the lower portion of the package (Pinheiros-Chiniquá and Santa Cruz Se­
quences) was deposited under more basic and dried environmental conditions and are dominated by synapsids,
while the top of the section (Candelária Sequence) is characterized by more acid and humid conditions and by a
shift in the faunal content, with diapsids as dominating forms and presenting an increase of diversity compared to
the lower biozones.

1. Introduction south Brazil) and is represented by 2 s-order sequences: the Sanga do

The Paraná Basin (PB) comprises an 8.000 m thick sedimentary
package, deposited in polycyclic events resulting from successive sedi­
mentation episodes related to the tectonic events that hit the SW portion
of the Gondwana (Milani, 1997; Milani et al., 1998). The Triassic beds of
this package, known worldwide for its continental tetrapod fauna, oc­
curs only in the southern portion of the basin (in Rio Grande do Sul State,
Cabral Supersequence (SCS - Early Triassic) and the Santa Maria
Supersequence (SMS - Middle-Late Triassic). In lithostratigraphic terms,
the first corresponds to Sanga do Cabral Formation, and the latter cor­
responds to Santa Maria and Caturrita Formations (Fig. 1). The SCS
fauna is traditionally considered Early Triassic and partially coeval to
the Lystrosaurus Assemblage Zone from the Karoo Basin due to the
presence of the widespread Gondwanan parareptile Procolophon

* Corresponding author.
E-mail addresses: cesar.schultz@ufrgs.br (C.L. Schultz), agustin_martinelli@yahoo.com.ar (A.G. Martinelli), marina.soares@mn.ufrj.br (M.B. Soares),
felipepinheiro@unipampa.edu.br (F.L. Pinheiro).

https://doi.org/10.1016/j.jsames.2020.102846
Received 25 March 2020; Received in revised form 21 July 2020; Accepted 24 August 2020
Available online 28 August 2020
0895-9811/© 2020 Elsevier Ltd. All rights reserved.
C.L. Schultz et al.
trigoniceps (Dias-da-Silva et al., 2006b; Cisneros, 2008). On its turn, the
Middle-Late Triassic Santa Maria Supersequence (SMS) is subdivided
into four third-order sequences, from base to top: Pinheiros-Chiniquá,
Santa Cruz, Candelária, and Mata Sequences (Zerfass et al., 2003; Horn
et al., 2014). Each of these sequences begins with fluvial deposition (low
sinuosity rivers) that is overlain by aeolian and shallow lacustrine de­
posits. This stacking is interpreted as cyclic basin subsidence, induced by
the tectonic uplifting of the source areas (Zerfass et al., 2004). The first
three of these sequences present a very rich record of fossil tetrapods,
divided into four successive faunal associations: the Dinodontosaurus
Assemblage Zone (Ladinian/Carnian, within the Pinheiros-Chiniquá
Sequence), the Santacruzodon AZ (early Carnian, in the Santa Cruz
Sequence), the Hyperodapedon AZ (Carnian) and the Riograndia AZ
(early Norian), respectively at the base and top of the Candelaria
Sequence. Absolute ages obtained for SMS by the U-Pb method in dating
detrital zircons (Langer et al., 2018; Philipp et al., 2018), indicate
minimal ages of 236.1 Ma (Santa Cruz Sequence), 233.23 Ma (Hyper­
odapedon AZ) and 225.42 Ma (Riograndia AZ). Both faunal content and
absolute ages correlate the Pinheiros-Chiniquá and Candelária se­
quences from Brazil, respectively, with the Chañares and Ischigualasto
formations from Argentina (Langer, 2005; Langer et al., 2007b; Martínez
et al., 2013). Here we present an up to date summary of the Triassic
faunal successions in Paraná Basin, correlating the tetrapod record of
each Assemblage Zone to the geological framework of Brazilian Triassic.
2. Institutional abbreviations
CAPPA/UFSM, Centro de Apoio à Pesquisa Paleontológica da Quarta
Colônia, Universidade Federal de Santa Maria, Rio Grande do Sul, Brazil;
Fig. 1. Stratigraphical chart of Sanga do Cabral and Santa Maria Supersequences, and their correspondent in lithostratigraphy, depositional environments, and
biostratigraphy. Lithologies based on Zerfass et al. (2003) and Horn et al. (2018a, b). Biostratigraphy from Soares et al. (2011b). Abbreviations: Ani, Anisian; Ind,
Induan; Ole, Olenekian.
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Journal of South American Earth Sciences 104 (2020) 102846
MCN-FZB-PV, Museu de Ciências Naturais, Fundação Zoobotânica do
Rio Grande do Sul (Coleção de Paleovertebrados), Porto Alegre, Brazil;
MCP-PV, Museu de Ciências e Tecnologia da Pontifícia Universidade
Católica do Rio Grande do Sul (Coleção de Paleovertebrados), Porto
Alegre, Brazil; MMACR-PV-T, Museu Municipal Aristides Carlos
Rodrigues (Coleção de Paleovertebrados-Triássico), Candelária, Brazil;
MCT, Museu de Ciências da Terra, Rio de Janeiro, Brazil; UFRGS-PV-T,
Universidade Federal do Rio Grande do Sul (Coleção de Paleoverte­
brados-Triássico), Porto Alegre, Brazil; UFSM, Universidade Federal de
Santa Maria, Laboratório de Estratigrafia e Paleobiologia (Santa Maria,
Brazil); UMVT, Universidade do Vale do Rio dos Sinos, Museu da Vida e
da Terra, São Leopoldo, Brazil; UNIPAMPA, Universidade Federal do
Pampa, Laboratório de Paleobiologia, São Gabriel, Brazil.
3. Early Triassic
3.1. Sanga do Cabral Supersequence - Procolophon AZ
3.1.1. Geological setting
The Sanga do Cabral Formation (SCF) of South Brazil was originally
proposed by Andreis et al. (1980) to designate very diagnostic succes­
sions of fine-grained sandstones intercalated by abundant intraforma­
tional conglomerates that crop out in an E–W belt spread throughout a
vast territory of the central Rio Grande do Sul state (see Dias-da-Silva
et al., 2017). The Sanga do Cabral Formation has been proposed as
laterally correlating to the Uruguayan Buena Vista Formation (Andreis
et al., 1996), but this later presents a distinct faunal content (Piñeiro
et al., 2003, 2007) and was dated as late Permian in a recent paper
(Ernesto et al., 2020).
C.L. Schultz et al.
Faccini (1989) postulated that the SCF represented a distinct allos­
tratigraphic unit, which he named the Eoscythian Sequence that was
separated from the underlying Permian package by an unconformity.
The underlying Upper Permian units are the fluvio-lacustrine Rio do
Rasto Formation (RRF) and the aeolian Piramboia Formation (PF), both
laterally interbedded. Posteriorly, Zerfass et al. (2003) enhanced the
allostratigraphic framework proposed by Faccini (1989) and the
Eoscythian Sequence was renamed as Sanga do Cabral Supersequence
(SCS), a tectonically controlled, second-order allostratigraphic unit
(Fig. 2), whose dominant facies association comprises intraformational
massive or trough cross-bedded conglomerates, and horizontal bedded
sandstones, sometimes with chute and pool structures and cyclic steps.
These facies were interpreted as low-sinuosity, high energy,
sheetflow-dominated rivers (Zerfass et al., 2003; Dario, 2017). Accord­
ing to Zerfass et al. (2003), the disconformity that delineates the basal
boundary of the SCS was correlated to Veevers et al. (1994) Gondwa­
nides I paroxysm, a tectonic event that would be responsible for the most
striking sedimentary feature of the SCS (with regard to the underlying
Permian rocks), i.e., a shift in the fluvial pattern that changed from
meandering to braided, reflecting an uplift in the relief. This change in
the fluvial style, from high- (Late Permian) to low-sinuosity (Early
Triassic) is also recorded in the Karoo Basin (Katberg Formation), which
has also been linked to the Gondwanides I paroxysm (Smith, 1995) or to
climatic shifts (Ward et al., 2000; Retallack et al., 2003). Classical SCF
exposures are located nearby Dilermando de Aguiar and São Vicente do
Sul municipalities, central Rio Grande do Sul State. More recently,
renewed prospections were able to recognize additional outcrops close
to the western border of the Paraná Basin Triassic belt. Among those, the
‘Bica São Tomé’ site (Da-Rosa et al., 2009) stands apart for producing a
comparatively large number of well-preserved specimens.
3.1.2. Tetrapod faunal content
Despite reasonably common, Procolophon AZ fossils often occur as
disarticulated/fragmented tetrapod remains, being fairly complete
specimens considered as exceptional occurrences (Pinheiro et al., 2016,
2019; De-Oliveira et al., 2020). Therefore, field prospections in SCS
were historically neglected, and its fossil content was often regarded as
monotonous or uninteresting.
Similar to what is common in other Early Triassic successions, SCS
tetrapod fauna (Figs. 2 and 3) is thus far composed of typically small to
medium-sized generalist taxa. Procolophonoid skulls and postcranial
elements are by far the most abundant occurrences, and temnospondyls
are also common in several outcrops. The archosauromorph record
became significant with renewed prospections (e.g., Da-Rosa et al.,
2009; Pinheiro et al., 2016; Oliveira et al., 2018) but, curiously, un­
ambiguous synapsid remains are completely absent (see Dias-da-Silva
et al., 2017).
The first SCS specimens of the parareptile genus Procolophon were
briefly reported by Barberena et al. (1981). Later, Lavina (1983)
described “Procolophon pricei”, which would differ from P. trigoniceps for
having a distinct palatal dentition pattern. An additional Procolophon
species (“P. brasiliensis”) was proposed by Cisneros and Schultz (2002)
for the SCS, also diagnosed by its peculiar palatal dentition. The two
Brazilian taxa were later considered as junior synonyms of a mono­
specific P. trigoniceps by Cisneros (2008). Further SCS P. trigoniceps
specimens were reported by Dias-da-Silva et al. (2006b, 2017), Da-Rosa
et al. (2009) and Silva-Neves et al. (2018). Notably, P. trigoniceps is the
best biostratigraphically constrained taxon so far reported for the Pro­
colophon AZ. In the South African Karoo, P. trigoniceps becomes abun­
dant in the upper Katberg Formation (Lystrosaurus Assemblage Zone,
Induan/Olenekian), in strata conserably higher than the Permo-Triassic
boundary (Botha and Smith, 2006).
Morphological disparity among recovered SCS procolophonoid
vertebrae and skull fragments suggests the presence of additional, non-
Procolophon taxa. Notably, a Kitchingnathus-like morphotype is recog­
nizable from an almost complete skull (Pinheiro et al. in prep.). So far,
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Journal of South American Earth Sciences 104 (2020) 102846
all procolophonoid specimens recovered from SCS (even unpublished
specimens) are consistent with an Induan/Olenekian age for the
formation.
Procolophon AZ temnospondyls are represented by several frag­
mented remains associated to indeterminate Stereospondyli, Rhytidos­
teidae, Mastodonsauroidea, and Plagiosteninae, besides two nominal
taxa known from partial skulls (e.g., Dias-da-Silva et al., 2005; Dia­
s-da-Silva and Schultz, 2008; Da-Rosa et al., 2009). The rhytidosteid
Sangaia lavinai (Dias-da-Silva et al., 2006a) was named based on a
partial left skull roof and palatal bones, whereas Tomeia witecki Eltink
et al., 2016 (Capitosauria) is originally known from a partial posterior
portion of the skull (including the occiput), in addition to scattered
dermal skull bones and a lower jaw fragment (Eltink et al., 2016). Recent
fieldwork in T. witecki type locality (10 years after the discovery of the
original remains) yielded a posterior skull fragment that is perfectly
complementary to the holotype, certainly corresponding to the same
specimen (Elesbão et al. in prep.). Sangaia lavinai has only limited use for
biostratigraphic correlations, as rhytidosteids are known from a broad
timespan that includes the Permo-Triassic boundary (see Dias-da-Silva
et al. (2017) for further discussion). Capitosauroids such as T. witecki,
however, seem to have their earliest occurrence in the Early Triassic
(Eltink et al., 2016; Dias-da-Silva et al., 2017), supporting an Indua­
n/Olenekian age for the Procolophon AZ.
Procolophon AZ archosauromorph record is mainly composed of
isolate vertebrae and other postcranial elements (e.g., Langer and Lav­
ina, 2000; Da-Rosa et al., 2009; Oliveira et al., 2018), as well as
exceptionally rare, fairly complete specimens (Pinheiro et al., 2016,
2019; De-Oliveira et al., 2020). Peculiar elongated cervical vertebrae
were identified as belonging to “Protorosauria” by Langer and Lavina
(2000). These specimens and other similar material were further
investigated by Oliveira et al. (2018), who recovered them as closely
related to the Tanystropheidae. In addition, the recently described
archosauromorph Elessaurus gondwanoccidens is worth noting for being
represented by a partial skeleton, composed by an almost complete
hindlimb, pelvic girdle, sacral and caudal vertebrae. The taxon was also
recovered as closely related to tanystropheids (De-Oliveira et al., 2020).
The Tanystropheidae is a peculiar clade of early-diverging arch­
osauromorphs with a mostly Tethyan distribution in the Middle to Late
Triassic. The clade is so far unknown in Permian strata, but Early
Triassic representatives (aside from the SCS specimens) are known from
Russia (Sennikov, 2011; Oliveira et al., 2018).
The most complete tetrapod thus far recovered from the Procolophon
AZ is the exceptionally preserved skull/cervical vertebrae of Teyujagua
paradoxa (Pinheiro et al., 2016). Although not particularly useful for
biostratigraphic correlations, T. paradoxa is of broad evolutionary in­
terest, as it shows character transitions that would ultimately lead to the
origin of the Archosauriformes, the ruling tetrapod clade in Mesozoic
terrestrial ecosystems (Pinheiro et al., 2016, 2019). Few years after the
discovery of T. paradoxa holotype, further fieldwork in the type locality
produced a fairly complete postcranium that, most likely, belongs to the
same individual.
Some other archosauromorph specimens were illustrated and briefly
described by Da-Rosa et al. (2009). These and other yet to be published
vertebrae and long bones display close resemblance to those of proter­
osuchid archosauriforms (FLP, personal observation).
As was previously mentioned, the SCS is peculiar for the (so far)
apparent absence of unambiguous synapsids in its succession. Putative
cynodont and Lystrosaurus remains were reported by Langer and Lavina
(2000), Abdala et al. (2002a), Da-Rosa et al. (2009) and Dias-da-Silva
and Da-Rosa (2011). Recent revaluation of these specimens, however,
showed that they are indistinguishable from procolophonoids, the most
common taxon in Procolophon AZ (Dias-da-Silva et al., 2017). Reasons
underlying the absence of synapsids in this zone still need investigation,
but taphonomic biases and limited collection efforts are not unlikely.
C.L. Schultz et al. Journal of South American Earth Sciences 104 (2020) 102846

Fig. 2. Chrono- and biostratigraphy of Triassic units with vertebrate Assemblages Zones from southern Brazil, and main localities (A) and the tetrapod fossil content
(B) of the Procolophon AZ. Absolute ages (indicated by arrows) as in Fig. 1. The ages of the column follow Gradstein et al., 2012. Abbreviations: Arch, sister taxon
Archosauriformes; Ca, Capitosauria; Ind. Induan; Ole, Olenekian; Rhyt, Rhytidosteidae; Ta, Tanystropheidae.

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C.L. Schultz et al. Journal of South American Earth Sciences 104 (2020) 102846

Fig. 3. Selected fossils from the Procolophon AZ. A, Procolophon trigoniceps (UFRGS-PV-231-T), skull in palatal view; B, Sangaia lavinai holotype (UMVT 4302), partial
skull in dorsal view; C, Teyujagua paradoxa holotype (UNIPAMPA 0653), skull in right lateral view; D, possible proterosuchid cervical vertebra in left lateral view; E,
Elessaurus gondwanoccidens holotype (UFSM 11471), articulated hind limb and axial elements in left lateral view; F, Tomeia witecki holotype (UFSM 11408), posterior
skull in palatal view; G, cervical vertebra with tanystropheid affinities (UFRGS-PV-492-T) in right lateral view.

