Journal of Systematic Palaeontology

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A new proterochampsid (Archosauriformes:

Proterochampsia) from the Late Triassic
of southern Brazil and the emergence of
archosaurian hind limb traits

Rodrigo Temp Müller, Mauricio Silva Garcia & André de Oliveira Fonseca

To cite this article: Rodrigo Temp Müller, Mauricio Silva Garcia & André de Oliveira Fonseca
(2022) A new proterochampsid (Archosauriformes: Proterochampsia) from the Late Triassic
of southern Brazil and the emergence of archosaurian hind limb traits, Journal of Systematic
Palaeontology, 20:1, 2128913, DOI: 10.1080/14772019.2022.2128913

To link to this article: https://doi.org/10.1080/14772019.2022.2128913

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Published online: 03 Nov 2022.

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 Journal of Systematic Palaeontology, 2022
Vol. 20, No. 1, 2128913
http://dx.doi.org/10.1080/14772019.2022.2128913

A new proterochampsid (Archosauriformes: Proterochampsia) from the Late

Triassic of southern Brazil and the emergence of archosaurian hind limb traits
Rodrigo Temp M€ullera,b
, Mauricio Silva Garciaa,b and Andre de Oliveira Fonsecac
a
Centro de Apoio a Pesquisa Paleontologica da Quarta Col^onia, Universidade Federal de Santa Maria, Rua Maximiliano Vizzotto,
598, 97230-000 S~ao Jo~ao do Pol^esine, RS, Brazil; bPrograma de Pos-Graduaç~ao em Biodiversidade Animal, Universidade Federal
de Santa Maria, 97105-120 Santa Maria, RS, Brazil; cLaboratorio de Geologia e Pedologia da Universidade Federal de Juiz de
Fora, 36036-900 Juiz de Fora, MG, Brazil
(Received 22 June 2022; accepted 21 September 2022)

Characterized by an elongated snout, proterochampsids are carnivorous non-archosaur archosauriforms. The clade is
endemic to South America and its fossil record extends from the early Carnian to the late Carnian/early Norian. Nesting
close to Archosauria, it is a key clade for understanding the origin and evolution of archosaurian traits. Unfortunately,
hind limb elements are usually poorly preserved for the group. Therefore, the hind limb anatomy of proterochampsids
still lacks detailed descriptions. In the present study, we partially fill this gap with the description of a new
proterochampsid represented by a complete and well-preserved hind limb. Stenoscelida aurantiacus gen. et sp. nov. was
excavated from the late Carnian/early Norian (Late Triassic) beds of southern Brazil. A phylogenetic investigation
recovers the new taxon as a non-rhadinosuchine proterochampsid. The species bears an unusual set of traits for the
group, which provides clues on the evolutionary origins of some muscle attachment structures. For instance, the femur
of Stenoscelida aurantiacus gen. et sp. nov. possesses an anterior trochanter and an anterolateral scar. So far, these
features have not been reported in other non-archosaurian archosauriforms. Therefore, the new specimen indicates that
some typical archosaurian features evolved earlier than previously thought. The taxon also carries additional uncommon
features for proterochampsids, such as an iliofibularis tubercle on the anterior margin of the fibula and a vestigial
phalanx in digit V. In sum, Stenoscelida aurantiacus has one of the best-preserved hind limbs within Proterochampsidae
and sheds light on the polarization of important traits regarding the evolutionary context of Archosauria.
http://zoobank.org/urn:lsid:zoobank.org:pub:C78B7CE3-AB9B-4543-833E-B2A3FEA957D9
Keywords: Archosauria; Archosauromorpha; Carnian; Gondwana; Proterochampsidae; South America

Introduction Arcucci 1990; Ezcurra et al. 2019) and from the

Santacruzodon AZ (Raugust et al. 2013) of Brazil; how-
Proterochampsids are non-archosaurian archosauriforms ever, the fossil record of proterochampsids from the
that are endemic to South America (Trotteyn et al. early Carnian beds of Brazil is far less abundant than
2013; Ezcurra et al. 2015, 2021). Characterized by an the Argentinean record. The younger members of the
elongated snout, members of the clade were usually clade are excavated from the middle to late Carnian/
regarded as semi-aquatic predators according to general early Norian beds of the Ischigualasto Formation
similarities with crocodilians (Reig 1959; Romer 1971; (Argentina: Sill 1967; Trotteyn et al. 2012) and the
Bonaparte 1978; Trotteyn et al. 2013). However, more Candelaria Sequence (Brazil: Huene 1938; Price 1946;
recently, palaeohistological data (Arcucci et al. 2019) Barberena 1982).
and morphofunctional studies (Ezcurra et al. 2021) have Perhaps one of the most interesting matters regarding
indicated more terrestrial habits for at least some mem- Proterochampsidae is its phylogenetic position. The
bers of this clade. The fossil record of proterochampsids most comprehensive studies regarding archosauromorph
ranges from the early Carnian to the late Carnian/early affinities have recovered proterochampsids close to
Norian (Ezcurra et al. 2017) and is restricted to Archosauria (Arcucci 1989; Sereno & Arcucci 1990;
Argentina and Brazil (Ezcurra et al. 2015). The oldest Sereno 1991; Ezcurra 2016; Ezcurra et al. 2017, 2020a;
proterochampsids have been unearthed from the M€uller et al. 2020a; Trotteyn & Ezcurra 2020; Kellner
Massetognathus-Chanaresuchus Assemblage Zone (AZ) et al. 2022). Therefore, Proterochampsidae is a key
of the Cha~ nares Formation, Argentina (Romer 1971; clade for investigating the origin and distribution of

