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First record of stereospondyls (Tetrapoda, Temnospondyli) in the Upper

Triassic of Southern Brazil

Article in Gondwana Research · February 2009

DOI: 10.1016/j.gr.2008.07.002

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GR Letter

First record of stereospondyls (Tetrapoda, Temnospondyli) in the Upper Triassic

of Southern Brazil
Sérgio Dias-da-Silva a,⁎, Eliseu Vieira Dias b, Cesar Leandro Schultz c
a
Centro de Ciências Rurais, Universidade Federal do Pampa, Campus de São Gabriel, Avenida Antônio Trilha, 1847, Bairro Centro, CEP 97.300-000, São Gabriel, Rio Grande do Sul, Brasil
b
Centro Universitário Positivo, Núcleo de Ciências Biológicas e da Saúde, Rua Prof. Pedro Viriato Parigot de Souza, 5.300, CEP 81.280-330, Curitiba, Paraná, Brasil
c
Laboratório de Paleovertebrados, Departamento de Paleontologia e Estratigrafia, Instituto de Geociências, Universidade Federal do Rio Grande do Sul, Avenida Bento Gonçalves,
9.500 prédio 43.127/110 B, CEP: 91.540-000, Porto Alegre, Rio Grande do Sul, Brazil

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

Article history: Stereospondyls survived the Permo-Triassic extinctions in a refuge probably located in the landmass that
Received 21 October 2007 nowadays comprises Australia. Subsequently, they radiated to other parts of Pangaea, reaching their highest
Received in revised form 16 July 2008 distribution and diversification during the Early Triassic. An incomplete interclavicle from the Caturrita
Accepted 16 July 2008
Formation represents their first record in the Upper Triassic of Brazil. Previously, Upper Triassic South
Available online 24 July 2008
American stereospondyls were restricted to Argentina. This new record reinforces a former hypothesis that
Keywords:
suggests the presence of a more diverse stereospondyl fauna in South America during the Late Triassic than
Temnospondyli previously assumed. Additionally, the presence of a stereospondyl and a phytosaur in the Caturrita Formation
Western Gondwana reinforces the hypothesis of a change to more humid climatic conditions in the Paraná Basin during the
Paleobiogeography Upper Triassic. The record of Early Triassic stereospondyls in South America suggests that they first colonized
Rosário do Sul Group Brazil and/or Uruguay, spreading from South Africa during the Early Triassic, subsequently reaching
Santa Maria Supersequence Argentina. Up till now, there is no record of Middle Triassic stereospondyls in either Argentina and Brazil,
probably due to either taphonomic bias or insufficient prospecting. Despite the lack of direct evidence, one
should not dismiss an earlier stereospondyl colonization of Argentina still during the Early or Middle Triassic.
© 2008 International Association for Gondwana Research. Published by Elsevier B.V. All rights reserved.

