The Science of Nature (2025) 112:28

https://doi.org/10.1007/s00114-025-01981-5

ORIGINAL ARTICLE

New proterochampsid remains from the Middle Triassic of Brazil

enhance the group's diversity during its origins
Rodrigo Temp Müller1

Received: 16 January 2025 / Revised: 5 March 2025 / Accepted: 13 March 2025

© The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature 2025

Abstract
Proterochampsidae is a clade of archosauriform reptiles that lived during the Triassic Period, with notable fossil records in
South America. Among these, Ladinian deposits in Brazil have yielded the oldest proterochampsids worldwide, offering
insights into the origins of the group. However, their record from this interval remains scarce, with only two species previ-
ously described: Pinheirochampsa rodriguesi and Retymaijychampsa beckerorum. Here, I report the first proterochampsid
specimen from the Posto site (Pinheiros-Chiniquá Sequence; Dinodontosaurus Assemblage Zone) in Dona Francisca, Rio
Grande do Sul. The new specimen, CAPPA/UFSM 0459, comprises the anterior tip of a rostrum, identified as a protero-
champsid based on its dorsoventrally flattened premaxilla with a dorsally facing external naris and anterior narial fossa.
Despite its fragmentary nature, CAPPA/UFSM 0459 represents the first non-archosaur archosauriform identified at the Posto
site, suggesting a small carnivorous member of this local fauna. Comparisons indicate it is unlikely to belong to any previ-
ously described species. This finding highlights the diversity and ecological significance of early proterochampsids within
Middle Triassic ecosystems dominated by therapsids, pseudosuchians, and early dinosauromorphs.

Keywords Archosauriformes · Archosauromorpha · Ladinian · Proterochampsidae · Santa Maria formation

Introduction Paes-Neto et al. 2024; Müller 2025). These fossils span a

temporal range from the Ladinian (Paes-Neto et al. 2024;
Proterochampsidae is a clade of archosauriform reptiles Müller 2025) to the early Norian (Reig 1959; Barberena
that lived during the Triassic Period (Trotteyn et al. 2013; 1982), highlighting the significance of proterochampsids
Ezcurra et al. 2021). Members of this group are character- as key components of Triassic ecosystems in the region.
ized by elongated skulls with dorsally positioned external In addition, as close relatives of archosaurs (Ezcurra 2016;
nares, asymmetrical feet, and a wide range of body forms Ezcurra and Sues 2021; Müller et al. 2022; Paes-Neto et al.
adapted to both terrestrial and semi-aquatic environments 2024), proterochampsids are critical to understanding the
(Reig 1959; Sill 1967; Arcucci et al. 2019; Ezcurra et al. early diversification of this dominant lineage of reptiles.
2021; García Marsa et al. 2023). Fossil evidence indicates The fossil record of proterochampsids and their close rela-
that proterochampsids were predominantly distributed in tives in Brazil is notably abundant, comprising two species
what is now South America, with their remains found in from the Ladinian (Paes-Neto et al. 2024; Müller 2025), one
Middle to Late Triassic deposits of Argentina and Brazil from the early Carnian (Paes-Neto et al. 2024), and four from
(Price 1946; Reig 1959; Barberena 1982; Romer 1971; the late Carnian beds (Huene 1946; Price 1946; Barberena
Trotteyn and Ezcurra 2014, 2020; Ezcurra et al. 2015; 1982; Müller et al. 2022). Whereas the record from the Car-
nian is more diverse, the record from the Ladinian beds is
relatively scarce, limited to specimens from the municipal-
Communicated by: Dany Azar ity of Candelária (Paes-Neto et al. 2024) and Paraíso do Sul
(Müller 2025). This record from the Ladinian is particularly
* Rodrigo Temp Müller
[email protected]; [email protected] significant as it represents the oldest proterochampsids world-
wide, shedding light on the origins of the group. Despite the
1
Centro de Apoio À Pesquisa Paleontológica, Universidade relative abundance of Ladinian deposits in Brazil (Martinelli
Federal de Santa Maria, Rua Maximiliano Vizzotto, et al. 2017; Schultz et al. 2020), only a few localities from
São João Do Polêsine, RS 598, 97230‑000, Brazil