Corresponding author. Email: [email protected]

# The Trustees of the Natural History Museum, London 2022. All rights reserved.

Published online 03 Nov 2022

2 R. T. M€uller et al.

archosaurian features. Unfortunately, hind limb elements Laboratorio de Estratigrafia e Paleobiologia,

are usually poorly preserved for the group. This portion Universidade Federal de Santa Maria, Brazil; ULBRA,
of the skeleton holds traits usually regarded as diagnos- Universidade Luterana do Brasil, Coleç~ao de
tic for a number of archosaurian clades (Nesbitt 2011; Paleovertebrados, Canoas, Brazil; UWBM, Burke
Ezcurra 2016; Baron et al. 2017). In the present study, Museum of Natural History and Culture, Seattle,
we describe a new proterochampsid from the Late Washington, USA.
Triassic of Brazil composed of a complete and articu-
lated hind limb. The new taxon is the first protero- Material
champsid from the Varzea do Agudo site and provides The specimen is housed in the palaeovertebrates collec-
insight on the distribution of some classical archosaurian tion of the Centro de Apoio a Pesquisa Paleontologica
traits. da Quarta Col^onia da Universidade Federal de Santa
Maria, S~ao Jo~ao do Pol^esine, Rio Grande do Sul, Brazil.
Its collection number is CAPPA/UFSM 0293.
Material and methods
Terminology
Institutional abbreviations This study employs traditional (Romerian) anatomical
BNM, B€ undner Naturmuseum, Chur, Switzerland; CA, terms (i.e. ‘anterior’ and ‘posterior’). The nomenclature
Colegio Anchieta, Porto Alegre, Rio Grande do Sul, for femoral tubers follows Ezcurra (2016), whereas the
Brazil; CAPPA/UFSM, Centro de Apoio a Pesquisa nomenclature for muscle attachment structures follows
Paleontol ogica da Quarta Col^onia da Universidade Griffin & Nesbitt (2016).
Federal de Santa Maria, Rio Grande do Sul, Brazil;
CPEZ, Paleontology Collection of the Museu Phylogenetic analysis
Paleontol ogico e Arqueologico Walter Ilha, S~ao Pedro The phylogenetic affinities of the new taxon were inves-
do Sul, Brazil; CRILAR-Pv, Paleontologıa de tigated employing the data matrix of Ezcurra & Sues
Vertebrados, Centro Regional de Investigaciones (2021). We rescored character 502, regarding the pres-
Cientıficas y Transferencia Tecnologica, Anillaco, ence and shape of the anterior trochanter (¼ lesser or
Argentina; FC-DPV, Coleccion de Vertebrados Fosiles, minor trochanter; iliofemoralis cranialis muscle inser-
Departamento de Paleontologıa, Facultad de Ciencias, tion) of the femur, from the state ‘absent (0)’ to ‘present
Universidad de la Republica, Montevideo, Uruguay; and forms a steep margin with the shaft but is com-
GR, Ruth Hall Museum of Paleontology at Ghost pletely connected to it (1)’ in Gualosuchus reigi (see
Ranch, New Mexico, USA; ISI, Indian Statistical Description, below, for further information). The ana-
Institute, Kolkata, India; NHMUK, Natural History lysis was conducted in TNT v. 1.5 (Goloboff &
Museum, London, UK; NHMW, Naturhistorisches Catalano 2016). Following Ezcurra & Sues (2021), 40
Museum Wien, Vienna, Austria; NMQR, National operational taxonomic units (ISIR 1132,
Museum, Bloemfontein, South Africa; NMT, National Protanystropheus antiquus, Trachelosaurus fischeri,
Museum of Tanzania, Dar es Salaam, Tanzania; Tanystropheus haasi, UFRGSPV- 492-T, Malerisaurus
PIMUZ, Pal€aontologisches Institut und Museum, all NA, Arctosaurus osborni, CRILAR-Pv 461,
Universit€at Z€urich, Z€urich, Switzerland; PIN, Borissiak CRILAR-Pv 462, CRILAR-Pv 497, Cha~nares rhyncho-
Paleontological Institute of the Russian Academy of saur, PVSJ 2728, Eorasaurus olsoni, Archosaurus com-
Sciences, Moscow, Russia; PULR, Paleontologıa, plete, FC-DPV 2641, UFSM 11444, UFSM 11394,
Museo de Ciencias Naturales, Universidad Nacional de Vonhuenia fredericki, C. rossicus combined, C. magnus
La Rioja, La Rioja, Argentina; PVSJ, Division of combined, Chasmatosuchus vjushkovi, Koilamasuchus
Vertebrate Paleontology of the Museo de Ciencias gonzalezdiazi, Kalisuchus rewanensis holotype, NMQR
Naturales de la Universidade Nacional de San Juan, San 3570, CRILAR-Pv 499, Shansisuchus kuyeheensis,
Juan, Argentina; SNSB-BSPG, Staatliche Uralosaurus combined, Osmolskina czatkoviensis,
Naturwissenschaftliche Sammlungen Bayerns- Osmolskina complete, Triopticus primus, Otter
Bayerische Staatssammlung f€ur Pal€aontologie und Sandstone archosaur, Dagasuchus santacruzensis,
Geologie, Munich, Germany; TMM, Jackson School of Ctenosauriscus koeneni, Hypselorhachis mirabilis,
Geosciences Vertebrate Laboratory, University of Texas Waldhaus poposauroid, Vytshegdosuchus zbeshartensis,
at Austin, Austin, Texas, USA; UFRGS, Bystrowisuchus flerovi, Bromsgroveia walkeri,
Paleovertebrate Collection of the Laboratorio de Lutungutali sitwensis and Nyasasaurus parringtoni) and
Paleovertebrados da Universidade Federal do Rio two characters (9 and 119) were set as deactivated. The
Grande do Sul, Porto Alegre, Brazil; UFSM, characters were equally weighted and characters 1, 2, 7,
 A new proterochampsid from Brazil 3