4. Middle-Late Triassic
4.1. Pinheiros-Chiniquá Sequence - Dinodontosaurus AZ
4.1.1. Geological setting
The Pinheiros-Chiniquá Sequence solely includes the Dinodontosau­
rus AZ (Horn et al., 2014), which represents the oldest faunal association
of the Santa Maria Supersequence. Outcrops containing the vertebrate
fauna of the Dinodontosaurus AZ are numerous (Fig. 3), distributed all
along the Triassic belt of Rio Grande do Sul (Barberena et al., 1985;
Schultz, 2005; Langer et al., 2007b; Martinelli and Soares, 2016; Mar­
tinelli et al., 2017a). As stated before, all sequences of Santa Maria
Supersequence begin with low-sinuosity, ephemeral braided rivers
characterized by greyish conglomeratic sandstones with planar and
trough-cross stratification (Zerfass et al., 2003; Horn et al., 2018). These
rocks are overlaid by metric layers of massive red mudstones with
pedogenetic alterations, as calcretes and slickensides (Da Rosa et al.,
2005; Horn et al., 2013). These rocks are interpreted as large loessic
plains with sporadic fluvial reworking (Horn et al., 2018).
The name Dinodontosaurus AZ was proposed by Barberena et al.
(1985), based on the formerly known Therapsida Cenozone (Barberena,
1977), restricted to the Alemoa Member of the Santa Maria Formation
and placed below the Rhynchosauria (=Scaphonyx) Cenozone (now
known as the Hyperodapedon AZ). Moreover, the Therapsida Cenozone
was established on the vertebrate components included in the Chiniquá
and Pinheiros Local Faunas (Barberena, 1977; Barberena et al., 1985;
Schultz et al., 2000), dominated by the dicynodont Dinodontosaurus and
the cynodonts Massetognathus and Chiniquodon. The Chiniquá Local
Fauna includes a series of outcrops (traditionally called Sangas; see the
meaning of this word in Beltrão, 1965, Langer et al., 2007b, and Mar­
tinelli et al., 2016) located in the Chiniquá region, at the municipality of
São Pedro do Sul. These outcrops have been intensely worked since the
beginning of the 20th century with the initial publications of Friedrich
von Huene (1935, 1936, 1942; Zingano and Cauduro, 1959; Beltrão,
1965). The most fossiliferous ones are the Sanga Cynodontier (also
known as Cynodont Sanga, Sanga dos Cinodontes, Sanga Béles, Sanga de
Theotônio Béles Xavier, Sanga north to the house of Theotônio Béles
Xavier), Sanga Baum (also known as Sanga da Árvore) and Sanga Weg
(also known as Sanga do Caminho or Sanga da Estrada) (Huene, 1935,
1936; 1942; Beltrão, 1965; Martinelli et al., 2017a). Additionally, there
are outcrops in this region that were cited in the literature as Rote Sanga
(=Sanga Vermelha), Sanga Leere (=Sanga Vazia), Sanga Cesar (=Sanga

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C.L. Schultz et al. Journal of South American Earth Sciences 104 (2020) 102846

César), and Sanga Sud-Weg (=Sanga south to Sanga Weg, =Sanga ao sul
da Sanga Weg) with less abundant fossil records (Martinelli et al., 2017a:
Supl. Info.).
Conversely, the Pinheiros Local Fauna was established to include the
fauna discovered in a series of outcrops located south of Candelária, in
the Pinheiro (erroneously used as Pinheiros or Vila Pinheiros) and Bom
Retiro regions, municipality of Candelária (Barberena, 1977; Barberena
et al., 1985; Schultz et al., 2000), in addition to other outcrops located in
the municipalities of Bom Retiro do Sul (e.g., Venâncio Aires), Vale
Verde (e.g., Vale Verde), Rio Pardo and Cachoeira do Sul. The most
fossiliferous outcrops in the Pinheiro region are: Sanga Pinheiro [also
known as Sanga Pinheiros, Sanga dos Fósseis (Fossil’s Sanga), Sanga
Carlos, and possibly Sanga do Forno (“Furnace’s Sanga”)], Sanga Bel­
miro (=Sanga Lili), Sanga Nicanor (=Sanga do Zé, Rincão do Simeão),

Fig. 4. Chrono- and biostratigraphy of Triassic units with vertebrate Assemblage Zones from southern Brazil, and main localities (A) and the tetrapod fossil content
(B) of the Dinodontosaurus AZ. Absolute ages (indicated by arrows) as in Fig. 1. The ages of the column follow Gradstein et al., 2012. Abbreviations: Ap, Aphano­
sauria; E, Erpetosuchidae; Ind. Induan; Ole, Olenekian; Ow, Owenettidae; P, Probainognathia; Ra, Rauisuchia; Rh, Rhynchosauria; T, Traversodontidae.

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C.L. Schultz et al. Journal of South American Earth Sciences 104 (2020) 102846

Sanga da Divisa, Sanga Janguta, Sanga do Ribeiro, among others (Bar­
berena, 1977; Barberena et al., 1985; Martinelli et al., 2016, 2017a).
With regard to Bom Retiro region, the most fossiliferous outcrops are:
Sanga Hintz, Sanga Pascual, Sanga Menezes, and Cerro dos Gomes
(Barberena, 1977, 1978; Martinelli et al., 2017a).
The location of several of these traditional outcrops representing the
Chiniquá and Pinheiros Local Faunas is likely imprecise due to
agricultural-related changes in the topography of those regions, since
the first discoveries (Beltrão, 1965; Dassie, 2014; Lacerda et al., 2016;
Martinelli et al., 2016, 2017a). Moreover, some outcrops correspond to
non-formal topographic names and thus, their location is even more
difficult to find out (Martinelli et al., 2016, 2017a).
Over the years, new outcrops containing faunal components of the
Dinodontosaurus AZ have been discovered, filling in the geographical
distribution of this AZ and enlarging their taxonomic context. Most
relevant ones are the outcrops known as Cortado (e.g., Da-Rosa et al.,
2004; Reichel et al., 2009), Linha Várzea (e.g., Da-Rosa et al., 2005),
Dona Francisca (Posto outcrop), and Bortolin sites (e.g., Bonaparte et al.,
2006; França et al., 2011; Pavanatto et al., 2020).

Fig. 5. Selected fossils from the Dinodontosaurus AZ. A, skull of Dinodontosaurus sp. in right lateral view; B, Skull and jaws of Prestosuchus sp. (UFRGS-PV-156-T) in
left lateral view; C, skull and jaws of Candelaria barbouri (UFSM 11076) in left lateral view; D, skull and jaws of Barberenasuchus brasiliensis (MCP-PV 220) in right
lateral view; E, skull and jaws of Protheriodon estudianti (UFRGS-PV-0962-T) in right lateral view; F, skull of Aleodon cromptoni (UFRGS-PV-146-T) in left lateral view;
G, skull and jaws of Chiniquodon theotonicus (UFRGS-PV-1331-T) in right lateral view; H, partial skull of Scalenodon ribeiroae (UFRGS-PV-0239-T) in ventral view; I,
skull of Massetognathus ochagaviae (MCP-PV 3871) in ventral view; J, skull and jaws of Bonacynodon schultzi (MCT-1716-R) in left lateroventral view.

7
C.L. Schultz et al.
4.1.2. Tetrapod faunal content
The fossiliferous composition of the Dinodontosaurus AZ (Figs. 4 and
5) includes procolophonians, diverse non-dinosaur archosauromorphs,
and dicynodont and non-mammaliaform cynodont therapsids. The
procolophonians are represented by the owenettid Candelaria barbouri,
discovered in two localities: Cortado and Pinheiro (erroneously written
in the literature as Pinheiros) regions (Price, 1947; Cisneros et al., 2004;
Da-Rosa et al., 2004).
The archosauromorphs include a rhynchosaur, a proterochampsid,
erpetosuchids, “rauisuchians”, and possibly avemetatarsalians. The
rhynchosaur is represented by the stenaulorhynchine Brasinorhynchus
mariantensis (formerly known as the “Mariante rhynchosaur”), based on
two specimens that were unearthed in only one locality: Porto Mariante
1, municipality of Bom Retiro do Sul (Schultz et al., 2016).
Proterochampsids are based on specimens collected at Pinheiro re­
gion several decades ago. They were originally related to Chanaresuchus
and Gualosuchus (Dornelles, 1995), but their taxonomy is uncertain, and
they are currently under revision. Archeopelta arborensis and Pagosve­
nator candelariensis are the two hitherto known erpetosuchids of the
Dinodontosaurus AZ (Desojo et al., 2011; Ezcurra et al., 2017; Lacerda
et al., 2018). The first species was discovered in the Chiniquá region,
whereas the second one comes from the municipality of Candelária or a
nearby region (see Lacerda et al., 2018).
The large-sized “rauisuchian” predators are well known in the
Dinodontosaurus AZ, represented by Prestosuchus chiniquensis and
Decuriasuchus quartacolonia. The first one is represented by several well-
preserved specimens, including almost complete skeletons, recovered in
several outcrops, such as in Chiniquá, Pinheiro, Dona Francisca and Vale
Verde (e.g., Barberena, 1978; Lacerda et al., 2016; Roberto-Da-Silva
et al., 2018; Mastrantonio et al., 2019). The second species is based on
several individuals recovered in association in the Posto outcrop at Dona
Francisca municipality (França et al., 2011). A recent revision of the
Brazilian specimens referred to Prestosuchus chiniquensis suggested more
than one species for this genus (see Desojo et al., 2020a).
Barberenasuchus brasiliensis and Spondylosoma absconditum are two
problematic taxa based on poorly preserved material. The only known
specimen of Barberenasuchus consists of skull with the first vertebra
discovered at Cortado outcrop. It was originally related to spheno­
suchians (Mattar, 1987) and, later on, to “rauisuchians” (Kischlat,
2000), indeterminate archosauriform (Irmis et al., 2013) or dinosaur­
omorphs (França et al., 2013). Spondylosoma was discovered in the
Chiniquá region and was first considered as a saurischian dinosaur
(Huene, 1942), a rauisuchian (Galton, 2000), but posteriorly interpreted
as an indeterminate basal dinosauriform (Langer, 2004) or more
recently as an aphanosaurian (a group of basal avemetatarsalians;
Nesbitt et al., 2017).
Therapsids are the most abundant component of the Dinodontosaurus
AZ, including two dicynodonts and several taxa of basal traversodontid
and probainognathian cynodonts. The dicynodonts are represented by
the toothed shansiodontid Dinodontosaurus pedroanum (for discussion
about the species-level taxonomy of this genus see Dassie, 2014), rep­
resented by several skeletons discovered in almost all localities ascribed
to the Dinodontosaurus AZ. Despite its high abundance, the complex
history of the genus and species (e.g., Cox, 1965; Lucas and Harris,
1996) claims for a profound restudy of the Dinodontosaurus specimens
and taxonomy. The other dicynodont taxon is represented by the
toothless stahleckeriid Stahleckeria potens, recovered in the Chiniquá
region (Huene, 1935) and in the municipality of Candelária (Pinheiro
and Bom Retiro areas; Romer and Price, 1944; Lucas, 1993; Peruzzo and
Araújo-Barberena, 1995; Vega-Dias et al., 2005).
Traversodontids are herbivorous/omnivorous gomphodont cyn­
odonts that are, in the Dinodontosaurus AZ, represented by six species:
Scalenodon ribeiroae, Massetognathus ochagaviae, M. pascuali, Luangwa
sudamericana, Traversodon stahleckeri, and Protuberum cabralense
(Huene, 1936; Barberena, 1974, 1981a, b; Sá-Teixeira, 1987; Abdala
and Sá-Teixeira, 2004; Liu et al., 2008; Reichel et al., 2009; Liu and
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Journal of South American Earth Sciences 104 (2020) 102846
Abdala, 2014; Pavanatto et al., 2016; Martinelli et al., 2017a; Melo et al.,
2017; Pavanatto et al., 2020). Scalenodon ribeiroae is based on a single
specimen found in Agudo (Melo et al., 2017), but another one from
Candelária is under study. Massetognathus ochagaviae was recovered in
most sites of this AZ, but apparently not in the Chiniquá region (Liu
et al., 2008; Martinelli et al., 2017a). Moreover, one record of
M. ochagaviae is known in the younger Santacruzodon AZ (Schmitt et al.,
2019). Only one putative Brazilian specimen was referred to M. pascuali
(a species firstly reported for Argentina), which was collected in Can­
delária (Sá-Teixeira, 1995; Liu et al., 2008). The holotype of Luangwa
sudamericana has an unknown provenance, but other specimens were
mentioned for Dona Francisca, Bortolin, Candelária and Vale Verde
(Hanich et al., 2013; Martinelli et al., 2017a; Pavanatto et al., 2020).
Traversodon stahleckeri is restricted until now to the Chiniquá region
(Huene, 1936; Barberena, 1981a; Liu and Abdala, 2014) and Protuberum
cabralense is based on material collected at Rincão do Pinhal and Cor­
tado outcrops (Reichel et al., 2009).
The non-mammaliaform probainognathians also include six species
but they are less abundant than traversodontids. They are the chini­
quodontids Chiniquodon theotonicus and Aleodon cromptoni (Huene,
1936; Teixeira, 1982; Abdala and Giannini, 2002; Martinelli et al.,
2017a), the probainognathid Bonacynodon schultzi (Martinelli et al.,
2016), and Protheriodon estudianti and Candelariodon barberenai (Bona­
parte et al., 2006; Oliveira et al., 2011b; Martinelli et al., 2016, 2017b),
which seem to be more closely related to mammaliaforms than the
previous mentioned taxa. Chiniquodon and Aleodon have a broad dis­
tribution along the Rio Grande do Sul outcrops (Martinelli et al., 2017a),
whereas the record of Bonacynodon is restricted the Pinheiro region
(Martinelli et al., 2016), Protheriodon to Dona Francisca (Bonaparte
et al., 2006), and Candelariodon to the Pinheiro and Cortado regions
(Oliveira et al., 2011b; Martinelli et al., 2017a, b).
Absolute dates for the Pinheiros-Chiniquá Sequence are still not
available, and its chronostratigraphic placement is mainly based on the
comparison of its tetrapod fauna with that of other units (e.g., Bar­
berena, 1977; Barberena et al., 1985; Schultz, 1995, 2005; Schultz et al.,
2000, 2016; Langer et al., 2007b, 2018; Abdala et al., 2009; Martinelli
et al., 2016, 2017a) and the recently obtained age for the overlying
Santa Cruz Sequence (Philipp et al., 2018; see also Schmitt et al., 2019).
The Dinodontosaurus AZ was traditionally correlated to the Chañares
Formation due to the shared occurrence of the genera Dinodontosaurus,
Massetognathus, the species Chiniquodon theotonicus (since Probelesodon
from Argentina was synonymized with Chiniquodon; see Abdala and
Giannini, 2002), and apparently one record of the Argentinean species
M. pascuali in Brazil (Liu et al., 2008).
Other faunal components are quite similar, but to a higher taxonomic
rank, such as the shared presence of large-sized rauisuchians (Luper­
osuchus and Prestosuchus, Nesbitt and Desojo, 2017), stenaulo­
rhynchines, erpetosuchids, proterochampsids and probainognathids
(Probainognathus and Bonacynodon). However, this scenario became
more complex due to new radiometric dates for the Chañares Formation
(Marsicano et al., 2016; Ezcurra et al., 2017) and the Santa Cruz
Sequence (Philipp et al., 2018), and the discovery of new faunal ele­
ments from the Chañares Formation and Pinheiros-Chiniquá and Santa
Cruz sequences (e.g., Martinelli et al., 2017a; Ezcurra et al., 2017; Melo
et al., 2017; Schmitt et al., 2019). One of the major implications was the
recognition of two AZs in the lower member of the Chañares Formation:
the Tarjadia and Massetognathus-Chanaresuchus AZs (Ezcurra et al.,
2017). The genera Massetognathus and Chiniquodon occur in the upper
AZ, whereas cf. Aleodon and aff. Scalenodon in the Tarjadia AZ. As
highlighted by Martinelli et al. (2017a; see also Abdala and Sá-Teixeira,
2004; Langer et al., 2007b), although the Dinodontosaurus AZ may have
hidden two sub-AZs, their recognition is almost impossibly because most
historical specimens have no precise provenance (possibly including
mixed taxa), the lateral correlation among outcrops is difficult to
establish, and the precise taxonomy of several taxa deserves more
studies. Also, the obtained age for the Santa Cruz Sequence (236.6 ± 1.5
C.L. Schultz et al. Journal of South American Earth Sciences 104 (2020) 102846

Ma - earliest Carnian; see Phillip et al., 2018) is similar to that obtained
for the base of the Chañares Formation (236.1 ± 0.6 Ma, Marsicano
et al., 2016; 236 ± 0.3 Ma for the basal portion, see also Ezcurra et al.,
2017), but the faunal composition is more similar when Dinodontosaurus
AZ is compared to both AZs of Chañares Formation (see discussions in
Martinelli et al., 2017a; Melo et al., 2017; Schmitt et al., 2019). How­
ever, a younger age for the Santacruzodon AZ is expected and congruent
with the high dating error (~1.5 Ma) of Santa Cruz Secuence.
Outside South America, the Dinodontosaurus AZ has a strong
biostratigraphic similarity with continental Africa: Namibia, Tanzania,
and Zambia. The presence of the cynodonts Aleodon and Scalenodon, the
rauisuchian Prestosuchus plus erpetosuchids and stenaulorhynchines in
the Dinodontosaurus AZ (e.g., Schultz et al., 2016; Martinelli et al.,
2017a; Melo et al., 2017; Lacerda et al., 2018; Mastrantonio et al., 2019;
Desojo et al., 2020a) allows a clear biostratigraphic correlation with the
Lifua Member of the Manda Beds in Tanzania, which bears the species
Aleodon brachyrhamphus and Scalenodon angustifrons (Crompton, 1955;
Peecook et al., 2017) and Prestosuchus nyassicus (Desojo et al., 2020a), as

Fig. 6. Chrono- and biostratigraphy of Triassic units with vertebrate Assemblages Zones from southern Brazil, and main localities (A) and the tetrapod fossil content
(B) of the Santacruzodon AZ. Absolute ages (indicated by arrows) as in Fig. 1. The ages of the column follow Gradstein et al., 2012. Abbreviations: Ind. Induan; Ole,
Olenekian; P, Probainognathia; Ra, Rauisuchia; T, Traversodontidae.

9
C.L. Schultz et al. Journal of South American Earth Sciences 104 (2020) 102846

well as distinctive genera and species of erpetosuchids and stenaulo­
rhynchines (e.g., Huene, 1938a; Nesbitt and Butler, 2012). Also, the
Brazilian dicynodont Stahleckeria potens, and the cynodonts Aleodon
cromptoni and Luangwa highlight a correlation with the upper Omin­
gonde Formation of Namibia, in which the same species of dicynodont
was recovered (Abdala et al., 2013), one specimen was tentatively
referred to Aleodon cromptoni (Martinelli et al., 2017a), and there is
record of Luangwa sp. (Abdala and Smith, 2009). The genus Luangwa
that was firstly recovered in the Ntawere Formation in Zambia (Kemp,
1980) is also reported in the Dinodontosaurus AZ (Abdala and

Fig. 7. Selected fossils from the Santacruzodon AZ. A, Massetognathus ochagaviae (UFRGS-PV-0712-T), skull in palatal view; B, Santacruzodon hopsoni holotype (MCN-
PV 2768), partial skull in palatal view; C, Massetognathus ochagaviae (UFRGS-PV-0712-T), lower jaw in dorsal view; D, Santacruzgnathus abdalai (UFRGS-PV-1121-T),
partial right dentary in medial view; E, Santacruzodon hopsoni (MCN-PV 2752), lower jaw in left lateral view; F, Menadon besairiei (UFRGS-PV-1164-T), skull in right
lateral view; G, Dagasuchus santacruzensis holotype (UFRGS-PV-1244-T and UFRGS-PV-1245-T), left ilium and ischium in lateral view; H, radinosuchine proter­
ochampsid (UFRGS-PV-0877-T), from left to right and clockwise: left femur in posteromedial view, partial left side of the skull roof in dorsal view, and partial right
dentary in lateral view.