1. Introduction (South America, South Africa, Madagascar, Antarctica, Australia, and

Stereospondyls are derived temnospondyls that invaded Gond-
wana during the Late Permian (Warren et al., 2000). According to
these authors, they probably survived the Permo-Triassic extinctions
in a refuge within the landmass that nowadays comprises Australia.
The sister taxon of Stereospondyli, the archegosaurs, is present in
Northern Gondwana (in Permian deposits from Brazil and India)
(Price, 1948, Cox and Hutchinson, 1991 and Warren et al., 2000). Their
absence from Kazakhstan and Siberia support the hypothesis of a
route across northern Gondwana, still during early Late Permian,
reaching a refuge in eastern Gondwana very close to the Permo-
Triassic boundary (Warren et al., 2000). The presence of a derived
stereospondyl (Trucheosaurus major) in Upper Permian deposits from
Australia supports this hypothesis (Marsicano and Warren, 1998;
Warren et al., 2000). Subsequently, they radiated to other parts of
Gondwana and Laurasia, reaching their highest distribution and
diversification during the Early Triassic. Indeed, all representative
clades of stereospondyls are widespread in deposits from Gondwana
⁎ Corresponding author. Fax: +55 5532326075.
E-mail addresses: sergiosilva@unipampa.edu.br (S. Dias-da-Silva),
eliseu.dias@unicenp.edu.br (E.V. Dias), cesar.schultz@ufrgs.br (C.L. Schultz).
India, see Schoch and Milner, 2000; Yates and Warren, 2000, Warren
and Marsicano, 2000, Damiani, 2001, Sidor et al., 2007). As stated by
Steyer (2002, 2003), the Early Triassic radiation of stereospondyls
could probably have been facilitated by their possible marine re-
adaptation (cf. the marine stereospondyls of Madagascar). However,
taking into account the high level of coalition of Pangaea during the
Triassic, a migration through fresh water bodies is also probable.
Temnospondyl stereospondyls are among the most frequently
encountered tetrapods in Triassic continental deposits. Late Triassic
stereospondyl temnospondyls were only known in South America
from deposits located in central-western Argentina. There are four
described Upper Triassic (Carnian to Norian) Argentinean stereospon-
dyl taxa: the mastodonsaurid Promastodontosaurus bellmanni from the
Ischigualasto Formation (Ischigualasto Basin); the chigutisaurids Pe-
lorocephalus mendonzensis and P. cacheutensis, both from the Cacheuta
Formation (Cuyana Basin) and P. tenax, from the Rio Blanco Formation
(Cuyana Basin). In addition, an undetermined brachyopoid is recorded
in the Cacheuta Formation (see Bonaparte, 1963; Marsicano, 1999,
2005). In contrast, the temnospondyl record in Brazil and Uruguay is a
mixture of non-stereospondyl temnospondyls, basal stereospondyls
and advanced stereospondyls restricted to Upper Permian–Lower
Triassic levels. From the Upper Permian, there are two long snouted
archegosaurids already described, Prionosuchus plummeri, from

1342-937X/$ – see front matter © 2008 International Association for Gondwana Research. Published by Elsevier B.V. All rights reserved.
doi:10.1016/j.gr.2008.07.002
132 S. Dias-da-Silva et al. / Gondwana Research 15 (2009) 131–136

Northern Brazil (Pedra de Fogo Formation, Parnaíba Basin) (Price,1948;
Cox and Hutchinson, 1991) and Bageherpeton longignathus, from
Southern Brazil (Morro Pelado Member, Rio do Rasto Formation,
Paraná Basin) (Dias and Barberena, 2001). In addition, basal repre-
sentatives of stereospondyls, namely rhinesuchids, are recorded in
Upper Permian deposits from Brazil: an unclassified mandible (from
Morro Pelado Member) (Malabarba et al., 2003), the long snouted
Australerpeton cosgriffi, and a short snouted unclassified rhinesuchid
skull, both from the locality of Serra do Cadeado (Rio do Rasto
Formation) (Barberena, 1998; Barberena and Dias, 1998; Dias and
Schultz, 2003). One could argue that subsequent stereospondyls found
in South American Triassic strata are descendants of rhinesuchids.
However, phylogenetic data of stereospondyls suggest that there is no
direct relationship between them and Lower Triassic taxa from South
America (see Yates and Warren, 2000). In other words, rhinesuchids
colonized Gondwana during the Permian and disappeared in the
landmass that nowadays comprises South America prior to the Permo-
Triassic extinctions. This group survived as a relict occurrence in South
Africa during the Lower Triassic. Hence, the subsequent stereospondyl
fauna from South America represents a reinvasion of this landmass
during the Early Triassic. Moreover, according to Damiani (2004), the
taxonomy of rhinesuchids is problematic and a forthcoming revision of
their anatomy and taxonomy is necessary in order to resolve their
phylogenetic relationships.
The record of stereospondyls from the Brazilian Lower Triassic
(Sanga do Cabral Formation, Paraná Basin) comprises dermal skull
fragments tentatively attributed to ‘lydekkerinids’ and rhytidos-
teids (Lavina and Barberena, 1985; Dias-da-Silva et al., 2005) and
the rhytidosteid Sangaia lavinai (Dias-da-Silva et al., 2006; Dias-da-
Silva and Marsicano 2006). In Uruguay, strata close to the Permo-
Triassic boundary have yielded a poorly preserved dvinosauroid
and mastodonsauroids, and the laidlerid Uruyiella liminea, both
from the Buena Vista Formation (Marsicano et al., 2000; Piñeiro et
al., 2007a,b,c).
The new stereospondyl described herein constitutes the first
undoubted occurrence of a member of this group in Upper Triassic
strata from Brazil (Caturrita Formation, Paraná Basin). Indeed, this is
the only stereospondyl record for the whole Santa Maria Super-
sequence (sensu Zerfass et al., 2003), which includes also the Santa
Maria Formation and its plentiful tetrapod record. The Caturrita
Formation also bears a rich tetrapod fauna including dinosaur remains,
non-mammalian therapsids, procolophonids and sphenodontids (see
Langer et al., 2007). The presence of an amphibian within this fauna
corroborates the sedimentological evidences towards a more humid
climate at the time of deposition of these layers, represented by an
increase in the rate of arenous fluvial/lacustrine deposits formed at the
transition from Santa Maria to Caturrita Formation (the Santa Maria 2
third order Sequence from Zerfass et al., 2003).