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the municipalities mentioned above have yielded fossils of taxonomic units (OTUs), with the new specimen incorpo-
this group. One of the most significant Ladinian outcrops in rated. The most parsimonious trees (MPTs) were recovered
Brazil is the Posto site, located in the municipality of Dona using the software TNT v. 1.5. (Goloboff and Catalano
Francisca. This fossiliferous site yielded a wide diversity of 2016), following the parameters employed by Müller (2025).
tetrapods (França et al. 2011; Martinelli et al. 2017; Pavanatto These parameters includes an equally weighted parsimony
et al. 2016; Roberto-da-Silva et al. 2020; Ugalde et al. 2020; analysis with characters 1, 2, 7, 12, 13, 19–23, 30, 31, 34, 36,
Müller 2021; Müller and Garcia 2022); however, no evidence 41, 44, 46, 52, 57, 58, 62, 76, 90, 111, 116, 118, 126–130,
of proterochampsids have been published so far. In the present 146, 151, 165, 167, 169, 181, 207, 216, 219, 222, 224, 227,
study, I describe the first proterochampsid specimen from the 235, 240, 241, 243, 247, 249, 255, 260, 262, 268, 269, 278,
Posto site, which not only extends the Ladinian record of the 283, 288, 290, 291, 293, 294, 296, 300, 301, 306, 310, 312,
group but also increases the diversity of reptiles known from 313, 315, 316, 325, 330, 338, 339, 343, 352, 359, 360, 364,
this locality. 370, 373, 374, 385, 388, and 389 set as ordered. In addition,
Institutional abbreviations. BSPG, Bayerische Staatssa- Euparkeria capensis was used to root the MPTs, which were
mmlung für Paläontologie und Geologie, München, Ger- recovered using the random addition sequence + tree bisec-
many; CA, Colégio Anchieta, Porto Alegre, Rio Grande do tion reconnection (TBR), which included 1000 replicates of
Sul, Brazil; CAPPA/UFSM, Centro de Apoio à Pesquisa Wagner trees (random seed = 0), TBR and branch-swapping
Paleontológica da Quarta Colônia, Universidade Federal de (holding 10 trees saved per replicate). Nodes without syna-
Santa Maria, Rio Grande do Sul, São João do Polêsine, Brazil; pomorphies were collapsed after the search. Decay indices
MCP, Museu de Ciências e Tecnologia, Pontifícia Universi- (Bremer support values) and bootstrap values (1000 repli-
dade Católica do Rio Grande do Sul, Rio Grande do Sul, Porto cates) were obtained with TNT v. 1.5.
Alegre, Brazil; PULR, Paleontología, Universidad Nacional
de La Rioja, La Rioja, Argentina; PVSJ, División de Pale- Systematic paleontology
ontologia de Vertebrados del Museo de Ciencias Naturales
y Universidad Nacional de San Juan, San Juan, Argentina; ARCHOSAUROMORPHA Huene 1946
UFRGS, Coleção de Paleontologia de Vertebrados, Universi- ARCHOSAURIFORMES Gauthier et al. 1988
dade Federal do Rio Grande do Sul, Rio Grande do Sul, Porto PROTEROCHAMPSIDAE Sill 1967
Alegre, Brazil. Material. CAPPA/UFSM 0459. Anterior tip of the ros-
trum, comprising both partial premaxillae (Fig. 2).
Geological setting Description. The left premaxilla preserves the main
body, the anterior portion of the posterolateral process, and
The specimen was collected from the reddish mudstones of the the anterior portion of the posteromedial (= internarial) pro-
Posto site (Fig. 1) site, which is located at the municipality of cess. The right premaxilla preserves the medial portion of
Dona Francisca (29°37′35.6″ S, 53°22.2′2.8″ W), Rio Grande the main body and the anterior portion of the posteromedial
do Sul state, southern Brazil. The outcrops at this site are part process. The external surface of the bone lacks ornamen-
of the Pinheiros-Chiniquá Sequence (Horn et al. 2014) of the tation (Fig. 2A), a condition shared with Pseudochampsa
Santa Maria Supersequence (Zerfass et al. 2003). According ischigualastensis (PVSJ 567; Trotteyn and Ezcurra 2014)
to geochronological data, the Pinheiros-Chiniquá Sequence and Chanaresuchus bonapartei (PULR 07; Trotteyn and
dates to the Ladinian (Middle Triassic; Philipp et al. 2023). Ezcurra 2020). In contrast, Proterochampsa nodosa (MCP-
The fossil content of the Posto site includes dicynodonts, 1694-PV; Barberena 1982) and Proterochampsa barri-
cynodonts, owenettids, pseudosuchians, and putative onuevoi (PVSJ 77; Reig 1959) exhibit well-developed protu-
dinosauromorphs (França et al. 2011; Martinelli et al. 2017; berances. There are some putative foramina located slightly
Pavanatto et al. 2016; Roberto-da-Silva et al. 2020; Ugalde dorsal to the alveolar margin (Fig. 2C).
et al. 2020; Müller 2021; Müller and Garcia 2022). In addition, The premaxillary body is dorsoventrally flattened, with
the occurrence of the dicynodont Dinodontosaurus assigns this its length being approximately 3 times its height. In Cerri-
fossil content to the Dinodontosaurus Assemblage Zone (AZ; tosaurus binsfeldi (CA without collection number; Trotteyn
Schultz et al. 2020). et al. 2013) and Pinheirochampsa rodriguesi (UFRGS-PV-
0464-T; Paes-Neto et al. 2024), the premaxillary body is
proportionally taller. In lateral view, the anterior tip of the
Material and methods bone is convex, the dorsal and ventral margins are straight.
At this point, the specimen differs from Proterochampsa
The phylogenetic affinities of the new specimen were nodosa (MCP-1694-PV; Simão-Oliveira et al. 2022) and
assessed using the data matrix of Müller (2025), which Proterochampsa barrionuevoi (PVSJ 606; Trotteyn et al.
includes 401 morphological characters and 27 operational 2013), in which the alveolar margin is convex in lateral view.
The Science of Nature (2025) 112:28 Page 3 of 7 28