10
C.L. Schultz et al.
Sá-Teixeira, 2004; Martinelli et al., 2017a; Pavanatto et al., 2020). The
Brazilian genus Chiniquodon is also known in Namibia and Madagascar
(Abdala and Smith, 2009; Kammerer et al., 2010), but due to its large
biochron (e.g., Martínez and Forster, 1996; Abdala and Giannini, 2002;
Kammerer et al., 2010) and some problems related to its taxonomy
(Martinelli et al., 2017a), this taxon has less biostratigraphic value than
the others.
4.2. Santa Cruz Sequence – Santacruzodon AZ
4.2.1. Geological setting
The Santacruzodon AZ, in the Santa Cruz Sequence, was discovered
more recently and is more geographically restricted (Fig. 6) than the
other faunal associations of the Santa Maria Supersequence (Abdala
et al., 2001; Abdala and Ribeiro, 2003; Horn et al., 2014; Soares et al.,
2011b). Accordingly, it also has the lowest species richness, with most
specimens belonging to traversodontid cynodonts (e.g., Schmitt et al.,
2019). It was initially named Traversodontid Biozone, based on the
abundance of these cynodonts in the type locality, the Schöenstatt
outcrop, in Santa Cruz do Sul (Abdala et al., 2001). Other fossiliferous
sites with similar faunal content, in Santa Cruz do Sul and in the nearby
cities of Venâncio Aires (Vila Estância Nova) and Vera Cruz (Carolina
Soil), were later assigned to this AZ (Horn et al., 2014; Reichel et al.,
2005), although the Schöenstatt outcrop remains the most taxonomi­
cally diverse locality (Abdala and Ribeiro, 2003; Lacerda et al., 2015;
Martinelli et al., 2016; Melo et al., 2015; Raugust et al., 2013).
This sequence is tectonically restricted by two lineaments, Rio Par­
dinho and Vigia-Roque Lineaments (Horn et al., 2014), comprising the
municipalities of Venâncio Aires, Santa Cruz do Sul and Vera Cruz.
Philipp et al. (2018) provided a maximum depositional age of 236.6 ±
1.5 Ma (earliest Carnian) for the Santa Cruz Sequence based on
LA-MC-ICPMS U-Pb detrital zircon data. The sequence presents the same
depositional sequence as the other sequences, initially with ephemeral
rivers, overlaid by loess deposits.
The depositional environment is very much the same as that of
Pinheiros-Chiniquá Sequence, consisting of basal white to reddish
sandstones and massive red mudstones with common calcretes and
paleosols on the top (Horn et al., 2018b).
4.2.2. Tetrapod faunal content
The faunal composition of the Santacruzodon AZ (Fig, 6, 7) includes
non-dinosaurian archosauriforms, non-mammaliaform cynodonts, and
dicynodont therapsids. Except for the cynodonts, most other species are
represented by punctual occurrences of fragmentary material from the
Schöenstatt outcrop (Abdala et al., 2001; Bertoni-Machado and Holz,
2006; Lacerda et al., 2015).
Non-dinosaurian archosauriforms are represented by the ‘raui­
suchian’ pseudosuchian Dagasuchus santacruzensis, known from a partial
pelvic girdle (Lacerda et al., 2015), and by a proterochampsid arch­
osauriform recognized as Chanaresuchus bonapartei by Raugust et al.
(2013), but considered an indeterminate radinosuchine by Ezcurra et al.
(2015).
Carnivorous/insectivorous probainognathian cynodonts are found in
the Schöenstatt outcrop in the form of the chiniquodontid Chiniquodon
sp. (awaiting detailed description) and the possible prozostrodontian
Santacruzgnathus abdalai (Abdala et al., 2001; Martinelli et al., 2016),
previously referred to as a juvenile cf. Probainognathus (Soares et al.,
2011a).
The traversodontid cynodonts Santacruzodon hopsoni, a massetog­
nathine, and Menadon besairiei, a gomphodontosuchine, are the most
abundant taxa (Abdala and Ribeiro, 2003; Melo et al., 2015). The former
occurs in all localities of the Santacruzodon AZ (Horn et al., 2014), while
the latter is restricted to the Schöenstatt outcrop (Melo et al., 2019,
2015). The recent work by Schmitt et al. (2019) confirmed the previous
identification of a skull and associated mandibles from the Schöenstatt
outcrop as Massetognathus ochagaviae.
11
Journal of South American Earth Sciences 104 (2020) 102846
The only other known vertebrate taxon that has been attributed to
the Santacruzodon AZ is the kannemeyeriiform dicynodont Dinodonto­
saurus sp., based on very fragmentary cranial material from the Vila
Estância Nova outcrop (e.g., Raugust et al., 2013; Martinelli et al.,
2017a; Schmitt et al., 2019). However, this specimen was collected from
a lower stratigraphic level, separated by an angular uncorformity from
the traversodontid-bearing (Santacruzodon) level, and possibly repre­
sents an older fauna than the Santacruzodon AZ (Horn et al., 2014). Also,
it lacks diagnostic characters, being more correctly identified as an
indeterminate tusked dicynodont.
The Santacruzodon AZ has been correlated to the basal Isalo II levels
(Besairie, 1972), also known as Makay Formation (Razafimbelo, 1987),
of the Morondava Basin, in Madagascar, by the shared presence of the
traversodontid Menadon besairiei and the probainognathian genus Chi­
niquodon, and by the close relation of the massetognathine tra­
versodontids Santacruzodon hopsoni and Dadadon isaloi, from Brazil and
Madagascar, respectively (Abdala et al., 2001; Abdala and Ribeiro,
2003; Flynn et al., 2000; Kammerer et al., 2010; Melo et al., 2015;
Nesbitt et al., 2015). The basal Isalo II and Santacruzodon AZ were
originally considered to be late Ladinian-early Carnian based on the
combination of tetrapod taxa found in “Ladinian” (Chañares Formation
and Dinodontosaurus AZ) and “Carnian” (Ischigualasto Formation and
Hyperodapedon AZ) faunas (Abdala et al., 2001; Flynn et al., 2000,
1999). Although the Chañares/Dinodontosaurus and the Ischigualasto/­
Hyperodapedon faunas were both shown to be mostly Carnian (Ezcurra
et al., 2017; Langer et al., 2018; Marsicano et al., 2016; Martínez et al.,
2011, 2013), the intermediate age of the Santacruzodon AZ and basal
Isalo II continues to be acknowledged (e.g., Kammerer et al., 2010; Melo
et al., 2015; Nesbitt et al., 2015). Chiniquodon is also recorded in the
Chañares Formation of Argentina (Abdala and Giannini, 2002), Omin­
gonde Formation of Namibia (Abdala and Smith, 2009), and in the
Dinodontosaurus AZ of the Santa Maria Supersequence (Abdala and
Giannini, 2002; Martinelli et al., 2017a; Huene, 1936), which are
probably older than the Santacruzodon AZ, and in the younger Ischi­
gualasto Formation of Argentina (Bonaparte, 1966; Martínez and For­
ster, 1996), suggesting the genus has a long range. The presence of
Massetognathus ochagaviae also indicates a longer range than formerly
known, as the genus is a key element in Triassic South American
biostratigraphy, previously found exclusively (and abundantly) in the
Chañares Formation (Romer, 1967; Ezcurra et al., 2017) and in the
Dinodontosaurus AZ (Abdala and Giannini, 2000; Liu et al., 2008).
The maximum depositional age provided by Philipp et al. (2018),
236.6 ± 1.5 Ma, overlaps with the ages obtained for the lower member
of the Chañares Formation (236.1 ± 0.6 Ma, Marsicano et al., 2016; 236
± 0.3 Ma for the basal portion; Ezcurra et al., 2017). The faunal and
isotopic age similarities do not exclude the possibility that the Santa­
cruzodon AZ is coeval with portions of the Chañares Formation and even
the Dinodontosaurus AZ, although new data (sedimentary, radiometric or
taxonomic) would be necessary to better constrain the age of deposition.
4.3. Candelária Sequence (base) – Hyperodapedon AZ
4.3.1. Geological setting
The Candelária Sequence, unlike the other sequences, includes the
Hyperodapedon and Riograndia Assemblage Zones (Fig. 8). On the base of
the sequence occur white to reddish sandstones with planar and trough-
cross stratification, sometimes with centimetric mudstone intraclasts
with plant fossils, interpreted as the ever-present ephemeral braided
rivers (Zerfass et al., 2003). These deposits are overlaid by laminated
reddish mudstones, very fine-grained, massive sandstones or stratified
with climbing and wave ripples. The loessic plains of the previous se­
quences is substituted by sheet deltas and ephemeral lakes, indicating a
humidity increase, in comparison with the other two sequences (Horn
et al., 2018b). To the top of the sequence, fine massive sandstone layers
begin to appear interlayered with the laminated mudstone deposits,
characterizing the transition from bottom to top. Locally, the transition
C.L. Schultz et al. Journal of South American Earth Sciences 104 (2020) 102846

Fig. 8. Chrono- and biostratigraphy of Triassic units with vertebrate Assemblages Zones from southern Brazil, and main localities (A) and the tetrapod fossil content
(B) of the Hyperodapedon AZ. The biostratigraghic location of Siriusgnathus is tentative in this AZ. Absolute ages (indicated by arrows) as in Fig. 1. The ages of the
column follow Gradstein et al., 2012. Abbreviations: A, Aetosauria; D, Dinosauria; Ind. Induan; L, Lagerpetidae; Ole, Olenekian; Orn, Ornithosuchidae; P, Probai­
nognathia; Pr, Protherochampsia; Ra, Rauisuchia; Rh, Rhynchosauria; T, Traversodontidae.

12
C.L. Schultz et al. Journal of South American Earth Sciences 104 (2020) 102846

is marked by channel excavations filled with fine-grained sandstones
(Horn et al., 2018a).
The Hyperodapedon AZ is distributed along the central portion of the
Rio Grande do Sul State and is well sampled in the municipalities of Vale
do Sol, Candelária, Agudo, São João do Polêsine, Santa Maria, São Pedro
do Sul, and Santana da Boa Vista (Schultz et al., 2000; Horn et al., 2015;
Pacheco et al., 2018). A U-Pb zircon geochronology study indicates a
233.23 ± 0.73 Ma age for the Alemoa site, Hyperodapedon AZ in Santa
Maria city (Langer et al., 2018). This datum places the Hyperodapedon
AZ in the middle Carnian, which is congruent with previous biostrati­
graphic proposals (e.g., Abdala et al., 2001; Langer et al., 2007b).
4.3.2. Tetrapod faunal content
The fossiliferous content of the Hyperodapedon Assemblage Zone
(=Rhynchosauria Cenozone; Schultz et al., 2000; Abdala et al., 2001),
includes (Figs. 8 and 9) a rich and diverse paleofauna of tetrapods
(Fig. 5), the fossil record of which is mostly composed of diapsids and
synapsids (e.g., Langer et al., 2005, 2007; Pavanatto et al., 2018; Garcia
et al., 2019). This assemblage zone, that was renamed by Lucas (2001)
from previous interpretations (e.g., Barberena, 1977, 1985; Schultz
et al., 2000; Abdala et al., 2001) and posteriorly redefined in subsequent
papers (e.g., Langer, 2005; Langer et al., 2007b), is characterized by the
presence and abundance of rhynchosaurs attributed to the genus
Hyperodapedon, a geographically widely distributed rhynchosaur, in
association to other taxa (see below). Moreover, this AZ is also

Fig. 9. Selected fossils from the lower portion of the Hyperodapedon AZ. A, skull of Hyperodapedon huenei (UFRGS-PV-0132-T) in ventral view; B, assembled skull
fragments of Compsocerops sp. (ULBRA PVT 059a-b) in dorsal view; C, skull and mandible of Gnathovorax cabreirai (CAPPA/UFSM 0009) in right lateral view; D,
partial skull and mandible of Clevosaurus hadroprodon (MMACR-PV-027-T) in right lateral view; E, partial right mandibular ramus of Prozostrodon brasiliensis (CAPPA/
UFSM 0123) in medial view; F, skull of Aetosauroides scagliai (UFSM 11505) in right lateral view; G, skull of Buriolestes schultzi (CAPPA/UFSM 0035) in right
lateral view.

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C.L. Schultz et al. Journal of South American Earth Sciences 104 (2020) 102846

remarkable by the presence of the oldest dinosaurs worldwide (e.g.,
Colbert, 1970; Langer et al., 1999; Cabreira et al., 2016; Müller et al.,
2018a,b; Pretto et al., 2019; Pacheco et al., 2019), as well as some of the
oldest aetosaurs, including one of the earliest diverging branches of
Aetosauria (Desojo and Ezcurra, 2011; Brust et al., 2018), as well as
possessing an important record of cynodonts (Abdala et al., 2002b;
Bonaparte and Barberena, 1975; Oliveira et al., 2010; Martinelli and
Soares, 2016; Martinelli et al., 2017c).
Additionally, the fossiliferous content of the Hyperodapedon AZ has
been divided into distinct portions according to the relative abundance

Fig. 10. Selected fossils from the upper portion of the Hyperodapedon AZ (Exaeretodon sub-AZ). A, skull of Teyumbayta sulcognathus (UFRGS-PV-0232-T) in ventral
view; B, skull of Proterochampsa nodosa (MCP 1694 PV) in dorsal view; C, skull and mandible of Exaeretodon riograndensis (CAPPA/UFSM 0033) in left lateral view; D,
skull and mandible of Siriusgnathus niemeyerorum (CAPPA/UFSM 0032) in left lateral view (but see discussion in the text); E, skull and mandible of Trucidocynodon
riograndensis (CAPPA/UFSM 0029) in left lateral view; F, skull of Bagualosaurus agudoensis (UFRGS-PV-1099-T) in right lateral view; G, assembled elements of the
skull and mandible of Dynamosuchus collisensis (CAPPA/UFSM 0248) in left lateral view.