10, 17, 19–21, 28, 29, 34, 36, 40, 42, 46, 50, 54, 66, biostratigraphically correlated with the Exaeretodon bio-
71, 74–76, 100, 122, 127, 146, 153, 156, 157, 171, 176, zone from the Ischigualasto Formation (Martınez et al.
177, 187, 202, 221, 227, 263, 266, 278, 279, 283, 324, 2013). A Bayesian age-model for the profile of the
327, 331, 337, 342, 345, 351, 352, 354, 361, 365, 368, Ischigualasto Formation at the Hoyada del Cerro Las
370, 377, 379, 386, 387, 398, 410, 414, 424, 425, 430, Lajas locality in Argentina recovered an age of
435, 446, 448, 454, 455, 458, 460, 463, 464, 470, 472, 227.94 þ 0.83/1.67 for the top of the Hyperodapedon
478, 482, 483, 485, 489, 490, 502, 504, 510, 516, 520, AZ (Desojo et al. 2020), indicating an age of c. 228 Ma
521, 529, 537, 546, 552, 556, 557, 567, 569, 571, 574, for the Exaeretodon biozone. A similar age (late
581, 582, 588, 636, 648, 652, 662, 701, 731, 735, 737, Carnian/early Norian, Late Triassic) for the Exaeretodon
738, 743, 749, 766, 784, 803, 810, 816, 850, 851, 872, sub-Assemblage Zone is inferred in the
875, 885 and 888 were treated as ordered (additive). Candelaria Sequence.
The most parsimonious trees were recovered from an
Ontogenetic assessment. The presence of some muscle
equally weighted parsimony analysis following the
attachment structures (e.g. anterior trochanter; anterolat-
search protocol by Ezcurra et al. (2020a): search
eral scar) in the femur of CAPPA/UFSM 0293 suggests
employing new technology search algorithms until 100
some degree of development if the ontogenetic pathways
optimal hits are reached, followed by a final round of
of other archosauriforms are considered (Griffin &
TBR branch swapping. Consistency and retention indi-
Nesbitt 2016). However, it is currently not possible to
ces were calculated with the script developed by
confirm whether the specimen reached its maximum
Spiekman et al. (2021) that does not take into account a
size or not.
priori deactivated terminals.
Diagnosis. Stenoscelida aurantiacus differs from all
other known proterochampsids in (
local autapomor-
Systematic palaeontology phies): possessing a slender femur; presence of an anter-
ior trochanter on the femoral head; presence of a raised
Archosauromorpha Huene, 1946 sensu Dilkes (1998) anterolateral scar above the anterior trochanter; fourth
Archosauriformes Gauthier, Kluge & Rowe, 1988 trochanter restricted to the proximal half of the femur;
sensu Gauthier et al. (1988) tibia approximately 84% the total length of the femur;
Proterochampsia Bonaparte, 1971 sensu in proximal view, the cnemial crest of the tibia is sub-
Kischlat (2000) equal in size to the proximal condyles; fibula with an
Proterochampsidae Sill, 1967 sensu Trotteyn (2011) iliofibularis tubercle on the proximal portion of the
Stenoscelida aurantiacus gen. et sp. nov. shaft
; proximal third of the fibular shaft is anteropos-
(Figs 1–5) teriorly expanded, tapering distally
; ratio between the
minimum midshaft width of metatarsal II and its total
length is 0.12; ratio between femoral length and meta-
Holotype. CAPPA/UFSM 0293, a complete and tarsal III is 2.5; and digit V with a phalanx
.
articulated right hind limb.
Etymology. The genus combines the Greek words
rsemό1 (¼ narrow) and rjeko1 (¼ hind leg), referring Description and comparison
to the slender leg of the creature. The specific epithet
derives from the Latin word aurantiacus (¼ orange), in Overview
allusion to the orange colour of the outcropping sedi- The preserved right hind limb of the holotype is entirely
ments of the Varzea do Agudo site (Fig. 1B). articulated (Fig. 2). Some portions of the specimen are
covered by a thick concretion layer, especially the femoral
Type locality. Varzea do Agudo site (¼ Janner site;
midshaft and the ankle. The external surface is well-
29 390 10.8900 S, 53 170 34.2000 W), Agudo, Rio Grande do
preserved in several places, revealing some muscle attach-
Sul, Brazil (Fig. 1).
ment points. However, the specimen was deformed by
Stratigraphic horizon. Lower portion of the Candelaria sedimentary compression. Therefore, the elements are lat-
Sequence (Horn et al. 2014) of the Santa Maria eromedially compressed, affecting the original shape. As
Supersequence (Zerfass et al. 2003), Parana Basin. The result, the midshaft of the limb bones is collapsed, show-
predominance of the cynodont genus Exaeretodon places ing artificial longitudinal grooves or sulci. Part of the fibu-
the site in the upper part (Exaeretodon sub-Assemblage lar midshaft is not preserved. The same is true for the
Zone; M€ uller & Garcia 2020) of the Hyperodapedon distal portion of metatarsal IV, precluding us from deter-
Assemblage Zone (Schultz et al. 2020), which is mining its total length. According to the femoral length
4 R. T. M€uller et al.