Fig. 1  Provenance of CAPPA/UFSM 0459. A. Location and geologi- of the site. C. Artistic representation of CAPPA/UFSM 0459 in life,
cal context of the Posto site, Dona Francisca, Rio Grande do Sul, Bra- created by Caio Fantini. D. General view of the specimen CAPPA/
zil (silhouette based on the artwork of Caio Fantini). B. General view UFSM 0459
 28 Page 4 of 7 The Science of Nature (2025) 112:28

Fig. 2  Premaxillae of CAPPA/

UFSM 0459. A, dorsal view. B,
ventral view. C, left lateral view.
Abbreviations: nf, narial fossa;
plp, posterolateral process;
pmx1–5, premaxillary tooth 1 to
5; tt, tooth

In dorsal view, the lateral margin of the premaxilla is The posteromedial process is taller than wide, whereas
convex, and it is uncertain whether a constriction was pre- the posterolateral process exhibits the inverse condition.
sent at the level of the external naris (the lateral margin is The main body of the left premaxilla preserves five alveoli
not entirely preserved). A transverse constriction at level (Fig. 2B), which are circular in ventral view. The anterior-
of premaxilla-maxilla suture occurs in Rhadinosuchus gra- most alveolus contains an erupting tooth. The second alveolus
cilis (BSPG AS XXV 50; Ezcurra et al. 2015) and Pin- houses an erupted but incomplete tooth, which shows an ovoid
heirochampsa rodriguesi (UFRGS-PV-0464-T; Paes-Neto cross-section of the crown. The teeth are not ankylosed to the
et al. 2024). Still in dorsal view, the medial margin of the premaxilla. Further details of the tooth morphology remain
bone contacts its counterpart through a straight suture. unclear due to the absence of most of its crown. In lateral view,
There is a depression on the posterior portion of the main each alveolus is well-defined, with concave margins.
body, forming a marked narial fossa at the anterior portion
of the external naris (Fig. 2A). As is characteristic of pro-
terochampsians (Trotteyn et al. 2013), the external naris Results
is entirely dorsally oriented and therefore not visible in
lateral view (Fig. 2C). The medial margin of the external The phylogenetic analysis recovered 11 MPTs, each with
naris is bordered by the posteromedial process, and the 914 steps (consistency index: 0.521; retention index: 0.557).
lateral margin is bordered by the posterolateral process. CAPPA/UFSM 0459 nests within Proterochampsidae in all
The Science of Nature (2025) 112:28 Page 5 of 7 28