14
C.L. Schultz et al.
of Hyperodapedon and the cynodont Exaeretodon (Langer et al., 2007a,b;
Oliveira and Schultz, 2007; Pretto et al., 2015; Garcia et al., 2019;
Müller and Garcia, 2019) (Figs. 9 and 10). Accordingly, the abundance
of Hyperodapedon is suggested to be high in the lower portion, also
named as Hyperodapedon Acme Zone (sensu Langer et al., 2007b), while
in the upper portion the representativity of Hyperodapedon declines,
while the presence of Exaeretodon increases (thus possibly character­
izing an Exaeretodon sub-AZ). A similar numerical faunistic replacement
is reported for the Ischigualasto Formation, in Argentina, where the top
of the section is dated as early Norian in age (Martínez et al., 2013).
Therefore, the temporal extension of the Hyperodapedon AZ might even
range from the middle Carnian to early Norian.
The record of temnospondyl amphibians in the Hyperodapedon AZ is
limited to the genus Compsocerops (Dias-da-Silva et al., 2012) and some
indeterminate stereospondyls (Dias-da-Silva et al., 2011). The same
genus is recovered from the upper Maleri Formation, in India (Segunpta,
1995), which is considered (?)late Norian to earliest Rhaetian in age
(Novas et al., 2010). However, the Brazilian records came from the
Pivetta site (São João do Polêsine), which lacks any material of Exaer­
etodon. Indeed, the fossil record of the Pivetta site resembles the typical
occurrences of the Hyperodapedon Acme Zone, with several specimens of
Hyperodapedon and no records of traversodontid cynodonts whatsoever.
Therefore, records of Compsocerops from Brazil and India may not be
coeval.
The Hyperodapedon AZ also yielded the oldest Gondwanan records of
sphenodonts, with the occurrence of Clevosaurus hadroprodon (Hsiou
et al., 2019; see also Romo de Vivar et al., 2019). Though fragmentary,
the specimen also represents the oldest true acrodont sphenodont, and
suggests that the Gondwanan strata preserve an important step on the
early radiation of sphenodonts.
The archosauromorph fossil record is far more representative. In
addition to Hyperodapedon, this AZ also includes the rhynchosaur
Teyumbaita sulcognathus (Azevedo and Schultz, 1987; Montefeltro et al.,
2010), recorded in the upper layers of the Alemoa site and in the Linha
Facão site. The presence of traversodontid remains (probably Exaer­
etodon, MMACR PV T 020) close to the type locality of Teyumbaita link
the taxon to the upper levels of the Hyperodapedon AZ (the Exaeretodon
sub-AZ). The fossil record of non-archosaur achosauromorphs also in­
cludes the two genera of Proterochampsidae excavated from the mu­
nicipality of Santa Maria: Rhadinosuchus gracilis (Huene, 1938a, b) and
Cerritosaurus binsfeldi (Price, 1946). These taxa are recorded from clas­
sical localities of the Hyperodapedon Acme Zone. Proterochampsids are
also present in the Exaeretodon-dominated portions of the Hyper­
odapedon AZ, with the record of the large-sized Proterochampsa nodosa
(Barberena, 1982).
Among Archosauria, aetosaurs are represented by three genera:
Aetosauroides scagliai (Casamiquela, 1960; Brust et al., 2018), Aeto­
barbakinoides brasiliensis (Desojo et al., 2012), and Polesinesuchus aurelioi
(Roberto-da-Silva et al., 2014). These aetosaurs have been recorded
from Hyperodapedon-abundant strata and represent some of the oldest
known aetosaurs (Desojo and Ezcurra, 2011). Aetosauroides scagliai is
also recorded from the Scaphonyx-Exaeretodon-Herrerasaurus biozone
(probably related to the Brazilian Hyperodapedon Acme Zone) and the
Exaeretodon biozone (equivalent to Brazilian Exaeretodon sub-AZ) of
Argentina (Martínez et al., 2013). Other members of Pseudosuchia from
the Brazilian Hyperodapedon AZ include the loricatan Rauisuchus tira­
dentes, which has been found as the sister taxon to a clade composed of
Postosuchus and Polonosuchus silesiacus (e.g., Roberto-da-Silva et al.,
2018). Polonoschus silesiacus was exhumed from Carnian beds of Poland
(Brusatte et al., 2009), while Postosuchus is recorded from Norian (lower
Cooper Canyon Formation, Dockum Group) strata of USA (e.g., Wein­
baum, 2013). Another pseudosuchian in the Hyperodapedon AZ is
Dynamosuchus collisensis, an ornithosuchid with close affinities to
Venaticosuchus rusconii (Müller et al., 2020). The only known specimen
of Venaticosuchus rusconii was excavated from the middle section of the
Ischigualasto Formation, at Cerro Las Lajas, La Rioja, Argentina (von
15
Journal of South American Earth Sciences 104 (2020) 102846
Baczko, 2018). This locality has yielded traversodontid cynodonts, pu­
tatively associated with the genus Exaeretodon (Desojo et al., 2020b).
The same is true for the type-locality of Dynamosuchus, which is domi­
nated by Exaeretodon riograndensis.
Ornithodiran archosaurs are well represented in the AZ through the
record of several dinosaurs and closely related forms. Staurikosaurus
pricei (Colbert, 1970) and Gnathovorax cabreirai (Pacheco et al., 2019)
are herrerasaurids from typical Hyperodapedon Acme Zone sites. The
first one was collected in the municipality of Santa Maria, from a locality
close to the Alemoa Complex, dated as c. 233 Ma, while the second came
from São João do Polêsine. Both taxa are related to Argentinean her­
rerasaurids from the Ischigualasto Formation. Except for Nhandumirim
waldsangae (Marsola et al., 2019), all the other dinosaurs from the
Hyperodapedon AZ have been recovered as members of Sau­
ropodomorpha (but see Pacheco et al., 2019, who recovered N. wald­
sangae as a sauropodomorph, sister-group to Saturnaliinae). The first
sauropodomorph described for the Hyperodapedon AZ was Saturnalia
tupiniquim (Langer et al., 1999), from the Alemoa site, Santa Maria.
Later, Pampadromaeus barberenai (Cabreira et al., 2011) was recovered
from the Janner site, Agudo, one of the localities with high abundance of
Exaeretodon. The Janner site also yielded Bagualosaurus agudoensis
(Pretto et al., 2019). From the Buriol site, at the municipality of São João
do Polêsine (close to the type-locality of Gnathovorax cabreirai), were
excavated the skeletons of Buriolestes schultzi (Cabreira et al., 2016;
Müller et al., 2018a,b). In the same excavation area also were exhumed
the specimens of Ixalerpeton polesinensis (Cabreira et al., 2016), a
lagerpetid dinosauromorph.
Both Buriolestes and Saturnalia (as well as Nhandumirim) are recov­
ered from Hyperodapedon dominated strata, while Pampadromaeus and
Bagualosaurus come from an Exaeretodon-rich outcrop. Indeed, both
Pampadromaeus and Bagualosaurus bear a set of dental traits more
related to an omnivorous/herbivorous diet when compared to other
putatively faunivorous sauropodomorphs, like Buriolestes schultzi and
Saturnalia tupiniquim (Cabreira et al., 2016; Bronzati et al., 2019; Pretto
et al., 2019). Considering that the herbivory seems to be a trait that
originated secondarily in the group, this seems in accordance with a
putative slight younger age for the Exaeretodon dominated strata, as
suggested by the Argentinean succession.
Synapsids are represented by cynodonts, while dicynodonts are so far
completely absent in the fossil record of the Hyperodapedon AZ. Curi­
ously, the coeval Argentinean beds from the Ischigualasto Formation
yielded several specimens of the stahleckeriid dicynodont Ischigualastia
jenseni (Cox, 1962). Probainognathian cynodonts are far less abundant
than traversodontids. On the other hand, the group is more diverse, with
about five genera described. Prozostrodon brasiliensis (Bonaparte and
Barberena, 2001), Therioherpeton cargnini (Bonaparte and Barberena,
2001), and Alemoatherium huebneri (Martinelli et al., 2017c) are pro­
zostrodontians known from typical localities of the Hyperodapedon Acme
Zone. In São João do Polêsine, Prozostrodon brasiliensis was found in
association to Hyperodapedon and Gnathovorax (see Pacheco et al.,
2019). Charruodon tetracuspidatus (Abdala and Ribeiro, 2000; Martinelli
et al., 2017c) was excavated from a fossiliferous locality in Candelária
that also yielded Exaeretodon and Proterochampsa. This cynodont taxon
is poorly known, given the fragmentary condition of the holotype
(Martinelli et al., 2017c). In contrast, the ecteniid Trucidocynodon riog­
randensis (Oliveira et al., 2010) is known on the basis of a nearly com­
plete skeleton (holotype). Like Charruodon tetracuspidatus, the specimens
of Trucidocynodon riograndensis have been discovered from strata (Jan­
ner site) that also yielded several specimens of Exaeretodon (Oliveira
et al., 2010; Stefanello et al., 2018). Unlike T. riograndensis, the Argen­
tinean ecteniniids (e.g., Ecteninion and Diegocanis) were all collected
from the lower portions of the Ischigualasto Formation, where Hyper­
odapedon is far more abundant than Exaeretodon (Martínez et al., 2013).
Traversodontid cynodonts are represented by Gomphodontosuchus bra­
siliensis (Huene, 1928), Exaeretodon riograndensis (Abdala et al., 2002b)
and possibly Siriusgnathus niemeyerorum (Pavanatto et al., 2018, but see
C.L. Schultz et al. Journal of South American Earth Sciences 104 (2020) 102846

Miron et al., 2020). The first one comes from the municipality of Santa
Maria and is poorly known, while Exaeretodon riogradensis is the best
sampled non-mammaliaform cynodont from Brazil. In the Janner site
(Agudo) the abundance of Exaeretodon riograndesis easily overwhelms all
the other vertebrate occurrences. The biostratigraphic assignment of
Siriusgnathus is, however, controverse. This taxon is the major compo­
nent of the sampled fauna at its type locality (Pavanatto et al., 2018),
and other fossil vertebrates do not help to refine its biostratigraphic
placement due to their fragmentary preservation and indeterminate
taxonomic status. However, Siriusgnathus was recently reported outside
its type-locality (Miron et al., 2020), in an outcrop which was suggested
to be correlated to the Sacisaurus site (Marsola et al., 2019), of (?)Norian
age (see also next section). Nonetheless, this second locality with Sir­
iusgnathus also lacks the typical index fossils from the Brazilian Triassic
AZs and other putative faunal components also found in the Sacisaurus
site. Therefore, though there is growing evidence of the presence of

Fig. 11. Chrono- and biostratigraphy of Triassic units with vertebrate Assemblages Zones from southern Brazil, and main localities (A) and the tetrapod fossil content
(B) of the Riograndia AZ. Absolute ages (indicated by arrows) as in Fig. 1. The ages of the column follow Gradstein et al., 2012. Abbreviations: Ar, Archosauria; D,
Dinosauria; Ind. Induan; Ole, Olenekian; P, Probainognathia.

16
C.L. Schultz et al. Journal of South American Earth Sciences 104 (2020) 102846

traversodontids in Norian Brazilian strata, with Siriusgnathus possibly
representing the youngest traversodontid cynodont record, further
chronostratigraphic and/or biostratigraphic data are needed to corrob­
orate this hypothesis.
The faunistic composition of Hyperodapedon AZ indeed resembles
that of the Ischigualasto Formation in Argentina, which was radioiso­
topically dated as Carnian to Norian (Martínez et al., 2011). However,
the presence of endemic taxa in some Brazilian outcrops results in a
complex scenario. For instance, the presence of ecteniniid cynodonts in
the Janner site differs from the pattern of Argentinean cynodont distri­
bution. Therefore, the turnover in the relative abundances of Hyper­
odapedon versus Exaeretodon, which took place in Argentina during the
Carnian (Martínez et al., 2013) may not have been coeval to the same
event in Brazil. Such issue demands further radioisotopic data for the
fossiliferous localities from Brazil and additional fieldwork in order to
reach a better sampled and more reliable picture of the faunal compo­
sition of each area. Nevertheless, a promising scenario is being revealed
along the last years. Several new specimens have been excavated and the
first U-Pb zircon geochronology investigations are being performed in
localities referred to the Hyperodapedon AZ. Better-constrained ages will
help to construct a more robust framework of this particular moment of
Earth’s life history when the first dinosaurs arose and synapsids expe­
rienced their last moment of supremacy during the Mesozoic Era.
4.4. Candelária Sequence (top) – Riograndia AZ
4.4.1. Geological setting
The main characteristic of the upper portion of the Candelária
Sequence (Fig. 11) is the dominance of fine sandstones, in contrast with
the base, dominated by mudstones. These sandstones are fine-grained
and dominantly massive, sometimes with disperse intraclasts. They
are interpreted as product of mass flows, caused by downpours in a
seasonal climate, in which a high grain fallout rate associated with flow
deceleration prevents the formation of tractive structures. Stratified
sandstones occur subordinately and show planar bedding with thick
lamination, low angle cross-stratification and climbing ripples. Mud­
stones occur as mud drapes or centimetric layers. These rocks are
interpreted as a hyperconcentred flow-dominated, ephemeral fluvial

Fig. 12. Selected fossils from the Riograndia AZ. A, skull and jaw of Riograndia guaibensis (UFRGS-PV-596-T) in right lateral view; B, skull and jaws of Brasilodon
quadrangularis (UFRGS-PV-1030-T, formerly holotype of Minicynodon) in left lateral view; C, left jaw of Botucaraitherium belarminoi (MMACR-PV-003-T) in medial
view; D, skull of Jachaleria candelariensis (UFRGS-PV-151-T) in left lateral view; E, partial left jaw of Cargninia enigmatica (UFRGS-PV-1027-T) in lateral view; F, left
jaw of Soturnia caliodon (UFRGS-PV-1234-T) in lateral view; G, skull of Clevosaurus brasiliensis (UFRGS-PV-748-T) in dorsal view; H, skull and jaws of Macrocollum
itaquii (CAPPA/UFSM 0001a) in left lateral view; I, snout fragment of Phytosauria indet. (MCN-FZB 1865) in ventral view.