Fig. 1. In A, Triassic Sequences from Southern Brazil. See the stratigraphic position of Mammaliamorpha Cenozone. Abreviations of the ages: Any, Anysian; Car, Carnian; Ind, Induan;
J, Jurassic; K, Cretaceous; Lad, Ladinian; Nor, Norian; Ole, Olenekian; P, Permian; Rha, Rhaetic. In B, geographic location of the Botucaraí Oucrop in South America. The simbol (⁎)
indicates the location of the outcrop. Modified from Zerfass et al. (2003), and Vega-Dias and Schultz (2004).
 S. Dias-da-Silva et al. / Gondwana Research 15 (2009) 131–136 133

Fig. 2. Composite photograph of the southern margin of the “Botucaraí Outcrop” (above) and its schematic section (below). D = Disconformity; SD= Subaqüous Dunes; SL= Sigmoidal Lobes;
TB= Lacustrine Turbidites. The arrows indicate the level from where the fossil was recovered. UFRGS PV 1059 T was found in a disconformity between the SD facies and the SL facies.

Institutional abbreviations. UFRGS, Universidade Federal do Rio
Grande do Sul; PV, Paleovertebrate collection; T, Triassic.
2. Geological Setting
The Triassic infilling of the Paraná Basin in southern Brazil (Rosário
do Sul Group) is entirely non-marine (Pierini et al., 2002; Zerfass et al.,
2003). In terms of sequence stratigraphy, according to Zerfass et al.
(2003) the Middle–Upper Triassic of the Rio Grande do Sul State
corresponds to a second order sequence (Fig. 1), denominated Santa
Maria Supersequence. It is divided into three third-order sequences:
the Santa Maria 1 (Ladinian age), Santa Maria 2 (Carnian age) and
Santa Maria 3 (Rhaetian–Early Jurassic age) sequences. Each of these
third-order sequences begins with fluvial deposits of low sinuosity
rivers covered by transgressive shallow lacustrine rocks.
The stereospondyl material here discussed was collected at the
northern margin of the so-called “Botucaraí Outcrop”, a road cut sited
at the margins of BR-287 road, near the Botucaraí Hillock, 6 km west of
the entrance to Candelária City, Rio Grande do Sul State (coordinates
29° 40′ 48″ S ; 52° 50′ 24″ W). The siltstones and fine-grained
sandstones beds that crop out in this area are lithostratigraphycally
included in the Upper Triassic Caturrita Formation (Andreis et al.,
1980) and are interpreted as deposited in a fluvial-deltaic environ-
ment (Fig. 2). This unit is part of the Santa Maria 2 third order
Sequence of Zerfass et al. (2003). Recently, the same author (Zerfass,
2007) after a detailed mapping of an area located about 100 km W