Figure 1. Provenance of Stenoscelida aurantiacus gen. et sp. nov. (CAPPA/UFSM 0293). A, location and geological context of the
Varzea do Agudo site (Agudo, Rio Grande do Sul, Brazil; modified from M€uller et al. 2020a); B, general view of the site; C,
reconstruction of the skeleton of Stenoscelida aurantiacus gen. et sp. nov. (CAPPA/UFSM 0293) depicting the preserved elements.

(147 mm), the specimen is smaller than the Argentinian Femur

taxa from the Ischigualasto Formation: Pseudochampsa The femur (Fig. 3) is sigmoid in lateral and medial
ischigualastensis (155 mm; Trotteyn & Ezcurra 2014) and views. The extremities are well ossified and expanded.
Proterochampsa barrionuevoi (179 mm; Trotteyn 2011). The femoral head is mainly anteriorly directed, whereas
A new proterochampsid from Brazil 5
6 R. T. M€uller et al.

in other proterochampsids it is more medially oriented. Gnathovorax cabreirai (0.16) and Buriolestes schultzi
Diagenetic processes might have exaggerated this condi- (0.15; M€uller & Garcia 2022).
tion. The general morphology of the femoral head The anterior surface of the distal portion bears an
resembles that of proterochampsids, whereas it differs extensor fossa (Fig. 3D), which results in a concave
from the anteromedially expanded femoral head of dino- anterior margin in distal view. On the opposite side, the
saurs (Nesbitt 2011) and from the hook-shaped head of popliteal fossa is well delimited (Fig. 3E). It is proximo-
lagerpetids and pterosaurs (Ezcurra et al. 2020a). The distally short, approximately 10% of the total length of
proximal articular surface bears a shallow straight the bone. This condition distinguishes the specimen
groove (Fig. 3C), which is usually absent in protero- from silesaurids and aphanosaurs, where the fossa is
champsids, except in an unnamed rhadinosuchine considerably longer (Nesbitt et al. 2010, 2017). The dis-
(Ezcurra et al. 2019; CRILAR-Pv 491) from the tal condyles are approximately at the same level in
Cha~ nares Formation. There is an anterior tuber (sensu anterior or posterior views. In addition, the lateral sur-
Ezcurra 2016) on the proximal portion of the femur, and face between the lateral condyle and the crista tibiofibu-
although the bone lacks a posteromedial tuber, the pos- laris is smooth, lacking a deep groove.
terior tuber is present. The greater trochanter is rounded
and tall (Fig. 3B), differing from the typical angled tro-
chanter of dinosaurs, and in the same way, there is no Tibia
dorsolateral trochanter on the proximal portion of the The tibia (Fig. 4) is 84% the total length of the femur
bone. Conversely, the anterolateral surface bears a raised (Table 1). This resembles the condition in
and rugose area (Fig. 3B) where the anterior (¼ lesser) Pseudochampsa ischigualastensis, where it is approxi-
trochanter is reported for a number of archosauriforms mately 82% (Trotteyn & Ezcurra 2014), and differs from
(e.g. ornithosuchids, aetosaurs, dinosauriforms). both Proterochampsa barrionuevoi (74.5%; Trotteyn
Whereas the anterior trochanter is absent in several pro- 2011) and Tropidosuchus romeri (100%; Arcucci 1990).
terochampsids (Trotteyn et al. 2013), the structure The bone is straight in anterior/posterior or medial/lateral
occurs in the holotype of Gualosuchus reigi (PULR-V views. The proximal portion is strongly expanded and is