17
C.L. Schultz et al.
system with lateral avulsions and frontal deconfinement, formation
sheetfloods (Horn et al., 2018a). Lithostratigraphycally, this portion
corresponds to the lower portion of the Caturrita Formation (Andreis
et al., 1980).
The upper portion, composed by arcosian sandstones with trough
cross stratifications, devoid of tetrapods but bearing silicified logs, was
understood as another third-order sequence by Zerfass et al. (2003) and
named Mata Sequence (Fig. 1).
4.4.2. Tetrapod faunal content
The faunal association of the Riograndia AZ (Figs. 11 and 12) is not so
diverse as those of Dinodontosaurus and Hyperodapedon AZs, but shows
high potential since systematic prospecting work in the fossiliferous
outcrops of the top of the Candelária sequence only started at the end of
the 1990s (Bonaparte et al., 2010). Nevertheless, it differs significantly
from other faunal assemblages by the remarkable prevalence of micro­
vertebrates (animals with less than 5 kg; Fisher, 1981). These are rep­
resented by procolophonids, non-rhynchocephalian and
rhynchocephalians lepidosauromorphs, non-mammaliaform cynodonts,
and a possible basal ornithodira. Larger forms include temnospondyls,
dicynodonts, phytosaurs, non-dinosaurian dinosauriforms, and di­
nosaurs (see Fig. 13).
The record of temnospondyls in Riograndia AZ is limited to an
indeterminate Mastodonsauroidea Stereropondlyli (Dias-da-Silva et al.,
2009) collected in the Botucaraí Hill site (Dias-da-Silva et al., 2009;
Bittencourt et al., 2012b).
Procolophonids and lepidosauromorphs are amongst the micro­
vertebrates that compose the Riograndia AZ. The procolophonid Soturnia
caliodon, from Linha São Luiz site, is a member of the Leptopleurinae
clade, corresponding to the first Gondwanan leptopleurine (Cisneros and
Schultz, 2003). The first known Lepidosauromorpha from the Riograndia
AZ was the eusphenodontian rhynchocephalian Clevosaurus brasiliensis
(Bonaparte and Sues, 2006), with dozens specimens collected in Linha
São Luiz and Sesmaria do Pinhal 1 sites (Ferigolo, 2000; Arantes et al.,
Fig. 13. Taxonomic tetrapod diversity among the biozones of the Brazilian Triassic. The numbers over the bars indicate the number of species or of novel taxonomic
representive of each group (based on the list of tetrapod content for each AZ used in previous figures).
18
Journal of South American Earth Sciences 104 (2020) 102846
2009; Hsiou et al., 2015; Romo de Vivar and Soares, 2015; Romo-de-­
Vivar et al., 2020a). A second taxon, Lanceirosphenodon ferigoloi, was
described by Romo de Vivar et al. (2020a) and represents one of the
basalmost eusphenodonts. Outside Rhynchocephalia is the lepidosaur­
omorph Cargninia enigmatica, from Linha São Luiz site (Bonaparte et al.,
2010a). An anatomical revaluation of Cargninia holotype provided by
Romo-de-Vivar et al. (2020b) suggested it could be a basal Lepidosauria.
The therapsids of the Riograndia AZ are represented by a single
dicynodont and several species of non-mammaliaform cynodonts. The
tuskless dicynodont Jachaleria candelariensis (Araújo and Gonzaga,
1980) is one of most derived kannemeyeriiform, closely related to the
Argentine J. colorata (Bonaparte, 1971; Vega-Dias et al., 2004). It was
the first tetrapod recovered from the beds of the Riograndia AZ, at the
Botucaraí Hill site.
Virtually all non-mammaliaform cynodonts of the Riograndia AZ fit
into the category of microvertebrates and belong to Prozostrodontia
clade of Probainognathia (Liu and Olsen, 2010). Riograndia guaibensis
(Bonaparte et al., 2001) is the most abundant non-mammaliaform cyn­
odont (that is why the name of the biozone). Its record in the Botucaraí
Hill site is restricted to an isolated incisor from the top of the outcrop
(Soares et al., 2011b). On the other hand, dozens of specimens were
recovered from the Sesmaria do Pinhal 1 and Linha São Luiz sites. A
postcanine tooth assigned to Riograndia (Ribeiro et al., 2011) was also
found in the Sacisaurus site, but Marsola et al. (2018) opted to treat it as
a Riograndia-like tooth, considering that it could pertain to another
prozostrodontian. Riograndia is a small basal ictidosaur closely related to
Tritheledontidae clade, in which Irajatherium hernandezi is nested
(Martinelli et al., 2005; Soares et al., 2011b; Oliveira et al., 2011a). The
record of Irajatherium is from Sesmaria do Pinhal 1 and Linha São Luız
sites (Martinelli et al., 2005; Oliveira et al., 2011a). Several specimens of
Brasilodon quadrangularis and Brasilitherium riograndensis also occur in
the Sesmaria do Pinhal 1 and especially in the Linha São Luiz site
(Bonaparte et al., 2003, 2005, 2012, ). A third record of Brasilitherium
was based on postcanine teeth found in the Sacisaurus site (Ribeiro et al.,
C.L. Schultz et al.
2011), but Marsola et al. (2018) interpreted them as belonging to a
likely new prozostrodontian. Bonaparte et al. (2005) erected the family
Brasilodontidae to nest Brasilodon and Brasilitherium and recovered it as
the sister-group of the Mammaliaformes. However, in subsequent ana­
lyses (e.g., Abdala, 2007; Liu and Olsen, 2010) Brasilodontidae appeared
as paraphyletic. Additionally, Liu and Olsen (2010) considered Brasili­
therium, as junior synonym of Brasilodon what was also raised by Mar­
tinelli and Bonaparte (2011; see also Martinelli, 2017; Martinelli et al.,
2017d). Despite these controversies, the status of Brasilodon and/or
Brasilitherium as the closest relatives of Mammaliaformes remains
consensual. Minicynodon maieri is a ‘brasilodontid’ from the Linha São
Luiz site (Bonaparte et al., 2010), but because of the strong resemblance
with Brasilitherium, it may be a juvenile of this taxon (or Brasilodon if the
former synonymization is confirmed; see Martinelli, 2017; Martinelli
et al., 2017d). Another prozoztrodontian closely related to Brasilodon is
Botucaraitherium belarminoi, exclusive from the Botucaraí Hill site
(Soares et al., 2014).
Remains of traversodontids are still scarce in the Riograndia AZ. They
include a lower incisor asssigned to Gomphodontosuchinae (Ribeiro
et al., 2011; Marsola et al., 2018) from the Sacisaurus site, and some
teeth and a fragment of maxilla resembling Exaeretodon (typical of the
Hyperodapedon AZ) collected in Sesmaria do Pinhal 3 site. If the taxo­
nomic identity is confirmed, these teeth would correspond to the
youngest record of traversodontids in South America.
Archosaurs are represented by a phytosaur and ornithodirans,
including dinosaurs. The only record of Phytosauria are rostral frag­
ments found scattered on the base of the Botucaraí Hill site (Kischlat and
Lucas, 2003). Although not providing enough anatomical information
for more accurate taxonomic identification, this specimen represents the
only phytosaur record in South America (Kischlat and Lucas, 2003).
Whitin Ornithodira, a possible basal form is the small Faxinalipterus
minima, found in the Linha São Luiz site (Bonaparte et al., 2010b; Soares
et al., 2013). Non-dinosaurian Dinosauriformes are represented by the
Silesauridae Sacisaurus agudoensis, the most conspicuous fossil of the
Sacisaurus site as it is alluded by the name of the outcrop (Ferigolo and
Langer, 2007; Langer and Ferigolo, 2013).
The dinosaur Guaibasaurus candelariensis comes from two localities of
Riograndia AZ, Sesmaria do Pinhal 2 (Bonaparte et al., 1999) and Linha
São Luiz sites (Bonaparte et al., 2007). Even though the postcranial
skeleton of Guaibasaurus is almost complete, nothing about the anatomy
of the skull and dentition is known. Bonaparte et al. (1999, 2007)
recognized Guaibasaurus as a basal saurischian, but Langer et al. (2007a,
2010) sustained Guaibasaurus as a theropod.
Several detached ziphodont teeth (Dornelles, 1990) were the first
signs of the presence of Dinosauria in the Botucaraí Hill site. Later,
isolated post-cranial bones from sauropodomorph dinosaurs (Bitten­
court et al., 2012a, 2012b; Pretto et al., 2017) and a fragmented femur of
a probable Neotheropoda (Pinheiro, 2016) were reported. Ezcurra
(2017) casted in doubt the interpretation of the femur as belonging to a
Neotheropoda but considers it may be one of the sister-taxa of the clade.
The occurrences of ‘prosauropods’ in the Riograndia AZ are more
problematic because they represent isolated records, making correla­
tions with other outcrops difficult. Unaysaurus tolentinoi was found
solely in the Água Negra site (Leal et al., 2004) whereas Macrocollum
itaqui is from the Wachholz site (Müller et al., 2015). Macrocollum was
found in the new clade Unaysauridae (the sister-group of Plateosauria)
that also gathers Unaysaurus and the Indian Jaklapallisaurus asymmetrica
(Müller et al., 2015). Additional findings of several isolated bones
resembling Unaysaurus came from Campinas, near Santa Maria, and
Sacisaurus site (Marsola et al., 2018).
The whole Riograndia AZ fauna allows some biostratigraphic corre­
lations. The record of the dicynodont Jachaleria candelariensis in the
Botucaraí Hill site enables a direct correlation with the base of the Los
Colorados Formation (early Norian) of the Ischigualasto-Villa Unión
Basin, Argentina, which records Jachaleria colorata (Fonseca and
Scherer, 1998). The early Norian age for the Botucaraí Hill site (Rubert
19
Journal of South American Earth Sciences 104 (2020) 102846
and Schultz, 2004; Soares et al., 2011) has remained stable, but from the
late 1990s, with fossils findings in other localities, the biostratigraphic
context became more complex. For example, the presence of the icti­
dosaur Riograndia and the tritheledontid Irajatherium would allow the
correlation of Riograndia AZ with the upper third of Los Colorados
Formation (La Esquina Local Fauna, Late Norian) where the trithele­
dontid Chaliminia musteloides comes from (Martinelli and Rougier, 2007;
Kent et al., 2014). However, the more basal phylogenetic position of the
Brazilian ictidosaurs could indicate an older age for Riograndia AZ. The
record of plateosaurian ‘prosauropods’ in the La Esquina Local Fauna (e.
g., Coloradisaurus, Riojasaurus) (Arcucci et al., 2004) also would aim a
correlation with the Riograndia AZ. Yet, some traits of the Brazilian
Unaysauridae are more plesiomorphic than those of the Argentine pla­
teosaurians, suggesting an early Norian age for the Brazilian taxa
(Müller et al., 2018a,b). Another Argentine unity whose faunal content
resembles the Riograndia AZ is Quebrada del Barro Formation, in San
Juan Province. It encompasses tritheledontids, rhynchocephalians, and
basal sauropodomorphs, among others. The resemblance with the
Riograndia AZ is especially due to the tritheledontid cynodonts and
sphenodonts. However, the Argentine opisthodontian Sphenotitan leyesi
is more derived than Lanceirosphenodon and Clevosaurus brasiliensis,
what sets the fauna of Quebrada del Barro as younger (Norian-Rhaetian
age) than those of Riograndia AZ (Martínez et al., 2013).
In sum, the biostratigraphic framework resulting from all studied
fossils has pointed out an early Norian age for the Riograndia AZ. This
relative dating found support in the radiometric dating performed by
Langer et al. (2018) (~225.42 ± 0.37 Ma). However, it is important to
emphasize that this dating was taken in one only point, the Linha São
Luiz site. Based on the faunal content, the correlation of the Linha São
Luiz site with at least Sesmaria do Pinhal 1 and 2 (Candelária) is well
grounded. The link with Botucaraí Hill site is still weak, once is solely
based on a supposed lower incisor of Riograndia from the higher levels of
the outcrop (Soares et al., 2011b). More collect efforts are needed in
Água Negra, Campinas, and Wachholz sites since, up to now, no fossil
guides typical of the Riograndia AZ (e.g., Riograndia, Jachaleria, Guai­
basaurus) were recorded there. Moreover, new radiometric dating on all
these localities would be fundamental to help in clarifying this matter.
5. Final remarks
The end of the Permian marked the end of a tectonic and depositional
cycle across southwestern Gondwana, both in the Chaco-Paraná Basin
and in the Karoo Basin. It is the end of the widespread continental
sedimentation of this basin. In Uruguay, a large gap separates the Buena
Vista Formation (Late Permian - Lopingian) from the Tacuarembó For­
mation (Early Jurassic). In southern Africa and Madagascar, a deposi­
tional gap close to the P-T boundary precedes the start of a new
depositional sequence, with different characteristics from the underly­
ing package. In southern Brazil, SCS represents the only Eotriassic
package containing tetrapods for the entire country. It deposition took
place on a compressive stage associated with Paleopacific margin of the
Southwest Gondwana (Zerfass et al., 2004). In paleofaunistic terms,
Procolophon AZ presents a typical recovery fauna of post P-T extinction,
with little diversity (Fig. 12) and composed of small animals that
evolved from the few remaining forms of Permian fauna (Erwin, 1994;
Benton and Twitchett, 2003; Benton et al., 2004; Sahney and Benton,
2008; Day et al., 2015). This deposition is associated with highly sea­
sonal rivers and arid climate. After a depositional hiatus of about 5
million years, extensional tectonic stresses took place in the SW margin
of Gondwana and led to the generation of an arch of small depositional
basins, aligned from western Argentina to India (Zerfass et al., 2003,
2004). It was in this new tectonic context that the deposition of SMS
began, with its 4 successive tetrapod biozones. In the Middle-earliest
Late Triassic, Dinodontosaurus AZ and Santacruzodon AZ already show
an increase in diversity in their tetrapod fauna (Fig. 12), with a pre­
dominance of therapsids. The sedimentological (predominance of loess
C.L. Schultz et al.
deposits) and geochemical (basic environment with low chemical wea­
tering) characteristics point to the time span in question the predomi­
nance of a dry and hot, strongly seasonal climate (Horn et al., 2018b;
Queiroz et al., submitted). Corroborating this evidence, the two AZs
mentioned above record a complete absence of amphibians and forms
that can be considered semi-aquatic (proterochampsids) are rare.
In the transition from Middle to Late Triassic, there is a considerable
change in the sedimentological (presence of underwater deposits) and
geochemical (more acidic conditions) patterns of the SSM rocks, indi­
cating a humidity increase, but still with a semi-arid CIA signature (Horn
et al., 2018b). Accordingly, there is a major change in the tetrapod
faunal content in the corresponding biozones. In the Hyperodapedon AZ,
diapsids (rhynchosaurs and dinosaurs) become the dominant herbivor­
es/omnivores, ending a synapsid echological supremacy that has lasted
since the Eopermian. In addition, this biozone has the highest degree of
diversity among all the AZs of the Brazilian Triassic (Fig. 2). Apparently,
this sudden climate change and the establishment of a period of higher
humidity coincide with the Carnian Pluvial Event (e.g., Simms and
Ruffell, 1989; Benton et al., 2018), reinforcing the hypothesis that this
event had global effects at that time. The sedimentological and faunal
evidences of Riograndia AZ, in turn, points towards a decrease in hu­
midity (although still far from the prevailing arid conditions present in
Mesotriassic) which is reflected in a slight loss of diversity in the
tetrapod fauna, with a predominance of dinosaurian forms and
small-sized derived non-mammaliaform probainognathians.
The younger epochs of the Triassic (end of the Norian and Rhaetian),
as well as nearly the entire Jurassic are not represented in the sedi­
mentary package of southern Brazil.
Author statements
Conceptualization: CLS; Investigation: all authors; Roles/Writing -
original draft: all authors; Writing - review & editing: all authors.
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.
Acknowledgements
Initially, we would like to thank Drs. Claudia Marsicano, Leandro
Gaetano and Randall Irmis for the invitation to produce a chapter for
this Special Volume on the Triassic at JSAES. For access to collections we
thank: Carlos Nunes Rodrigues (MMACR), Belarmino Stefanello
(MMACR), Ana Maria Ribeiro (MCN-FZB), Jorge Ferigolo (MCN-FZB),
Marco Brandalise de Andrade (MCP-PUC), Rodrigo Machado (MCT),
and Diógenes de Almeida Campos (MCT). Several photographs of this
contribution were skillfully taken by Luiz Flávio Lopes (UFRGS). MBS is
supported by grant from Conselho Nacional de Desenvolvimento Cien­
tífico e Tecnológico (CNPq process numbers 407969/2016-0). FLP is
supported by grants from Conselho Nacional de Desenvolvimento
Científico e Tecnológico (CNPQ process numbers 407969/2016-0,
305758/2017-9) and Fundação de Amparo à Pesquisa do Estado do Rio
Grande do Sul (FAPERGS process number 16/2551-0000271-1). Finally,