Fig. 3. Stereospondyl interclavicle UFRGS PV 1059 T from the Upper Triassic of Southern Brazil in ventral view. In A photograph; in B, interpretative drawing; in C, estimated outline.
Scale bar equals 10 mm (C not to scale).
134 S. Dias-da-Silva et al. / Gondwana Research 15 (2009) 131–136

from the Botucarai Outcrop, encountered a disconformity between the vicle the distance between ridges is even more obvious. The dorsal
Santa Maria and Caturrita formations and interpreted the last one as surface of UFRGS PV 1059 T is flat, with no signal of processes or
an individual third-order sequence, called Caturrita Sequence. If such depressions. Therefore is uninformative for taxonomic purposes.
disconformity could be extended to the area of the Botucaraí Outcrop,
it would be placed below the level where the stereospondyl was found
(so this level clearly belongs to the Caturrita Formation, based on its
facies and paleofauna). The nearest fossil bearing outcrop from the
Santa Maria Formation (which includes Hyperodapedon, Barberena-
champsa and Exaeretodon) is located 7 km W (and at least 30 m below)
from the Botucaraí Outcrop.

2.1. The age and faunal content of the Caturrita Formation

Rubert and Schultz (2004) proposed the “Ictidosaur Association

Zone” for the fossiliferous beds of the Caturrita Formation based on the
presence of a distinct tetrapod association characterized by little non-
mammalian “ictidosaurian” cynodonts, including Riograndia guaiben-
sis, Irajatherium hernandezi, Brasilodon quadrangularis and Brasilither-
ium riograndensis. In addition to the non-mammalian cynodonts, the
procolophonid Soturnia caliodon, the dinosaurs Guaibasaurus cande-
lariensis and Unaysaurus tolentinoi, the dicynodont Jachaleria cande-
lariensis, the sphenodontid Clevosaurus brasiliensis, an indeterminate
phytosaur and footprints attributed to prosauropod dinosaurs also
characterized the Caturrita tetrapod fauna. For the complete set of
taxonomic references, see da Silva et al. (2007) and Langer et al.
(2007). The putative “ictidosaurs” mentioned above were recently
moved (Soares, 2004) to Tritheledontidae (Riograndia and Irajather-
ium) and Brasilodontidae (Brasilodon and Brasilitherium) which are
considered stem-groups of Mammalia, and thus included in the clade
Mammaliamorpha (Soares and Schultz, 2006). Therefore, Schultz and
Soares (2006) proposed to change the name “Ictidosaur Association
Zone” for the fossiliferous levels of the Caturrita Formation to the
“Mammaliamorpha Cenozone”. This cenozone is correlated with the
“early Coloradian stage” from Argentina, which is considered early
Norian in age (Rubert and Schultz 2004).

3. Systematic Paleontology

Temnospondyli von Zittel 1888

Stereospondylomorpha Yates and Warren 2000
Stereospondyli von Zittel 1888
Mastodonsauroidea? (sensu Damiani, 2001) indet.

3.1. Material

UFRGS PV 1059 T, an incomplete interclavicle (Fig. 3A, B).

3.2. Locality and Horizon

Botucaraí Hillock, 6 km west of the entrance to Candelária City, Rio

Grande do Sul State, Brazil. Upper Carnian to Lower Norian of Paraná
Basin (Rubert and Schultz, 2004; Bonaparte and Sues, 2006; Langer
et al., 2007).