MPTs. In the strict consensus tree (Fig. 3), CAPPA/UFSM anterior narial fossa (Dilkes and Arcucci 2012; Trotteyn
0459 groups with typical rhadinosuchines and Tropido- et al. 2013; Paes-Neto et al. 2024). A less inclusive taxo-
suchus romeri in a large polytomy. In the majority of the nomic assignation is challenging due to the fragmentary
MPTs, CAPPA/UFSM 0459 is recovered within Rhadi- condition of the material. Nevertheless, additional compar-
nosuchinae, though in floating positions. There is no evi- isons with coeval specimens and remarks on the putative
dence in the analysis suggesting the placement of CAPPA/ taxonomic assignment of the new specimen are provided
UFSM 0459 within Proterochampsinae. The structure of the below.
most parsimonious trees resembles that recovered by Mül- CAPPA/UFSM 0459 represents the first non-archosaur
ler (2025), with Proterochampsidae divided into two main archosauriform recorded at the Posto site. Regarding the
groups: Proterochampsinae and forms closely related to Rha- archosauriforms from this locality, there are records of the
dinosuchinae. Due to the limited data provided by CAPPA/ pseudosuchian Prestosuchus chiniquensis (Mastrantonio
UFSM 0459, further observations on the distribution and et al. 2019; Roberto-da-Silva et al. 2020) and an indeter-
evolution of character states will not be explored. minate dinosauromorph (CAPPA/UFSM 0282; Müller and
Garcia 2022). Decuriasuchus quartacolonia (França et al.
2011) is another pseudosuchian from the Posto site. How-
Discussion and conclusion ever, some studies have suggested that it represents skel-
etally immature individuals of Prestosuchus chiniquensis
Despite the fragmentary nature of CAPPA/UFSM 0459, the (Farias et al. 2024; Corecco et al. 2025). Whereas Pres-
new specimen is identified as a proterochampsid archosauri- tosuchus chiniquensis and Decuriasuchus quartacolonia
form based on its dorsoventrally flattened premaxilla, which are unequivocally carnivorous organisms, the diet of the
features a dorsally facing external naris and a pronounced indeterminate dinosauromorph remains uncertain due to

Fig. 3  Time-calibrated most parsimonious tree showing the phyloge- reconstruction of CAPPA/UFSM 0459 by Caio Fantini. Smaller illus-
netic position of CAPPA/UFSM 0459. Number on nodes represent trations by Matheus Fernandes Gadelha
Bremer support values and Bootstrap values higher than 50%. Life
 28 Page 6 of 7 The Science of Nature (2025) 112:28

its fragmentary and poorly preserved condition. Therefore, could have played an important ecological role in ecosys-
the record of a proterochampsid at the Posto site represents tems mainly dominated by therapisds, large pseudosuchians,
the first evidence of a small carnivorous archosauriform and the first members of the dinosaurian lineage.
within this local fauna. Actually, the tooth morphology of
Acknowledgements I thank curator Everton Paz (MCP) for grant-
CAPPA/UFSM 0459 is poorly preserved, making it difficult ing access to the specimens under his care, paleoartists Caio Fantini
to determine its diet with certainty. On the other hand, all and Matheus Fernandes Gadelha for providing the illustrations used
proterochampsids known so far are carnivorous (Trotteyn in figures 1 and 2, and the Willi Hennig Society for making the TNT
et al. 2013). Thus, the feeding behavior of CAPPA/UFSM software freely available. I extend my gratitude to the two anonymous
reviewers for their valuable comments and suggestions. This work was
0459 is inferred based on this context. carried out with aid of the Conselho Nacional de Desenvolvimento
Regarding the record of proterochampsids from other Científico e Tecnológico (CNPq 404095/2021-6; 303034/2022-0; and
Ladinian localities in southern Brazil, two species have 406902/2022-4).
been described to date: Pinheirochampsa rodriguesi Paes-
Neto et al. 2024, a rhadinosuchine from the municipality Declarations
of Candelária; and Retymaijychampsa beckerorum Müller Conflict of interest The authors declare no competing interests.
2025, a proterochampsine from the municipality of Paraíso
do Sul. There is a putative additional specimen identified as
an indeterminate rhadinosuchine (MCP-4195-PV; Paes-Neto
et al. 2024). Initially, this specimen was tentatively assigned References
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