05; M. D. Ezcurra, pers. comm.). A trochanteric shelf is triangular in shape in proximal view (Fig. 4C). The anter-
not associated with the anterior trochanter, and the prox- ior half of the proximal margin is proximally projected in
imal-most portion of this trochanter is completely con- medial view. In medial or lateral views, the well-devel-
nected to the shaft. Slightly above the proximal tip of oped cnemial crest extends anteriorly, and in proximal
the anterior trochanter there is an additional scar, which view it is straight and the anterior margin is rounded. The
resembles the anterolateral scar (sensu Griffin & Nesbitt cnemial crest is sub-equal in size regarding the proximal
2016) of aphanosaurs and dinosauriforms. There is a condyles, distinguishing the new species from
similar scar in Gualosuchus reigi (PULR-V 05; M. D. Proterochampsa barrionuevoi, where the crest is propor-
Ezcurra, pers. comm.). tionally smaller. The posterior and lateral condyles are
The transition from the femoral head to the shaft is sub-equal in size. The lateral condyle is offset regarding
smooth. The fourth trochanter rests on the posterior surface the posterior condyle. The posterior condyle does not
of the proximal third of the femur (Fig. 3A). It is crest-like taper posteriorly. The presence of any crest on the lateral
and symmetrical in shape. Its medial surface is ornamented surface of the proximal portion of the bone is uncertain.
with muscle scars. The distal portion of the fourth trochan- The midshaft is ovoid in cross-section. The bone wall
ter does not extend further down along the femoral shaft. is thick, distinct from the thin walls of several pan-avi-
In contrast, the fourth trochanter is strongly proximodistally ans and some crocodylomorphs (Kellner et al. 2022).
developed in Gualosuchus reigi and Chanaresuchus bona- The distal portion of the bone is moderately expanded.
partei (Ezcurra et al. 2019). The midshaft is slender. The It lacks a posterolateral process. There is a proximodis-
robustness index (RI, sensu Wilson & Upchurch 2003; i.e. tally oriented groove on the lateral surface of the distal
average of the greatest widths of the proximal end, mid- portion (Fig. 4F). It is more pronounced in
shaft and distal end of the element divided by the length Proterochampsa barrionuevoi (Trotteyn 2011). The dis-
of the element) is 0.14. For comparison, this is slightly tal outline of the tibia is elliptical. It is anteroposteriorly
slenderer than that of some early dinosaurs, such as longer than transversely wide.

3
Figure 2. Holotype of Stenoscelida aurantiacus gen. et sp. nov. (CAPPA/UFSM 0293). Right hind limb in A, medial and B, lateral
views. Abbreviations: 4t, fourth trochanter; a, astragalus; als, anterolateral scar; at, anterior trochanter; c, calcaneum; cc, cnemial
crest; f, femur; fh, femoral head; fi, fibula; ift, iliofibularis tubercle; mc, medial condyle; mt, metatarsal; pf, popliteal fossa; ph,
phalanx; t, tibia.
 A new proterochampsid from Brazil 7

Figure 3. Right femur of Stenoscelida aurantiacus gen. et sp. nov. (CAPPA/UFSM 0293) in A, anterior, B, lateral, C, proximal, D,
distal, E, medial and F, posterior views. Abbreviations: 4t, fourth trochanter; als, anterolateral scar; at, anterior trochanter; ctf, crita
tibiofibularis; ef, externsor fossa; fh, femoral head; gt, greater trochanter; lc, lateral condyle; mc, medial condyle; pf, popliteal fossa;
pg, proximal groove; pt, posterior tuber.