the comments made by the reviewers Max C. Langer and Martín D.
Ezcurra greatly improved the ms.
References
Abdala, F., Giannini, N.P., 2000. Gomphodont cynodonts of the Chañares Formation: the
analysis of an ontogenetic sequence. J. Vertebr. Paleontol. 20, 501–506. https://doi.
org/10.1671/0272-4634(2000)020[0501:GCOTCA]2.0.CO;2.
Abdala, F., Ribeiro, A.M., 2000. A new therioherpetid cynodont from the Santa Maria
Formation (middle Late Triassic), southern Brazil. Geodiversitas 22 (4), 589–596.
20
Journal of South American Earth Sciences 104 (2020) 102846
Abdala, F., Giannini, N.P., 2002. Chiniquodontid cynodonts: systematic and
morphometric considerations. Palaeontology 45 (6), 1151–1170.
Abdala, F., Ribeiro, A.M., 2003. A new traversodontid cynodont from the Santa Maria
Formation (Ladinian-Carnian) of southern Brazil, with a phylogenetic analysis of
Gondwanan traversodontids. Zool. J. Linn. Soc. 139, 529–545. https://doi.org/
10.1111/j.1096-3642.2003.00096.x.
Abdala, F., Sá-Teixeira, A.M., 2004. A traversodontid cynodont of African affinity in the
South American Triassic. Palaeontol. Afr. 40, 11–22.
Abdala, F., Smith, R.M.H., 2009. A middle Triassic cynodont fauna from Namibia and its
implications for the biogeography of Gondwana. J. Vertebr. Paleontol. 29, 837–851.
Abdala, F., Ribeiro, A.M., Schultz, C.L., 2001. A Rich Cynodont Fauna of Santa Cruz Do
Sul, Santa Maria Formation (Middle-Late Triassic), Southern Brazil. Neu Jb Geol
Paläont, Mh, pp. 669–687, 2001.
Abdala, F., Dias-da-Silva, S., Cisneros, J.C., 2002a. First record of nonmammalian
cynodonts (Therapsida) in the Sanga do Cabral Formation (early Triassic) of
southern Brazil. Palaeontol. Afr. 38, 93–98.
Abdala, F, 2007. Redescription of Platycraniellus elegans (Therapsida, Cynodontia) from
the Lower Triassic of South Africa, and the cladistic relationships of eutheriodonts.
Palaeontology 50, 591–618.
Abdala, F., Barberena, M.C., Dornelles, J., 2002b. A new species of the traversodontid
cynodont Exaeretodon from the Santa Maria Formation (Middle/Late Triassic) of
southern Brazil. J. Vertebr. Paleontol. 22 (2), 313–325.
Abdala, F., Martinelli, A.G., Soares, M.B., de la Fuente, M., Ribeiro, A.M., 2009. South
American Middle Triassic continental faunas with amniotes: biostratigraphy and
correlation. Palaeontol. Afr. 44, 83–87.
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 (3), 1151–1162.
Andreis, R.R., Bossi, G.E., Montardo, D.K., 1980. O Grupo Rosário do sul (triássico) no
Rio Grande do sul-Brasil. XXXI Congresso Brasileiro de Geologia, Camboriú,
Sociedade Brasileira de Geologia. Anais 2, 659–673.
Andreis, R.R., Ferrando, L., Herbst, R., 1996. Terrenos Carboniferos y Pérmicos de
Republica Oriental del Uruguay. In: Archangelsky, S. (Ed.), El Sistema Pérmico en la
República Argentina y en la República Oriental del Uruguay. Academia Nacional del
Uruguay, Córdoba, Argentina, pp. 309–343.
Arantes, B.A., Soares, B.M., Schultz, C.L., 2009. Clevosaurus brasiliensis (Lepidosauria,
Sphenodontia) do Triássico Superior do Rio Grande do Sul: Anatomia pós-craniana e
relações filogenéticas. Rev. Bras. Palaontol. 12 (1), 199–210.
Araújo, D.C., Gonzaga, T.D., 1980. Uma nova espécie de Jachaleria (Therapsida,
Dicynodontia) do Triássico do Brasil. Actas del II Congreso Argentino de
Paleontología y Bioestratigrafía. I Congreso Latinoamericano de Paleontología,
Buenos Aires, pp. 159–174, 1980.
Arcucci B, A, Marsicano A, C, Caselli T, A, 2004. Tetrapod association and
palaeoenvironment of the Los Colorados Formation (Argentina): a significant sample
from Western Gondwana at the end of the Triassic. Geobios 37, 557–568.
Azevedo A, S, Schultz L, C, 1987. Scaphonyx sulcognathus (sp. nov.), um novo
rincossaurídeo do neotriássico do Rio Grande do Sul, Brasil. In: Moura A, J,
Gilson MN, H, Campos A, D, Beurlen, G, Macedo CM, A, Brito AM, I (Eds.),
X Congresso Brasileiro de Paleontologia. Sociedade Brasileira de Paleontologia, Rio
de Janeiro, pp. 99–113.
Barberena, M.C., 1974. Contribuição Ao Conhecimento Dos Cinodontes Gonfodontes
(Cynodontia, Tritylodontoidea) Do Brasil. Universidade Federal do Rio Grande do
Sul, p. 194. Livre Docência Thesis.
Barberena, M.C., 1977. Bioestratigrafia preliminar da Formação Santa Maria. Pesquisas
7, 111–129.
Barberena, M.C., 1978. A huge thecodont from the Triassic of Brazil. Pesquisas 9, 62–75.
Barberena, M.C., 1981a. Uma nova espécie de Massetognathus (Massetognathus
ochagaviae, sp. nov.) da Formação Santa Maria, Triássico do Rio Grande do Sul.
Pesquisas 14, 181–195.
Barberena, M.C., 1981b. Novos materiais de Traversodon stahleckeri da Formação Santa
Maria (Triássico do Rio Grande do Sul). Pesquisas 14, 149–162.
Barberena, M.C., Lavina, E.L., Becker, M.R., 1981. Sobre a presença de tetrápodos na
Formação Sanga do Cabral (Grupo Rosário do Sul), Triássico do Rio Grande do Sul,
Brasil. In: Congresso Latino-Americano de Palaeontologia. Porto Alegre, Anais, vol.
2, pp. 295–306.
Barberena, M, 1982. Uma nova especie de Proterochampsa (P. nodosa, sp. nov.) do
Triassico do Brasil. Anais da Academia Brasileira de Ciencias 54, 127–141.
Barberena, M.C., Araújo, D.C., Lavina, E.L., 1985. Late Permian and Triassic tetrapods of
southern Brazil. Natl. Geogr. Res. 1, 5–20.
Beltrão, R., 1965. Paleontologia de Santa Maria e São Pedro do Sul, Rio Grande do Sul,
Brasil, vol. 2. Boletim do Instituto de Geociências, pp. 5–114.
Benton, M.J., Tverdokhlebov, V.P., Surkov, M.V., 2004. Ecosystem remodeling among
vertebrates at the Permian-Triassic boundary in Russia. Nature 432, 97–100.
Benton, M.J., Twitchett, R.J., 2003. How to kill (almost) all life: the end-Permian
extinction event. Trends Ecol. Evol. 18, 358–365.
Benton, M.J., Bernardi, M., Kinsella, C., 2018. The Carnian Pluvial episode and the origin
of dinosaurs. J. Geol. Soc. 175 (6), 2018–2049.
Bertoni-Machado, C., Holz, M., 2006. Biogenic fossil concentration in fluvial settings: an
example of a cynodont taphocoenosis from the Middle Triassic of southern Brazil.
Rev. Bras. Palaontol. 9, 273–282.
Besairie, H., 1972. Géologie de Madagascar. I. Les terrains sédimentaires. Ann.
Géologiques Madagascar, Service des Mines 35, 1–463.
Bittencourt, J.S., Leal, L.A., Langer, M.C., Azevedo, S.A., 2012a. An additional basal
sauropodomorph specimen from the Upper Triassic Caturrita Formation, southern
Brazil, with comments on the biogeography of plateosaurids. Alcheringa 36,
269–278.
C.L. Schultz et al.
Bittencourt, J.S., Da-Rosa, A.A.S., Schultz, C.L., Langer, M.C., 2012b. Dinosaur remains
from the ‘Botucaraí Hill’ (Caturrita Formation), late Triassic of South Brazil, and
their stratigraphic context. Hist. Biol. 25, 81–93. https://doi.org/10.1080/
08912963.2012.694881.
Bonaparte, J.F., 1966. Chiniquodon Huene (Therapsida-Cynodontia) en el Triasico de
Ischigualasto, Argentina. Acta Geol. Lilloana 8, 157–169.
Bonaparte, J.F., 1971. Los tetrápodos del sector superior de la Formación Los Colorados,
La Rioja, Argentina (Triásico Superior). I Parte. Opera Lilloana 22, 1–183.
Bonaparte F, J, Barberena C, M, 1975. A possible mammalian ancestor from the Middle
Triassic of Brazil. (Therapsida-Cynodontia). J. Paleontol. 49, 931–939.
Bonaparte, J.F., Barberena, M.C., 2001. On two advanced carnivorous cynodonts from
the Late Triassic of southern Brazil. Bull. Mus. Comp. Zool. 156 (1), 59–80.
Bonaparte, J.F., Sues, H.-D., 2006. A new species of Clevosaurus (Lepidosauria:
Rhynchocephalia) from the upper Triassic of Rio Grande do sul, Brazil.
Palaeontology 49 (4), 917–923.
Bonaparte, J.F., Ferigolo, J., Ribeiro, A.M., 1999. A new early late Triassic saurischian
dinosaur from Rio Grande do sul state, Brazil. Proceedings of the second gondwanan
dinosaur Symposium. Natl. Sci. Mus. Monogr. 15, 89–109.
Bonaparte, J.F., Ferigolo, J., Ribeiro, A.M., 2001. A primitive late Triassic ‘ictidosaur’
from Rio Grande do sul, Brazil. Palaeontology 44 (4), 623–635.
Bonaparte, J.F., Martinelli, A.G., Schultz, C.L., 2005. New information on Brasilodon and
Brasilitherium (Cynodontia, Probainognathia) from the late Triassic of southern
Brazil. Rev. Bras. Palaontol. 8, 25–46.
Bonaparte, J.F., Soares, M.B., Schultz, C.L., 2006. A new non-mammalian cynodont from
the Middle Triassic of southern Brazil and its implications for the ancestry of
mammals. Bull. N. M.Mus. Nat. Hist. Sci. 37, 599–607.
Bonaparte, J.F., Soares, M.B., Martinelli, A.G., 2012. Discoveries in the Late Triassic of
Brazil improve knowledge on the origin of mammals. Historia Natural, Fundación
Felix de Azara, Tercera Serie 2, 5–30.
Bonaparte, J.F., Martinelli, A.G., Schultz, C.L., Rubert, R., 2003. The sister group of
mammals: small cynodonts from the Late Triassic of southern Brazil. Rev. Bras.
Palaontol. 5, 5–27.
Bonaparte, J.F., Brea, G., Schultz, C.L., Martinelli, A.G., 2007. A new specimen of
Guaibasaurus candelariensis (basal Saurischia) from the late Triassic Caturrita
Formation of southern Brazil. Hist. Biol. 19 (1), 73–82.
Bonaparte, J.F., Schultz, C.L., Soares, M.B., Martinelli, A.G., 2010a. La fauna local de
Faxinal do Soturno, Triásico tardío de Rio Grande do Sul, Brasil. Rev. Bras.
Palaontol. 13 (3), 233–246.
Bonaparte, J.F., Schultz, C.L., Soares, M.B., 2010b. Pterosauria from the late Triassic of
southern Brazil. In: Bandyopadhyay, S. (Ed.), New Aspects of Mesozoic Biodiversity,
Lecture Notes in Earth Sciences, vol. 132, pp. 63–71.
Botha, J., Smith, R.M.H., 2006. Rapid vertebrate recuperation in the Karoo Basin of
South Africa following the end-Permian extinction. J. Afr. Earth Sci. 45, 502–514.
Bronzati, M., Müller, R.T., Langer, M.C., 2019. Skull remains of the dinosaur Saturnalia
tupiniquim (Late Triassic, Brazil): with comments on the early evolution of
sauropodomorph feeding behaviour. PloS One 14 (9), e0221387.
Brusatte, S.L., Butler, R.J., Sulej, T., Niedźwiedzki, G., 2009. The taxonomy and anatomy
of rauisuchian archosaurs from the Late Triassic of Germany and Poland. Acta
Palaeontol. Pol. 54 (2), 221–231.
Cabreira, S.F., Schultz, C.L., Bittencourt, J.S., Soares, M.B., Fortier, D.C., Silva, L.R.,
Langer, M.C., 2011. New stem-sauropodomorph (Dinosauria, Saurischia) from the
Triassic of Brazil. Naturwissenschaften 98 (12), 1035–1040.
Brust C, A, Desojo B, J, Schultz L, C, Paes-Neto D, V, Da-Rosa A, A, 2018. Osteology of the
first skull of Aetosauroides scagliai Casamiquela 1960 (Archosauria: Aetosauria) from
the Upper Triassic of southern Brazil (Hyperodapedon Assemblage Zone) and its
phylogenetic importance. Plos One 13 (8), e0201450.
Cabreira, S.F., Kellner, A.W.A., Dias-da-Silva, S., da Silva, L.R., Bronzati, M., de Almeida
Marsola, J.C., Carrilho, R., 2016. A unique Late Triassic dinosauromorph assemblage
reveals dinosaur ancestral anatomy and diet. Curr. Biol. 26 (22), 3090–3095.
Casamiquela, R.M., 1960. Sobre dos nuevos estagonolepoideo argentinos. Ameghiniana 2
(1), 3–9.
Cisneros, J.C., 2008. Taxonomic status of the reptile genus Procolophon from the
gondwanan Triassic. Palaeontol. Afr. 43, 7–17.
Cisneros, J.C., Schultz, C.L., 2002. Procolophon brasiliensis n. sp., a new procolophonid
reptile from the Lower Triassic of southern Brazil. Neues Jahrbuch Geol. Palaontol.
Monatsh. 641–648, 2002.
Cisneros, J.C., Schultz, C.L., 2003. Soturnia caliodon n. g. n. sp., a procolophonid reptile
from the upper Triassic of Southern Brazil. Neues Jahrbuch Geol. Palaontol. Abhand.
227 (3), 365–380.
Cisneros, J.C., Damiani, R., Schultz, C.L., Da-Rosa, Á.A.S., Schwanke, C., Neto, L.W.,
Aurélio, P.L.P., 2004. A procolophonoid reptile with temporal fenestration from the
Middle Triassic of Brazil. Proc. Roy. Soc. Lond. [Biol] 271 (1547), 1541–1546.
Crompton, A.W., 1955. On some Triassic cynodonts from Tanganyka. Proc. Zool. Soc.
Lond. 125, 617–669.
Colbert, E.H., Price, L.I., White, T.E., 1970. A saurischian dinosaur from the Triassic of
Brazil. Am. Mus. Novit. 2405, 1–39.
Cox, C.B., 1962. Preliminary diagnosis of Ischigualastia, a new genus of dicynodont from
Argentina. Breviora 156, 8–9.
Cox, C.B., 1965. New Triassic dicynodonts from South America, their origins and
relationships. Philos. Trans. R. Soc. Lond. B Biol. Sci. 248, 457–516.
Da-Rosa, Á.A.S., Schwanke, C., Aurélio, P.L.P., Poitevin, M., Neto, L.W., 2005. Sítio Linha
Várzea - uma nova assembléia fossilífera do Triássico Médio do sul do Brasil.
Geociencias 24, 115–129.
Da-Rosa, Á.A.S., Schwanke, C., Cisneros, J.C., Neto, L.W., Aurélio, P.L.P., Poitevin, M.,
2004. “Sítio Cortado” - uma nova assembléia fossilífera para o Triássico Médio do sul
do Brasil. Rev. Bras. Palaontol. 7, 289–300.
21
Journal of South American Earth Sciences 104 (2020) 102846
Da-Rosa, Á.A.S., Piñeiro, G., Dias-da-Silva, S., Cisneros, J.C., Feltrin, F.F., Neto, L.W.,
2009. Bica São Tomé, um novo sítio fossilífero para o Triássico Inferior do sul do
Brasil. Rev. Bras. Palaontol. 12 (67), 76.
Dario, E.M., 2017. Arquitetura de fácies e modelo deposicional dos depósitos fluviais
efêmeros da Formação Sanga do Cabral, Triássico inferior da Bacia do Paraná, na
região central do Rio Grande do Sul. Universidade Federal do Rio Grande do Sul
(UFRGS, p. 59. Unpublished Monography.
Dassie, E.C.G., 2014. Tetrápodes triássicos brasileiros: uma investigação envolvendo
banco de dados e análise de cluster. Universidade de São Paulo (USP). Unpublished
Master thesis.
Day, M.O., Ramezani, J., Bowring, S.A., Sadler, P.M., Erwin, D.H., Abdala, F., Rubidge, B.
S., 2015. When and how did the terrestrial mid-Permian mass extinction occur?
Evidence from the tetrapod record of the Karoo Basin, South Africa. Proc. Roy. Soc.
Lond. B 282, 20150834.
De-Oliveira, T.M., Pinheiro, F.L., Da-Rosa, A.A.S., Dias-da-Silva, S., Kerber, L., 2020.
A new archosauromorph from South America provides insights on the early
diversification of tanystropheids. PloS One 15 (5), e0233216.
Desojo, J.B., Ezcurra, M., Schultz, C.L., 2011. An unusual new archosauriform from the
Middle-Late Triassic of southern Brazil and the monophyly of Doswelliidae. Zool. J.
Linn. Soc. 161, 839–871.
Desojo, J.B., Ezcurra, M.D., Kischlat, E.E., 2012. A new aetosaur genus (Archosauria:
Pseudosuchia) from the early Late Triassic of southern Brazil. Zootaxa 3166, 1–33.
Desojo, J.B., von Baczko, M.B., Rauhut, O.W.M., 2020a. 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. 23 (1), 4.
Desojo, J.B., Fiorelli, L.E., Ezcurra, M.D., Martinelli, A.G., Ramezani, J., Da Rosa, Á.A.S.,
von Baczko, M.B., Trotteyn, M.J., Montefeltro, F.C., Ezpeleta, M., Langer, M.C.,
2020b. The late Triassic Ischigualasto formation at Cerro Las Lajas (La Rioja,
Argentina): fossil tetrapods, high-resolution chronostratigraphy, and faunal
correlations. Sci. Rep.
Dias-da-Silva, S., Schultz, C.L., 2008. Early Triassic postcranial temnospondyl remains
from southern Brazil (Sanga do Cabral Formation, Paraná Basin). Rev. Bras.
Palaontol. 11, 51–58.
Dias-da-Silva, S., Da-Rosa, Á.A.S., 2011. Granja Palmeiras, a new fossiliferous site for the
Lower Triassic of southern Brazil. Rev. Bras. Palaontol. 14, 157–168.
Dias-da-Silva, S., Marsicano, C., Schultz, C.L., 2005. Early Triassic temnospondyl skull
fragments from southern south America (Paraná Basin, Brazil). Rev. Bras. Palaontol.
8, 165–172.
Dias-da-Silva, S., Marsicano, C., Schultz, C.L., 2006a. Rhytidosteid temnospondyls in
Gondwana, a new taxon from the lower Triassic of Brazil. Palaeontology 49,
381–390.
Dias-da-Silva, S., Modesto, S.P., Schultz, C.L., 2006b. New material of Procolophon
(Parareptilia, Procolophonoidea) from the lower Triassic of Brazil, with remarks on
the ages of the Sanga do cabral and Buena Vista formations of south America. Can. J.
Earth Sci. 43, 1695–1693.
Dias-da-Silva, S., Dias, E.V., Schultz, C.L., 2009. First record of stereospondyls
(Tetrapoda, Temnospondyli) in the upper Triassic of southern Brazil. Gondwana Res.
15 (1), 131–136.
Dias-da-Silva, S., Cabreira, S.F., Da Silva, L.R., 2011. Occurrence of giant stereospondyl
remains in the Santa Maria Formation (Middle/Upper Triassic of southern Brazil).
Alcheringa 35 (1), 11–19.
Dias-da-Silva, S., Sengupta, D.P., Cabreira, S.F., Da Silva, L.R., 2012. The presence of
Compsocerops (Brachyopoidea: Chigutisauridae) (Late Triassic) in southern Brazil
with comments on chigutisaurid palaeobiogeography. Palaeontology 55 (1),
163–172.
Dias-da-Silva, S., Pinheiro, F., Da-Rosa, Á.A.S., Martinelli, A.G., Schultz, C.L., Silva-
Neves, E., Modesto, S.P., 2017. Biostratigraphic reappraisal of the Lower Triassic
Sanga do Cabral Supersequence from South America, with a description of new
material attributable to the parareptile genus Procolophon. J. South Am. Earth Sci.
79, 281–296.
Dornelles, J.E.F., 1990. Registro sobre a ocorrência de dentes de um arcossaurio para a
Formacão Caturrita, Triássico superior do Rio Grande do Sul. Cienc. Nat. 12, 99–101.
Dornelles, J.E.F., 1995. Um tecodonte proterosuchídeo (Chanaresuchus sp.) do Triássico
do Rio Grande do Sul. Comunicações do Museu de Ciências e Tecnologia UBEA/