3.3. Description

The material consists of an incomplete interclavicle of flattened

rhomboidal shape (11 mm tick, 112 mm wide, and 161 mm long). The
ventral surface of the interclavicles is covered by a coarse spider web
reticulate pattern, characterized by straight ridges that radiate from
the ossification centre. The ridges are shallow (deepness of about
1.5 mm) and are interconnected by relatively wide crests; nodes or
pustules are absent. This type of ornamentation is similar to that
present in Almasaurus (Warren and Snell, 1991), Siderops (Warren and
Marsicano, 2000) and indeterminate brachyopoid material from
Argentina (Marsicano, 1993). Nevertheless, in the Brazilian intercla-
Fig. 4. Interclavicular pattern amongst different groups of stereospondyls. In A,
Metoposauridae (modified from Chowdhury, 1965); in B, mastodonsauroid Mastodon-
saurus giganteous (modified from Schoch and Milner, 2000); in C indeterminate brachyopoid
from Argentina (modified from Marsicano,1993); in D, mastodonsauroid Promastodonsaurus
bellmanni with attached clavicles (modified from Bonaparte, 1963). Since these elements are
for comparison to UFRGS PV 1059 T (being of different sizes), none of them is to scale.
 S. Dias-da-Silva et al. / Gondwana Research 15 (2009) 131–136
According to the strong ossification degree of this dermal bone and its
heavy ornamentation, the specimen probably belonged to an adult
individual (see Steyer 2000, and Steyer et al., 2004 for the somatic age
criteria of the temnospondyls).
In spite of the poor preservation of the external borders of the
material it presents the flattened rhomboidal shape, little longer than
wide, typical of stereospondylomorphs. A strip of ornamentation,
which presumably extended to the anterior tip of the interclavicle
thus, separates the clavicular articular facets; the clavicles when
articulated did not completely overlap the interclavicular blade.
According to Warren and Snell (1991), this is the condition present
in Mastodonsauridae and Almasauridae, and some Permian taxa as
Eryops. Conversely, in the Triassic Metoposauridae, Trematosauridae,
Capitosauridae, Benthosuchidae, Plagiosauridae, and Siderops the
clavicular facets meet anteriorly so the interclavicular stem lacks any
ornamentation (Warren and Snell, 1991).
The centre of ossification of the interclavicle is apparently located
posterior to the level of the maximum width of the interclavicular
blade. In metoposaurids (Fig. 4A), the centre of ossification is situated
markedly posterior to this point, while in mastodonsaurids (including
Promastodontosaurus) and the rhinesuchid Australerpeton just poster-
iorly located (Warren and Snell, 1991; Dias and Schultz, 2003). This
condition presumably occurs in the Brazilian specimen.
4. Discussion
Non-stereospondyl temnospondyls are quite rare in Mesozoic
deposits. In fact, by the beginning of the Triassic all major
stereospondyl clades were already established in Pangaea (e.g. Schoch
and Milner, 2000; Warren and Marsicano, 2000; Yates and Warren,
2000; Damiani, 2001). Although the interclavicle here described is not
associated with cranial material that would assure a more accurate
taxonomic assignation, it is more likely to represent a stereospondyl
temnospondyl taxon. Within Stereospondyli, the general rhomboidal
shape, the pattern of sculpture and the thickness of the Brazilian
interclavicle relate it with those described for mastodonsauroids
(Schoch and Milner, 2000, and Damiani, 2001) (Fig. 4B) and
almasaurids (Warren and Snell, 1991).
Upper Triassic South American stereospondyls with interclavicular
elements preserved are restricted to the mastodonsauroid Promasto-
dontosaurus bellmanni and indeterminate brachyopoid material
both from the Upper Triassic of central-western Argentina (Fig. 4C)
(Bonaparte, 1963; Marsicano, 1993).
In UFRGS PV 1059 T, the centre of ossification is presumably located
behind the level of the maximum width of the interclavicular blade. A
condition quite similar to that found to the neighbor Argentinean
mastodonsauroid Promastodontosaurus (Bonaparte, 1963) (Fig. 4D). In
fact, UFRGS PV 1059 T is remarkably similar to the interclavicle of
Mastodonsaurus giganteous (Fig. 4B), especially regarding the pattern
of ornamentation. Therefore, it is reasonable to assume that this group
might have been present in Upper Triassic environments from
Southern Brazil. Further corroboration provided by more complete
material is necessary to reinforce this hypothesis.
The tetrapod content of the Upper Triassic Caturrita Formation is
rich and diversified; however, stereospondyls were unknown from