Fibula The proximal third of the midshaft is anteroposteriorly

The total length of the fibula (Fig. 4) is uncertain expanded, being wider than the tibia (Fig. 4F).
because a portion of the midshaft is not preserved. The Conversely, the shaft becomes extremely slender for the
bone is straight in anterior or lateral views, and the next two-thirds, being approximately two times narrower
proximal articular surface is anteroposteriorly expanded than the tibia. This is an unusual condition and distin-
and transversely compressed (Fig. 4C). It differs from guishes the specimen from other proterochampsids.
the rounded to elliptical proximal end of some pseudo- Furthermore, there is an iliofibularis tubercle on the prox-
suchians (e.g. Dynamosuchus collisensis, Prestosuchus imal portion of the shaft (Fig. 4E, F), a feature absent in
chiniquensis). In proximal view, the lateral margin is other proterochampsids. The condition in Stenoscelida
convex but the medial margin is concave. In lateral aurantiacus differs from the more distally located tuber-
view, the posterior margin projects posteriorly, whereas cle of several pseudosuchians and rhynchosaurs.
the anterior margin lacks any anterior expansion. The distal portion of the fibula is gently expanded. It
Conversely, in non-proterochampsid proterochampsians is approximately 0.45 times the maximum length of the
(e.g. Vancleavea campi; Litorosuchus somnii; proximal articular surface (Table 1). There is a faint lon-
Jaxtasuchus salomoni) the proximal portion is symmet- gitudinal ridge running on the anterolateral margin. The
rical to nearly symmetrical in lateral view. The posterior distal articular surface is flat to concave. In distal view
margin of the proximal third of the bone is sharp. (Fig. 4D), it is ovoid to triangular.
8 R. T. M€uller et al.

Figure 4. Right tibia and fibula of Stenoscelida aurantiacus gen. et sp. nov. (CAPPA/UFSM 0293) in A, medial, B, anterior, C,
proximal, D,
distal, E, posterior and F, lateral views.
Only fibula is visible in distal view. Abbreviations: cc, cnemial crest; fi,
fibula; ift, iliofibularis tubercle; lc, lateral condyle; lg, lateral groove; pc, posterior condyle; t, tibia.

Proximal tarsals Metatarsals

The astragalus and calcaneum are poorly preserved (Figs As in other proterochampsids, the metatarsals (Fig. 5)
2, 5). The astragalus is transversely wider (20.5 mm) than overlap each other. Metatarsal I is shorter than metatar-
anteroposteriorly long (11 mm). The anterior margin of sals II–IV (Fig. 5C). It is almost three times shorter
the astragalus is concave in dorsal view (Figs 2B, 5B). A than metatarsal III (Table 1). The proximal articular sur-
transverse groove runs on the posterior surface of the face is transversely expanded. There is a longitudinal
bone. The presence or absence of foramina on this pos- smooth crest on the dorsomedial corner of the proximal
terior groove is uncertain because it is badly preserved. half of the bone. The distal end is moderately expanded.
As in other proterochampsids, the tibial facet is wider The lateral condyle is more pronounced dorsally than
than the fibular facet. The latter facet is dorsolaterally the medial one. There is a shallow extensor fossa on the
oriented, resembling the condition of Chanaresuchus dorsal surface of the distal portion. The presence of col-
bonapartei and Proterochampsa barrionuevoi, whereas in lateral ligament pits on the condylar sides is uncertain.
Pseudochampsa ischigualastensis it is dorsally oriented The distal articular surface is not strongly ginglymoid.
(Trotteyn & Ezcurra 2014). The ventral surface of the Metatarsal II is the stoutest metatarsal. The proximal
bone is transversely convex. The calcaneum is 10 mm in articular surface is strongly expanded (Table 1). The
width (approximately half of the astragalar width). There ratio between the proximal articular surface and the total
is a posterolaterally oriented calcaneal tuber. This struc- length is 0.39. It is 0.36 in Rhadinosuchus gracilis
ture is sub-rectangular, with a straight lateral margin. In (Ezcurra et al. 2015). Conversely, the shaft is stouter in
dorsal view, the posterolateral corner of the calcaneal Rhadinosuchus gracilis, where the ratio between the
tuber is rounded (Fig. 5B). minimum midshaft width and the total length of
 A new proterochampsid from Brazil 9

Figure 5. Right pes of Stenoscelida aurantiacus gen. et sp. nov. (CAPPA/UFSM 0293) in A, medial, B, lateral and C, dorsal views.
Abbreviations: a, astragalus; c, calcaneum; mt, metatarsal; ph, phalanx; t, tibia.
10 R. T. M€uller et al.