PUCRS (Série Ciências da Terra) 1, 63–68.
Eltink, E., Da-Rosa, Á.A.S., Dias-da-Silva, S., 2016. A capitosauroid from the lower
Triassic of south America (Sanga do cabral Supersequence: Paraná Basin), its
phylogenetic relationships and biostratigraphic implications. Hist. Biol. 29,
863–874. https://doi.org/10.1080/08912963.2016.1255736.
Ernesto, M., Demarco, P.N., Xavier, P.A., Sanchez, L.B., Schultz, C.L., Piñeiro, G., 2020.
Age constraints on the Paleozoic Yaguarí-Buena Vista succession from Uruguay:
paleomagnetic and paleontologic information. J. South Am. Earth Sci. 98, 102489.
https://doi.org/10.1016/j.jsames.2019.102489.
Erwin, D.H., 1994. The Permo-Triassic extinction. Nature 367, 231–236.
Ezcurra, M.D., 2017. A new early Coelophysoid neotheropod from the late Triassic of
northwestern Argentina. Ameghiniana 54, 506–538.
Ezcurra, M.D., Desojo, J.B., Rauhut, O.W.M., 2015. Redescription and phylogenetic
relationships of the proterochampsid Rhadinosuchus gracilis (Diapsida:
Archosauriformes) from the early late Triassic of southern Brazil. Ameghiniana 52,
391–417.
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.
C.L. Schultz et al.
Faccini F, U, 1989. O Permo–Triássico do Rio Grande do Sul. Uma análise sob o ponto de
vista das seqüências deposicionais. MSc Dissertation, Universidade Federal do Rio
Grande do Sul, Porto Alegre 1–133.
Ferigolo, J., 2000. Esfenodontídeos do Neo-triássico/?Jurássico do Estado do Rio Grande
do sul. In: Holz, M., De Ros, L.F. (Eds.), Paleontologia Do Rio Grande Do Sul. CIGO/
UFRGS, Porto Alegre, pp. 236–245.
Ferigolo, J., Langer, M.C., 2007. A Late Triassic dinosauriform from south Brazil and the
origin of the ornithischian predentary bone. Hist. Biol. 19 (1), 23–33.
Fisher, D.C., 1981. Crocodilian scatology, microvertebrate concentrations, and enamel-
less teeth. Paleobiology 7, 262–275.
Flynn, J.J., Parrish, J.M., Rakotosamimanana, B., Simpson, W.F., Whatley, R.L., Wyss, A.
R., 1999. A Triassic fauna from Madagascar, including early dinosaurs. Science 286,
763–765. https://doi.org/10.1126/science.286.5440.763.
Flynn, J.J., Parrish, J.M., Rakotosamimanana, B., Ranivoharimanana, L., Simpson, W.F.,
Wyss, A.R., 2000. New traversodontids (Synapsida: Eucynodontia) from the Triassic
of Madagascar. J. Vertebr. Paleontol. 20, 422–427. https://doi.org/10.1671/0272-
4634(2000)020[0422:NTSEFT]2.0.CO, 2.
Fonseca, M.M., Scherer, C.M.S., 1998. The Meso and late Triassic of Southbrazilian
Gondwanaland: a process-oriented analysis and the fluvial deposits. In:
Epicontinental Triassic International Symposium, vol. 1. Halle, pp. 51–52, 1998.
Abstracts.
França A, M, Bittencourt, J, Langer C, M, 2013. Reavaliação taxonômica de
Barberenasuchus brasiliensis (Archosauriformes), Ladiniano do Rio Grande do Sul
(Zona-Assembléia de Dinodontosaurus). Boletim de Destaque, Sociedade Brasileira de
Paleontologia 2013, 230-230.
França, M.A., Ferigolo, J., Langer, M.C., 2011. Associated skeletons of a new middle
Triassic "Rauisuchia" from Brazil. Naturwissenschaften 98 (5), 389–395.
Galton, P.M., 2000. Are Spondylosoma and Staurikosaurus (Santa Maria Formation,
middle-upper Triassic, Brazil) the oldest saurischian dinosaurs? Paläontol. Z. 74 (3),
393–423.
Garcia, M.S., Müller, R.T., Da-Rosa, Á.A., Dias-da-Silva, S., 2019. 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.
Gradstein M, F, Ogg G, J, Schmitz D, M, Ogg, G, 2012. The Geologic Time Scale, 1st.
Elsevier.
Hanich, D., Bertoni, R.S., Abdala, F., Ribeiro, A.M., 2013. Traversodontidae da Zona
Assembleia de Dinodontosaurus (Triassico Medio), Dona Francisca, RS, Brasil. In:
XXIII Congresso Brasileiro de Paleontologia, 2013. Boletim de Resumos XXIII
Congresso Brasileiro de Paleontologia, Gramado, pp. 234–235, 2013.
Horn, B.L.D., Pereira, V., Schultz, C.L., 2013. Calcretes of the Santa Maria
Supersequence, middle Triassic, Rio Grande do sul, Brazil: classification, genesis and
paleoclimatic implications. Palaeogeogr. Palaeoclimatol. Palaeoecol. 376, 39–47.
https://doi.org/10.1016/j.palaeo.2013.02.013.
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.
Horn, B.L.D., Goldberg, K., Schultz, C.L., 2018a. Interpretation of massive sandstones in
ephemeral fluvial settings: a case study from the upper Candelária sequence (upper
Triassic, Paraná Basin, Brazil). J. South Am. Earth Sci. 81, 108–121.
Horn, B.L.D., Goldberg, K., Schultz, C.L., 2018b. A loess deposit in the Late Triassic of
southern Gondwana, and its significance to global paleoclimate. J. South Am. Earth
Sci. 81, 189–203.
Horn, B, Schultz L, C, Figueiredo E, A, Motta, F, 2015. Recognition of the Hyperodapedon
Assemblage Zone (Late Triassic) in a relictual occurrence over the Sul-Rio-Grandense
Shield. Revista Brasileira de Paleontologia 18, 91–96.
Hsiou, A.S., de França, M.A.G., Ferigolo, J., 2015. New data on the Clevosaurus
(Sphenodontia: Clevosauridae) from the upper Triassic of southern Brazil. PloS One
10 (9), e0137523.
Hsiou, A.S., Nydam, R., Simões, T., Pretto, F., Onary, S., Martinelli, A.G., Liparini, A.,
Romo de Vivar Martínez, P., Soares, M.B., Schultz, C.L., Caldwell, M., 2019. A new
clevosaurid from the Triassic (Carnian) of Brazil and the rise of sphenodontians in
Gondwana. Sci. Rep. 9, 11821. https://doi.org/10.1038/s41598-019-48297-9.
Huene, F.v., 1928. Ein Cynodontier aus der Trias Brasiliens. Centralblatt für Mineralogie,
Geologie und Paläontologie B 1928, 251–270.
Huene, F.v., 1935. Lieferung 1. Anomodontia. Die Fossilen Reptilien des
Südamerikanischen Gondwanalandes. Ergebnisse der Sauriergrabungen in
Südbrasilien 1928/29. C. H. Beck’sche Verlagsbuchhandlung, München, pp. 1–82.
Huene, F.v., 1936. Lieferung 2. Cynodontia. Die Fossilen Reptilien des
Südamerikanischen Gondwanalandes. Ergebnisse der Sauriergrabungen in
Südbrasilien 1928/29. C. H. Beck’sche Verlagsbuchhandlung, München, pp. 83–160.
Huene, F.v., 1938a. Stenaulorhynchus, ein Rhynchosauride der ostafrikanischer Obertrias.
Nova Acta Leopoldina, Neue Folge 6, 83–121.
Huene, F.v., 1938b. Die fossilen Reptilien des südamerikanischen Gondwanalandes.
Neues Jahrb Geol Palaontol Abh Referate 3, 142–151.
Huene, F.v., 1942. Lieferungen 3/4. Pseudosuchia, Saurischia, Rhynchosauridae und
Schlussabschnitt. Die Fossilen Reptilien des Südamerikanischen Gondwanalandes.
Ergebnisse der Sauriergrabungen in Südbrasilien 1928/29. C. H. Beck’sche
Verlagsbuchhandlung, München, pp. 161–332.
Irmis B, R, Nesbitt J, S, Sues -D, H, 2013. Early Crocodylomorpha. Geological Society,
London, Special Publications, pp. 275–302, 379.
Kammerer, C.F., Flynn, J.J., Ranivoharimanana, L., Wyss, A.R., 2010. The first record of
a probainognathian (Cynodontia: Chiniquodontidae) from the Triassic of
Madagascar. J. Vertebr. Paleontol. 30 (6), 1889–1894.
22
Journal of South American Earth Sciences 104 (2020) 102846
Kemp, T.S., 1980. Aspects of the structure and functional anatomy of the Middle Triassic
cynodont Luangwa. J. Zool. 191, 193–239.
Kent V, D, Malnis S, P, Colombi E, C, Alcober A, O, Martínez N, R, 2014. Age constraints
on the dispersal of dinosaurs in the Late Triassic from magnetochronology of the Los
Colorados Formation (Argentina). Proceedings of the National Academy of Sciences
111, 7958–7963.
Kischlat, E.E., 2000. Tecodôncios: a aurora dos arcossáurios no Triássico. In: Holz, M., De
Ros, L.F. (Eds.). Paleontologia do Rio Grande do Sul, pp. 273–316, 2000.
Kischlat, E.E., Lucas, S.G., 2003. Phytosaur from the upper Triassic of Brazil. J. Vertebr.
Paleontol. 23 (2), 464–467.
Lacerda, M.B., Schultz, C.L., Bertoni-Machado, C., 2015. First “rauisuchian” archosaur
(Pseudosuchia, Loricata) for the middle Triassic Santacruzodon assemblage zone
(Santa Maria Supersequence), Rio Grande do sul state, Brazil. PloS One 10,
e0118563. https://doi.org/10.1371/journal.pone.0118563.
Lacerda, M.B., França, M.A., Schultz, C.L., 2018. A new erpetosuchid (Pseudosuchia,
Archosauria) from the middle–late Triassic of southern Brazil. Zool. J. Linn. Soc. 184
(3), 804–824.
Lacerda, M., Mastrantonio, B.M., Fortier, D.C., Schultz, C.L., 2016. New insights on
Prestosuchus chiniquensis Huene, 1942 (Pseudosuchia, Loricata) based on new
specimens from the “tree Sanga” outcrop, Chiniquá region, Rio Grande do sul, Brazil.
PeerJ 4, e1622.
Langer, M.C., 2004. Basal saurischians. In: Weishampel, D., Dodson, P., Osmólska, H.
(Eds.), The Dinosauria, second ed. University of California Press, Berkeley,
pp. 25–46. 2004.
Langer, M.C., 2005. Studies on continental Late Triassic tetrapod biochronology. I. The
type locality of Saturnalia tupiniquim and the faunal succession in south Brazil.
J. South Am. Earth Sci. 19 (2), 205–218.
Langer, M.C., Lavina, E.L., 2000. Os amniotas do Neopermiano e Eotriássico da Bacia do
Paraná–répteis e “répteis mamaliformes”. In: Holz, M., de Ros, L.F. (Eds.),
Paleontologia Do Rio Grande Do Sul, pp. 210–235.
Langer, M.C., Ferigolo, J., 2013. The late Triassic dinosauromorph Sacisaurus agudoensis
(Caturrita Formation; Rio Grande do sul, Brazil): anatomy and affinities. Geol. Soc.
Lond. Spec. Publ. 379, 353–392.
Langer, M.C., Bittencourt, J.S., Schultz, C.L., 2007a. The inclusivity and phylogenetic
position of Guaibasaurus candelariensis: a basal dinosaur from the Late Triassic of
Brazil. J. Vertebr. Paleontol. 27 (3), 103A–104A.
Langer, M.C., Bittencourt, J.S., Schultz, C.L., 2010. A reassessment of the basal dinosaur
Guaibasaurus candelariensis, from the Late Triassic Caturrita Formation of south
Brazil. Earth Environ. Sci. Trans. Roy. Soc. 101, 301–332.
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.
Langer, M.C., Abdala, F., Richter, M., Benton, M.J., 1999. A sauropodomorph dinosaur
from the Upper Triassic (Carman) of southern Brazil. C. R. Acad. Sci. 329 (7),
511–517.
Langer, M.C., Ribeiro, A.A.M., Schultz, C.L., Ferigolo, J., 2007b. The continental
tetrapod-bearing Triassic of south Brazil. Bull. N. M.Mus. Nat. Hist. Sci. 41, 201–218.
Lavina, E.L., 1983. Procolophon pricei sp. n. um novo réptil procolofonídeo do Triássico
do Rio Grande do Sul. Iheringia Ser. Zool. 9, 51–78.
Leal, L.A., Azevedo, S.A.K., Kellner, A.W.A., Da-Rosa, Á.A.S., 2004. A new early dinosaur
(Sauropodomorpha) from the Caturrita Formation (late Triassic), Parana basin,
Brazil. Zootaxa 690, 1–24.
Liu, J., Olsen, P., 2010. The phylogenetic relationships of Eucynodontia (Amniota:
Synapsida). J. Mamm. Evol. 17, 151–176.
Liu, J., Abdala, F., 2014. Phylogeny and taxonomy of the traversodontidae. In:
Kammerer, C.F., Angielczyk, K.D., Fröbisch, J. (Eds.), Early Evolutionary History of
the Synapsida. Springer, pp. 255–279.
Liu, J., Soares, M.B., Reichel, M., 2008. Massetognathus (Cynodontia, traversodontidae)
from the Santa Maria Formation of Brazil. Rev. Bras. Palaontol. 11 (1), 27–36.
Lucas, S.G., 1993. Barysoma lenzii (Synapsida: Dicynodontia) from the middle Triassic of
Brazil, a synonym of Stahleckeria potens. J. Paleontol. 67, 318–321.
Lucas, S.G., 2001. Age and correlation of Triassic tetrapod assemblages from Brazil.
Albertiana 26, 13–20. https://doi.org/10.1016/S0031-0182(98)00117-5.
Lucas G, S, Harris K, S, 1996. Taxonomic and biochronological significance of specimens
of the Triassic dicynodont Dinodontosaurus Romer 1943 in the Tübingen collection.
Paläontologische Zeitschrift 70, 603–622.
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. Unit. States Am. 113
(3), 509–513.
Marsola, J.C., Bittencourt, J.S., Butler, R.J., Da-Rosa, Á.A., Sayão, J.M., Langer, M.C.,
2019. A new dinosaur with theropod affinities from the late Triassic Santa Maria
Formation, south Brazil. J. Vertebr. Paleontol. 38 (5), e1531878.
Marsola CA, J, Bittencourt S, J, Da-Rosa A, A, Martinelli G, A, Ribeiro M, A, Ferigolo, J,
Langer C, M, 2018. New sauropodomorph and cynodont remains from the Late
Triassic Sacisaurus site in southern Brazil and its stratigraphic position in the Norian
Caturrita Formation. Acta Palaeontologica Polonica 63, 653–669.
Martinelli, A.G., 2017. Contribuição ao conhecimento dos cinodontes probainognátios
(Therapsida, Cynodontia, Probainognathia) do Triássico da América do Sul e seu
impacto na origem dos Mammaliaformes. Universidade Federal do Rio Grande do
Sul, Brazil, p. 645. PhD Thesis.
Martinelli, A.G., Rougier, G.W., 2007. On Chaliminia musteloides Bonaparte (Cynodontia,
Tritheledontidae) and the phylogeny of the Ictidosauria. J. Vertebr. Paleontol. 27,
442–460.
Martinelli, A.G., Bonaparte, J.F., 2011. Postcanine replacement in Brasilodon and
Brasilitherium (Cynodontia, Probainognathia) and its bearing in cynodont evolution.
In: Calvo, J., Porfiri, J., González Riga, B., dos Santos, D. (Eds.), Dinosaurios y
Paleontología desde América Latina, Anales del III Congreso Latinoamericano de
C.L. Schultz et al.
Paleontología, Neuquén, 2008. Editorial de la Universidad Nacional de Cuyo,
Mendoza, pp. 179–186.
Martinelli, A.G., Soares, M.B., 2016. Evolution of south American non-mammaliaform
cynodonts (Therapsida, Cynodontia). In: Agnolin, F.L., Lio, G.L., Egli, F.B.,
Chimento, N.R., Novas, F.E. (Eds.), Contribuciones del MACN: Historia Evolutiva y
Paleobiogeográfica de los Vertebrados de América del Sur, pp. 183–196. Buenos
Aires.
Martinelli, A.G., Soares, M.B., Schwanke, C., 2016. Two new cynodonts (Therapsida)
from the middle-early late Triassic of Brazil and comments on south American
probainognathians. PloS One 11, e0162945.
Martinelli, A.G., Bonaparte, J.F., Schultz, C.L., Rubert, R., 2005. A new tritheledontid
(Therapsida, Eucynodontia) from the Late Triassic of Rio Grande do Sul (Brazil), and
its phylogenetic relationships among carnivorous non-mammlain eucynodonts.
Ameghiniana 42 (1), 191–208.
Martinelli, A.G., Kammerer, C.F., Melo, T.P., Paes-Neto, V.D., Ribeiro, A.M., Da-Rosa, A.
A.S., Schultz, C.L., Soares, M.B., 2017a. The African cynodont Aleodon (Cynodontia,
Probainognathia) in the Triassic of southern Brazil and its biostratigraphic
significance. PloS One 12 (6), e0177948.
Martinelli, A.G., Soares, M.B., Oliveira, T.V., Rodrigues, P.G., Schultz, C.L., 2017b. The
Triassic eucynodont Candelariodon barberenai revisited and the early diversity of
stem prozostrodontians. Acta Palaeontol. Pol. 62 (3), 527–542.
Martinelli, A.G., Eltink, E., Da-Rosa, Á.A., Langer, M.C., 2017c. A new cynodont from the
Santa Maria formation, south Brazil, improves Late Triassic probainognathian
diversity. Pap. Palaeontol. 3 (3), 401–423.
Martinelli, A.G., Corfe, I.J., Gill, P.G., Kallonen, A., Rayfield, E.J., Rodrigues, P.B.,
Schultz, C.L., Soares, M.B., 2017d. Brasilodon quadrangularis, Brasilitherium
riograndensis and Minicynodon maieri (Cynodontia): taxonomy, ontogeny and tooth
replacement. In: Congresso Brasileiro de Paleontologia, vol. 25. Ribeirão Preto,
p. 189.
Martínez, R.N., Forster, C.A., 1996. The skull of Probelesodon sanjuanensis sp. nov., from
the late Triassic Ischigualasto formation of Argentina. J. Vertebr. Paleontol. 16 (2),
285–291.
Martínez, 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 (6014), 206–210.
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. https://doi.org/10.1080/
02724634.2013.818546.
Mastrantonio, B., von Baczko, B.M., Desojo, J., Schultz, C.L., 2019. The skull anatomy
and cranial endocast of the pseudosuchid archosaur Prestosuchus chiniquensis from
the Triassic of Brazil. Acta Palaeontol. Pol. https://doi.org/10.4202/
app.00527.2018.
Mattar, L.C.B., 1987. Descrição osteólogica do crânio e segunda vértebrata cervical de
Barberenasuchus brasiliensis Mattar, 1987 (Reptilia, Thecodontia) do Mesotriássico do
Rio Grande do Sul, Brasil. An. Acad. Brasil. Cienc. 61, 319–333.
Melo, T.P., Abdala, F., Soares, M.B., 2015. The Malagasy cynodont Menadon besairiei
(Cynodontia; traversodontidae) in the middle–upper Triassic of Brazil. J. Vertebr.
Paleontol. 35, e1002562 https://doi.org/10.1080/02724634.2014.1002562.
Melo, T.P., Martinelli, A.G., Soares, M.B., 2017. A new gomphodont cynodont
(traversodontidae) from the middle-late Triassic Dinodontosaurus assemblage zone of
the Santa Maria Supersequence, Brazil. Palaeontology 1–12. https://doi.org/
10.1111/pala.12302.
Melo, T.P., Ribeiro, A.M., Martinelli, A.G., Soares, M.B., 2019. Early evidence of
molariform hypsodonty in a Triassic stem-mammal. Nat. Commun. 10, 2841.
https://doi.org/10.1038/s41467-019-10719-7.
Milani, E.J., 1997. Evolução tectono-estratigráfica da Bacia do Paraná e seu
relacionamento com a geodinâmica fanerozóica do Gondwana Sul-ocidental.
Universidade Federal do Rio Grande do Sul. PhD Thesis.
Milani, E.J., Faccini, U.F., Scherer, C.M., Araújo, L.M., Cupertino, J.A., 1998. Sequences
and Stratigraphic Hierarchy of the Paraná Basin (Ordovician to Cretaceous),
Southern Brazil, vol. 29. Boletim do Instituto de Geociências - USP, pp. 125–173.
Série Científica.
Miron, L.R., Pavanatto, A.E.B., Pretto, F.A., Müller, R.T., Dias-da-Silva, S., Kerber, L.,
2020. Siriusgnathus niemeyerorum (Eucynodontia: Gomphodontia): the youngest