this unit until now. In spite of the difficulty to assign this new
specimen to a less inclusive group, its presence in the Upper Triassic of
Brazil corroborate the hypothesis of Marsicano (2005), who suggested
that temnospondyls were probably more diversified in this part of
Gondwana during the Late Triassic than it is evidenced by the fossil
record.
So far, the discovery of this stereospondyl constitutes a single
record for the Middle–Upper Triassic package of southern Brazil. This
package includes the Santa Maria and Caturrita formations, both
characterized by the presence of abundant fossil terrestrial tetrapod
faunas. Nevertheless, the presence of aquatic or even semi-aquatic
135
forms within these faunas, in the whole Santa Maria 1 Sequence and in
the lower part of Santa Maria 2 Sequence is rare (restricted only to a
few proterochampsid archosaurs (Chañaresuchus, Cerritosaurus and
Barberenachampsa). Besides, no fossil fishes or aquatic invertebrates
were encountered associated to the tetrapods. In spite of the
preservation of these fossils in floodplain deposits, the availability of
permanent water bodies would be insufficient to maintain aquatic
forms of life. Towards the top of the Santa Maria 2 Sequence, the
transition from Santa Maria to Caturrita Formation is characterized by
an increase of sandy facies represented by fluvial\deltaic deposits. This
indicates a change to more humid climatic conditions at that time, and
the presence of a phytosaur and an amphibian in the faunal
assemblage of the Caturrita Formation reinforces such hypothesis.
Regarding the paleobiogeography of advanced stereospondyls in
South America, the presence of rhytidosteids in Lower Triassic
deposits from Brazil, as well as mastodonsaurids and the enigmatic
Uruyiella liminea in Permo-Triassic deposits from Uruguay (Dias-da-
Silva et al., 2005, 2006; Piñeiro et al., 2007a,b,c) suggests that they first
colonized Brazil and Uruguay, probably spreading from South Africa.
Later, they reached Argentina during late Early, Middle or Late Triassic.
It is important to point out that Lower Triassic deposits are quite rare
in Argentina. Therefore, an alternative scenario cannot be dismissed,
with stereospondyls colonizing the Argentinean Gondwana still
during Early or Middle Triassic. Unfortunately, the small amount of
information regarding the Lower Triassic in Argentina is an obstacle to
clarify the sequence of events related to the diversification of
stereospondyls in this part of Gondwana. Besides, to the present,
there is no record of stereospondyls in Middle Triassic deposits from
both Argentina and Brazil. Alternative explanations for their absence
in the Middle Triassic of South America are either taphonomic bias or
insufficient prospecting in deposits of this age. Stereospondyls (and
other aquatic animals), tend to be less represented than other
vertebrate groups, due to taphonomic bias, because the fluvial style
causes a direct impact on the taphonomic mode of preservation.
According to Fonseca and Scherer (1998) and Pires et al. (2005) the
Triassic was marked by a large degree of seasonality, with large dry
periods replaced by shorter periods of humidity. The increasing of
humidity towards the top of the Triassic succession modified the
fluvial environments from anastomosed to braided systems (Fonseca
and Scherer, 1998). In braided systems, intense reworking tends to
destroy delicate bones. Therefore, animals with slender skeletons (e. g.
stereospondyls and phytosaurs) are likely to be less represented in
this kind of depositional environment. In conclusion, further pro-
specting efforts are necessary in order to clarify the likely sequence of
events related with the diversification of stereospondyls in South
American Gondwana which would further contribute to our under-
standing of the evolution of West Gondwana (Vaughan and Pankhurst,
2008), as well as the paleobiogeography of crustal fragments in the
erstwhile Gondwana assembly (cf. Meert and Lieberman, 2008).
Acknowledgements
SDS would like to thank the Universidade Federal do Pampa/UFSM,
for the facilities provided during the present study, Claudia Marsicano
for a critical review of an earlier version of this manuscript, Cristina
Machado-Bertoni for her useful taphonomic insights, Juan Carlos
Cisneros for the interpretative drawing of the Brazilian interclavicle
(Fig. 3B), and Lúcia Helena do Canto Vinadé for English revision. In
addition, we thank Jean Sébastien Steyer and an anonymous referee
for their suggestions that greatly improved the manuscript.
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