Table 1. Measurements (in mm) of the right hind limb of Stenoscelida aurantiacus (CAPPA/
UFSM 0293).
Element Length Maximum proximal width Maximum distal width
Femur 147 29 24
Tibia 124 31 21
Fibula – 22 10
Metatarsal I 21 10 7
Metatarsal II 48 19 12
Metatarsal III 58 14 9
Metatarsal IV – – –
Metatarsal V 12 7 3

Table 2. Measurements (in mm) of the phalanges of covered by thick sediment). The lateral condyle is
Stenoscelida aurantiacus (CAPPA/UFSM 0293). slightly more pronounced laterally than the lateral con-
Element Length dyle of metatarsal II.
Digit I Metatarsal IV is incomplete (Fig. 5B). The element
Phalanx 1 13 does not preserve the distal portion. Therefore, its total
Phalanx 2 – length is uncertain. The preserved part is 30 mm in
Digit II length. Despite its condition, the element is longer than
Phalanx 1 19
Phalanx 2 17 metatarsals I and V. The proximal articular surface is
Phalanx 3 18 not transversely expanded. The general morphology of
Digit III the element is simple. It comprises an elongated and
Phalanx 1 19 compressed shaft. Distinct from the previous metatar-
Phalanx 2 13
sals, metatarsal IV is plate-like in cross-section.
Phalanx 3 9
Phalanx 4 10 Metatarsal V is the shortest (Fig. 5B), being slightly
Digit IV shorter than phalanx 1 of digit I. In this sense,
No phalanges Stenoscelida aurantiacus differs from Pseudochampsa
Digit V ischigualastensis, where metatarsal V is slightly longer
Phalanx 1 7
than the phalanx 1 from digit I (Trotteyn & Ezcurra
2014). The proximal articular surface is moderately
expanded and ‘U’-shaped in ventral view. The shaft
metatarsal II is 0.17. In Stenoscelida aurantiacus it is tapers distally to an unexpanded and featureless distal
0.12. On the dorsal surface of the distal portion, there is end, which bears no distal condyles.
a triangular extensor fossa. It is not possible to deter-
mine if the articular surface is ginglymoid. There is no
evidence of a collateral ligament pit on the medial sur- Phalanges
face of the medial condyle. On the other hand, the lat- The phalangeal formula is 2-3-4-?-1 (Fig. 5). Other pro-
eral surface of the lateral condyle bears a well-delimited terochampsids lack phalanges in digit V. The first phal-
collateral ligament pit (Fig. 5B). In dorsal view, the lat- anx of digit I is longer than broad. This is the same
eral condyle is straight, whereas the medial condyle is pattern observed in the other phalanges of the specimen.
slightly medially oriented. Both condyles are equally This phalanx is shorter than the first phalanx from digits
expanded distally, which is distinct from Rhadinosuchus II and III (Table 2). The dorsal intercondylar process is
gracilis, where the lateral condyle is far more extended moderately developed and the distal articular surface is
distally (Ezcurra et al. 2015). ginglymoid. There is an extensor depression on the dor-
Metatarsal III is the longest (Table 1), whereas its sal surface of the distal portion. This surface receives
midshaft is slenderer than that of metatarsal II. Its prox- the extensor tubercle from the ungual phalanx. The
imal articular surface is moderately expanded (maximum ungual (distal-most) phalanx of digit I lacks its distal
proximal width is 14 mm), far less than the proximal half. The phalanx is transversely compressed and tapers
articular surface of metatarsal II (19 mm). The distal distally (at least, the preserved portion). The flexor
articular surface resembles that of metatarsal II, with a tubercle is poorly developed (Fig. 5A). The first phalanx
marked extensor fossa on the dorsal surface and a col- of digit II is similar in shape to that of digit I. However,
lateral ligament pit on the lateral surface of the lateral the phalanx from digit II is larger (Fig. 5C). Phalanx 2
condyle (Fig. 5B). The presence of the collateral liga- is slightly smaller. Its anatomy is obscured by a thick
ment pit on the medial condyle is uncertain (it is layer of concretion. The ungual phalanx of digit II is
 A new proterochampsid from Brazil 11