south American traversodontid? J. South Am. Earth Sci. 97, 102394.
Müller, R.T., Garcia, M.S., 2019. Rise of an empire: analysing the high diversity of the
earliest sauropodomorph dinosaurs through distinct hypotheses. Hist. Biol. 1–6.
Müller, R.T., von Baczko, M.B., Desojo, J.B., Nesbitt, S.J., 2020. The first ornithosuchid
from Brazil and its macroevolutionary and phylogenetic implications for Late
Triassic faunas in Gondwana. Acta Palaeontol. Pol. (in press).
Müller, R.T., Langer, M.C., Bronzati, M., Pacheco, C.P., Cabreira, S.F., Dias-Da-Silva, S.,
2018a. Early evolution of sauropodomorphs: anatomy and phylogenetic
relationships of a remarkably well-preserved dinosaur from the Upper Triassic of
southern Brazil. Zool. J. Linn. Soc. 184 (4), 1187–1248.
Montefeltro C, F, Langer C, M, Schultz L, C, 2010. Cranial anatomy of a new genus of
hyperodapedontine rhynchosaur (Diapsida, Archosauromorpha) from the Upper
Triassic of southern Brazil. Earth Environ. Sci. Trans. Roy. Soc. Edinburgh 101,
27–52.
Müller, R.T., Da-Rosa, A.A.S., Silva, L.R., Aires, A.S.S., Pacheco, C.P., Pavanatto, A.E.B.,
Dias-da-Silva, S., 2015. Wachholz, a new exquisite dinosaur-bearing fossiliferous site
from the Upper Triassic of southern Brazil. J. South Am. Earth Sci. 61, 120–128.
Müller, R.T., Langer, M.C., Silva, S.D., 2018b. An exceptionally preserved association of
complete dinosaur skeletons reveals the oldest long-necked sauropodomorphs. Biol.
Lett. 14 (11), 1744–9561. https://doi.org/10.1098/rsbl.2018.0633.
23
Journal of South American Earth Sciences 104 (2020) 102846
Nesbitt, S.J., Butler, R.J., 2012. Redescription of the archosaur Parringtonia gracilis from
the middle Triassic Manda beds of Tanzania, and the antiquity of Erpetosuchidae.
Geol. Mag. 150 (2), 225–238.
Nesbitt, S.J., Flynn, J.J., Pritchard, A.C., Parrish, J.M., Ranivoharimanana, L., Wyss, A.R.,
2015. Postcranial osteology of Azendohsaurus madagaskarensis (?Middle to upper
Triassic, Isalo group, Madagascar) and its systematic position among stem archosaur
Reptiles. Bull. Am. Mus. Nat. Hist. 398, 1–126. https://doi.org/10.1206/amnb-899-
00-1-126.1.
Nesbitt, S., Desojo, J.B., 2017. The osteology and phylogenetic position of Luperosuchus
fractus (Archosauria: Loricata) from the latest middle Triassic or earliest late Triassic
of Argentina. Ameghiniana 54 (3), 261–282.
Novas, F.E., Ezcurra, M.D., Chatterjee, S., Kutty, T.S., 2010. New dinosaur species from
the upper Triassic upper Maleri and lower Dharmaram formations of central India.
Earth Environ. Sci. Trans. Roy. Soc. 101 (3–4), 333–349.
Oliveira V, T, Schultz L, C, 2007. La predominancia de Exaeretodon Cabrera 1943 en una
sección Triásica de Brasil y su probable correlación conel mismo evento en la porción
mediana superior de la Formación Ischigualasto (Triásico de Argentina). XXIII
Jornadas Argentinas de Paleontología de Vertebrados – Programa de
Comunicaciones Científicas y Libro de Resúmenes, Trelew, Argentina, 9-9.
Oliveira, T.V., Soares, M.B., Schultz, C.L., 2010. Trucidocynodon riograndensis gen. nov. et
sp. nov. (Eucynodontia), a new cynodont from the Brazilian Upper Triassic (Santa
Maria Formation). Zootaxa 2382 (1), 1–71.
Oliveira, T.V., Martinelli, A.G., Soares, M.B., 2011a. New material of Irajatherium
hernandezi Martinelli, Bonaparte, schultz & Rubert 2005 (Eucynodontia,
Tritheledontidae) from the upper Triassic (Caturrita Formation, Paraná Basin) of
Brazil. Paläontol. Z. 85, 67–82.
Oliveira, T.V., Soares, M.B., Schultz, C.L., Rodrigues, P., 2011b. A new carnivorous
cynodont (Synapsida, Therapsida) from the Brazilian Middle Triassic (Santa Maria
Formation): Candelariodon barberenai gen. et sp. nov. Zootaxa 3027, 19–28.
Oliveira, T.M., Oliveira, D., Schultz, C.L., Kerber, L., Pinheiro, F.L., 2018. Tanystropheid
archosauromorphs in the lower Triassic of Gondwana. Acta Palaeontol. Pol. 63 (4),
713–723.
Pacheco, C.P., Martinelli, A.G., Pavanatto, A.E.B., Soares, M.B., Dias-da-Silva, S., 2018.
Prozostrodon brasiliensis, a probainognathian cynodont from the Late Triassic of
Brazil: second record and improvements on its dental anatomy. Hist. Biol. 30 (4),
475–485.
Pacheco, C., Müller, R.T., Langer, M., Pretto, F.A., Kerber, L., da Silva, S.D., 2019.
Gnathovorax cabreirai: a new early dinosaur and the origin and initial radiation of
predatory dinosaurs. PeerJ 7, e7963.
Pavanatto E, A, Müller T, R, Da-Rosa AS, A, Dias-da-Silva, S, 2016. New information on
the postcranial skeleton of Massetognathus ochagaviae Barberena, 1981
(Eucynodontia, Traversodontidae), from the Middle Triassic of Southern Brazil. Hist.
Biol. 28, 978–989.
Pavanatto, A.E.B., Pretto, F.A., Kerber, L., Müller, R.T., Da-Rosa, Á.A.S., Dias-da-Silva, S.,
2018. A new Upper Triassic cynodont-bearing fossiliferous site from southern Brazil,
with taphonomic remarks and description of a new traversodontid taxon. J. South
Am. Earth Sci. 88, 179–196.
Pavanatto, A.E.B., Da-Rosa, Á.A.S., Temp-Müller, R., Roberto-da-Silva, L., Ribeiro, A.M.,
Martinelli, A.G., Dias-da-Silva, S., 2020. Bortolin site, a new fossiliferous locality in
the Triassic (Ladinian/Carnian) of southern Brazil. Rev. Bras. Palaontol. 23 (2),
123–137.
Peecook R, B, Steyer S, J, Tabor J, N, Smith M, R, 2017. Updated geology and vertebrate
paleontology of the Triassic Ntawere Formation of northeastern Zambia, with special
emphasis on the archosauromorphs. J. Vertebr. Paleontol. Supl1, 8–38.
Peruzzo, C.S., Araújo-Barberena, D.C., 1995. Sobre a ocorrência do gênero Ischigualastia
Cox, 1962 na Formação Santa Maria, triássico do Rio Grande do sul. An. Acad. Brasil.
Cienc. 67, 175–181.
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.
Piñeiro, G., Verde, M., Ubilla, M., Ferigolo, J., 2003. First basal synapsids (“pelycosaurs”)
from south America, late Permian? Early Triassic of Uruguay. J. Paleontol. 77,
389–392.
Piñeiro, G., Marsicano, C., Lorenzo, N., 2007. A new temnospondyl from the Permo-
Triassic Buena Vista formation of Uruguay. Palaeontology 50, 627–640.
Pinheiro, F.L., 2016. A fragmentary dinosaur femur and the presence of Neotheropoda in
the Upper Triassic of Brazil. Rev. Bras. Palaontol. 19 (2), 211–216.
Pinheiro, F.L., França, M.A.G., Lacerda, M.B., Butler, R.J., Schultz, C.L., 2016. An
exceptional fossil skull from South America and the origins of the archosauriform
radiation. Sci. Rep. 6, 22817.
Pinheiro, F.L., Simão-Oliveira, D., Butler, R.J., 2019. Osteology of the archosauromorph
Teyujagua paradoxa and the early evolution of the archosauriform skull. Zool. J.
Linn. Soc. 20, 1–40.
Pretto, F.A., Langer, M.C., Schultz, C.L., 2019. A new dinosaur (Saurischia:
Sauropodomorpha) from the Late Triassic of Brazil provides insights on the evolution
of sauropodomorph body plan. Zool. J. Linn. Soc. 185 (2), 388–416.
Pretto A, F, Schultz L, C, Langer C, M, 2015. New dinosaur remains from the Late Triassic
of southern Brazil (Candelária Sequence, Hyperodapedon Assemblage Zone).
Alcheringa 39, 1–10.
Pretto A, F, Veiga H, F, Langer C, M, Schultz L, C, 2017. A juvenile sauropodomorph tibia
from the ‘Botucaraí Hill’, Late Triassic of Southern Brazil. Revista Brasileira de
Paleontologia 19, 407–414.
Price, L.I., 1946. Sobre um novo peudosuquio do Triássico Superior do Rio Grande do
Sul. Departmento Nacional da Produção Mineral 120, 1–38.
C.L. Schultz et al.
Price, L.I., 1947. Um procolofonídeo do triássico do Rio Grande do sul. Boletim da Divisão
de Geologia e Paleontologia, DNPM 122, 7–27.
Queiroz, L.C., Pereira, V.P., Soares, M.B., Schultz, C.L., 2020. Geochemical study of the
vertebrate assemblage zones of the Santa Maria Supersequence (middle to late
Triassic), Paraná Basin, Brazil. Brazilian J. Geol. submitted for publication.
Raugust, T., Lacerda, M., Schultz, C.L., 2013. The first occurrence of Chanaresuchus
bonapartei Romer 1971 (Archosauriformes, Proterochampsia) of the middle Triassic
of Brazil from the Santacruzodon assemblage zone, Santa Maria Formation (Parana
basin). Geol. Soc. London, Spec. Publ. 379, 303–318. https://doi.org/10.1144/
SP379.22.
Razafimbelo, E., 1987. Le basin de Morondava (Madagascar). Synthèse géologique
Struct. Université Louis Pasteur Strasbourg, France.
Reichel, M., Schultz, C.L., Pereira, V.P., 2005. Diagenetic pattern of vertebrate fossils
from the traversodontidae biozone, Santa Maria Formation (Triassic), southern
Brazil. Rev. Bras. Palaontol. 8, 173–180.
Reichel, M., Schultz, C.L., Soares, M.B., 2009. A new traversodontid cynodont
(Therapsida, Eucynodontia) from the middle Triassic Santa Maria Formation of Rio
Grande do sul. Palaeontology 52 (1), 229–250.
Retallack J, G, Sheldon D, N, Cogoini, M, Elmore D, R, 2003. Magnetic susceptibility of
early Paleozoic and Precambrian paleosols. Palaeogeogr. Palaeoclimatol. Palaeoecol.
198, 373–380.
Ribeiro, A.M., Abdala, F., Bertoni, R.S., 2011. Traversodontid cynodonts
(Therapsida–Eucynodontia) from two upper Triassic localities of the Paraná Basin,
southern Brazil. Ameghiniana 48 (Supplement), R111.
Roberto-Da-Silva, L., Desojo, J.B., Cabreira, S.F., Aires, A.S., Mueller, R.T., Pacheco, C.P.,
Dias-Da-Silva, S., 2014. A new aetosaur from the upper Triassic of the Santa Maria
Formation, southern Brazil. Zootaxa 3764 (3), 240–278.
Roberto-Da-Silva, L., Müller, R.T., França, M.A.G.D., Cabreira, S.F., Dias-Da-Silva, S.,
2018. An impressive skeleton of the giant top predator Prestosuchus chiniquensis
(Pseudosuchia: Loricata) from the Triassic of Southern Brazil, with phylogenetic
remarks. Hist. Biol. 1–20.
Romer, A.S., 1967. The Chañares (Argentina) Triassic reptile fauna. III. Two new
gomphodonts Massetognathus pascuali and M. teruggii. Breviora 264, 1–25.
Romo de Vivar R, P, Martinelli G, A, Paez Neto D, V, Scartezini, C, Lacerda B, M,
Rodrigues, C, Soares B, M, 2019. New rhynchocephalian specimen in the Late
Triassic of southern Brazil and comments on the palatine bone of Brazilian
rhynchocephalians. Hist. Biol. https://doi.org/10.1080/08912963.2019.1602616.
Romo de Vivar, M.P.R., Soares, M.B., 2015. Dentary morphological variation in
Clevosaurus brasiliensis (Rhynchocephalia, Clevosauridae) from the upper Triassic of
Rio Grande do sul, Brazil. PloS One 10 (3), e0119307.
Romer R, A, Price I, L, 1944. Stahleckeria lenzii, a giant Triassic Brazilian dicynodont.
Bull. Mus. Comp. Zool. 93, 463–491.
Romo de Vivar, P., Martinelli, A.G., Hsiou, A.S., Soares, M.B., 2020a. A new
rhynchocephalian from the Late Triassic of southern Brazil enhances
eusphenodontian diversity. J. Syst. Palaeontol. 18, 1103–1126. https://doi.org/
10.1080/14772019.2020.1732488.
Romo-de-Vivar, P.R., Martinelli, A.G., Fonseca, P.H.M., Soares, M.B., 2020b. To be or not
to be: the hidden side of Cargninia enigmatica and other puzzling remains of
Lepidosauromorpha from the Upper Triassic of Brazil. J. Vertebr. Paleontol.
Rubert, R.R., Schultz, C.L., 2004. Um Novo Horizonte de Correlação para o Triássico
Superior do Rio Grande do Sul. Pesqui. em Geociencias 31 (1), 71–88. https://doi.
org/10.22456/1807-9806.19569.
Sá-Teixeira, A.M., 1987. Novas observações osteológicas e taxonômicas sobre
Massetognathus ochagaviae Barberena, 1981 (Reptilia, Therapsida, Cynodontia).
Paula-Coutiana 1, 39–49.
Sá-Teixeira, A.M., 1995. A família Traversodontidae (Therapsida, Cynodontia) no sul do
Brasil e suas relações com formas afins no domínio gonduânico. Universidade
Federal do Rio Grande do Sul (UFRGS), p. 144. PhD Thesis.
Sahney, S., Benton, M.J., 2008. Recovery from the most profound mass extinction of all
time. Proc. R. Soc. Ser. B 275, 759–765.
Schmitt, M.R., Martinelli, A.G., Melo, T.P., Soares, M.B., 2019. On the occurrence of the
traversodontid Massetognathus ochagaviae (Synapsida, Cynodontia) in the early late
Triassic Santacruzodon assemblage zone (Santa Maria Supersequence, southern
Brazil): taxonomic and biostratigraphic implications. J. South Am. Earth Sci. 93,
36–50.
Schultz, C.L., 1995. Subdivisão do Triássico do RS com base em macrofósseis: problemas
e perspectivas. Comunicações do Museu de Ciências e Tecnologia, UBEA/PUCRS,
Série Ciências da Terra 1, 25–32.
Schultz, C.L., 2005. Biostratigraphy of the non-marine Triassic: is a global correlation
based on tetrapod faunas possible? In: Koutsoukos, E.A.M. (Ed.) Applied
Stratigraphy. 1ed, pp. 123–145 (Org.), 2005.
24
Journal of South American Earth Sciences 104 (2020) 102846
Schultz, C.L., Langer, M.C., Montefeltro, F.C., 2016. A new rhynchosaur from south
Brazil (Santa Maria Formation) and rhynchosaur diversity patterns across the
Middle-Late Triassic boundary. Paläontol. Z. 90, 593–609.
Schultz, C.L., Scherer, C.M., Barberena, M.C., 2000. Bioestratigraphy of southern
Brazilian middle-upper Triassic. Rev. Bras. Geociencias 30, 495–498.
Sengupta, D.P., 1995. Chigutisaurid temnospondyls from the late Triassic of India and a
review of the family Chigutisauridae. Palaeontology 38 (2), 313–339.
Sennikov, A.G., 2011. New tanystropheids (Reptilia: Archosauromorpha) from the
Triassic of Europe. Paleontol. J. 45, 90–104.
Simms, M.J., Ruffell, A.H., 1989. Synchronicity of climatic change and extinctions in the
Late Triassic. Geology 17 (3), 265–268. https://doi.org/10.1130/0091-7613(1989)
017.
Silva-Neves, E., Modesto, S.P., Dias-da-Silva, S., 2018. A new, nearly complete skull of
Procolophon trigoniceps Owen 1876 from the Sanga do Cabral Supersequence, Lower
Triassic of Southern Brazil, with phylogenetic remarks. Hist. Biol. https://doi.org/
10.1080/08912963.2018.1512106.
Smith MH, R, 1995. Changing fluvial environments across the Permian–Triassic
boundary in the Karoo Basin, South Africa and possible causes of tetrapod
extinctions. Palaeogeogr. Palaeoclimatol. Palaeoecol. 117, 81–104.
Soares, M.B., Abdala, F., Bertoni-Machado, C., 2011a. A sectorial toothed cynodont
(Therapsida) from the Triassic Santa Cruz do sul fauna, Santa Maria Formation,
southern Brazil. Geodiversitas 33, 265–278. https://doi.org/10.5252/g2011n2a4.
KEY.
Soares, M.B., Schultz, C.L., Horn, B.L.D., 2011b. New information on Riograndia
guaibensis Bonaparte, Ferigolo & Ribeiro, 2001 (Eucynodontia, Tritheledontidae)
from the late Triassic of southern Brazil: anatomical and biostratigraphic
implications. An. Acad. Brasil. Cienc. 83 (1), 329–354.
Soares, M.B., Dalla Vecchia, F.M., Schultz, C.L., Kellner, A.W.A., 2013. On the supposed
pterosaurian nature of Faxinalipterus minima Bonaparte et al. (2010) from the Upper
Triassic of Rio Grande do Sul, Brazil. In: Sayão, J.M., Costa, F.R., Bantim, R.A.M.,
Kellner, A.W.A. (Eds.), International Symposium on Pterosaurs, Rio Ptero 2013,
Short Communications. Universidade Federal do Rio de Janeiro, pp. 95–97.
Soares, M.B., Martinelli, A.G., Oliveira, T.V., 2014. A new prozostrodontian cynodont
(Therapsida) from the late Triassic Riograndia assemblage zone (Santa Maria
Supersequence) of southern Brazil. An. Acad. Brasil. Cienc. 86, 1673–1691.
Stefanello, M., Müller, R.T., Kerber, L., Martínez, R.N., Dias-da-Silva, S., 2018. Skull
anatomy and phylogenetic assessment of a large specimen of Ecteniniidae
(Eucynodontia: Probainognathia) from the Upper Triassic of southern Brazil.
Zootaxa 4457 (3), 351–378.
Teixeira, A.M.S., 1982. Um novo cinodonte carnivoro (Probelesodon kitchingi sp. nov.) do
Triassico do Rio Grande do Sul, Brasil. Comunicações do Museu de Ciências da
PUCRS 24, 1–31.
Vega-Dias, C., Maisch, M.W., Schwanke, C., 2005. The taxonomic status of Stahleckeria
impotens (Therapsida, Dicynodontia): redescription and discussion of its
phylogenetic position. Rev. Bras. Palaontol. 8, 221–228.
Veevers J, J, Cole I, D, Cowan J, E, 1994. Southern Africa: Karoo basin and Cape Fold
Belt. Geological Society of America, Memoir 184, 223–279.
Vega-Dias, C., Maisch, M.W., Schultz, C.L., 2004. A new phylogenetic analysis of Triassic
dicynodonts (Therapsida) and the systematic position of Jachaleria candelariensis
from the Upper Triassic of Brazil. Neues Jahrbuch Geol. Palaontol. Abhand. 231,
145–166.
von Baczko, M.B., 2018. Rediscovered cranial material of Venaticosuchus rusconii enables
the first jaw biomechanics in Ornithosuchidae (Archosauria: Pseudosuchia).
Ameghiniana 55 (4), 365–380.
Ward D, P, Montgomery R, D, Smith, R, 2000. Altered river morphology in South Africa
related to the Permian-Triassic extinction. Science 289, 1740–1743.
Weinbaum, J.C., 2013. Postcranial skeleton of Postosuchus kirkpatricki (Archosauria:
Paracrocodylomorpha), from the upper Triassic of the United States. Geol. Soc. Spec.
Publ. 379 (1), 525–553.
Zerfass, H, Chemale Jr, F, Schultz L, C, Lavina L, E, 2004. Tectonics and sedimentation in
South America during Triassic. Sediment. Geol. 166, 265–292.
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.
Zingano, A.G., Cauduro, A.D., 1959. Afloramentos Fossilíferos Do Rio Grande Do Sul, vol.
8. Boletim Instituto de Ciências Naturais, Ministério de Educação e Cultura,
Universidade do Rio Grande do Sul, pp. 1–48.