clearly the largest ungual. It is transversely compressed a trichotomy including Rhadinosuchus gracilis,
and tapers to a sharp point. The ungual is not ventrally Pseudochampsa ischigualastensis and Chanaresuchus
recurved and lacks a well-developed flexor tubercle. bonapartei. Except for the polytomy at the base of
Digit III is the longest (Fig. 5B). The first phalanx from Proterochampsidae, the topology of the strict consensus
digit III is sub-equal in length to the first phalanx of tree is identical to that recovered by Ezcurra &
digit II (Table 2). On the other hand, the shaft of this Sues (2021).
phalanx is more gracile. The extensor fossa is deep on
the dorsal surface of the distal portion. The subsequent
phalanges of this digit are smaller. The ungual is Discussion
remarkably smaller than that of digit II. The number of
phalanges in digit IV is unknown and the distal portion Taxonomic status
of metatarsal IV is not preserved. Therefore, the absence In addition to a series of cranial features, proterochamp-
of phalanges is ambiguous. The phalanx of digit V (Fig. sids are characterized by a metatarsal II with a robust
5A) is reduced to an elongated structure with a concave midshaft, whereas metatarsal IV has a slender profile
proximal articular surface. No condyles or other struc- (Trotteyn et al. 2013). The new taxon possesses this
tures are present in this vestigial element. particular combination of traits. Therefore, Stenoscelida
aurantiacus is unequivocally assigned to
Proterochampsidae due to the shape of metatarsals.
Phylogenetic results Moreover, this is supported by our phylogenetic ana-
lysis. Unfortunately, the absence of overlapping ele-
The phylogenetic analysis recovered 39,200 most parsi- ments between Stenoscelida aurantiacus and several
monious trees (MPTs) of 6287 steps (consistency index other proterochampsids (especially the Brazilian forms)
¼ 0.18375; retention index ¼ 0.63457). Stenoscelida aur- precludes a clear assessment of its specific phylogenetic
antiacus nests within Proterochampsidae in all MPTs relationships. Rhadinosuchus gracilis is the only
(Fig. 6A). The clade is supported by 19 synapomorphies, Brazilian proterochampsid from the Hyperodapedon AZ
of which three are coded for Stenoscelida aurantiacus: 1) that preserves any hind limb elements. Actually, the
midshaft diameter of metatarsal II is more than the mid- holotype preserves a left metatarsal II (Ezcurra et al.
shaft diameter of metatarsal I (ch. 572: 0!1); 2) mid- 2015). It is the only comparable element between both
shaft diameter of metatarsal IV is lower than that of taxa. The latter is a member of Rhadinosuchinae
metatarsal III (ch. 573: 0!1); and 3) crest-like fourth tro- (Ezcurra et al. 2015), a less inclusive clade within
chanter, dorsoventrally/anteroposteriorly taller than or Proterochampsidae, a nesting reinforced by our results
equal to the shaft at its minimum depth (ch. 803: 1!2). (see above). On the other hand, Stenoscelida aurantia-
Proterochampsidae is the sister group to Doswelliidae, cus is not recovered as a rhadinosuchine in our analysis,
which is composed of Rugarhynchus sixmilensis, and despite the few comparable elements, there are
Sphodrosaurus pennsylvanicus, Doswellia kaltenbachi some differences between the metatarsal II of CAPPA/
and Jaxtasuchus salomoni. The clade supporting UFSM 0293 (Stenoscelida aurantiacus) and SNSB-
Proterochampsidae þ Doswelliidae is in a trichotomy with BSPG AS XXV 50 (Rhadinosuchus gracilis). In SNSB-
Polymorphodon adorfi and an unnamed clade composed BSPG AS XXV 50 the midshaft is stouter and the prox-
of Vancleavea campi and Litorosuchus somnii. These imal articular surface is proportionally smaller than in
clades are included within Proterochampsia. CAPPA/UFSM 0293. The medial and lateral condyles
Regarding the internal relationships of of SNSB-BSPG AS XXV 50 are uneven distally,
Proterochampsidae, there is a polytomy composed of whereas in CAPPA/UFSM 0293 both condyles are in
Stenoscelida aurantiacus, Tropidosuchus romeri, the same plane. A more detailed comparison demands
Cerritosaurus binsfeldi, Gualosuchus reigi, both species additional overlapping elements. In sum, CAPPA/UFSM
of Proterochampsa, and Rhadinosuchinae in the strict 0293 cannot be ascribed to Rhadinosuchus gracilis
consensus tree (Fig. 6A). Given the absence of overlap- according to the lack of anatomical correspondence and
ping elements between the new taxon and several other distinct phylogenetic positions.
proterochampsids (e.g. Proterochampsa nodosa, Although the holotype (CA unnumbered) of
Cerritosaurus binsfeldi), these unresolved affinities are Cerritosaurus binsfeldi preserves some postcranial ele-
unsurprising. The new taxon, however, does not nest ments, there are no hind limb bones (Price 1946;
within Rhadinosuchinae in any of the MPTs, thus Trotteyn et al. 2013). Despite the absence of overlap-
Stenoscelida aurantiacus is a non-rhadinosuchine proter- ping bones between Cerritosaurus binsfeldi and
ochampsid. The clade Rhadinosuchinae is composed of Stenoscelida aurantiacus, the former is far smaller, with
12 R. T. M€uller et al.

Figure 6. Results of the phylogenetic analysis depicting the position of Stenoscelida aurantiacus. A, time-calibrated reduced strict
consensus tree (number on nodes represent Bremer support values higher than 1); B, strict consensus tree depicting the occurrence
and distribution of the anterior trochanter (at) and the anterolateral scar (als) in the proximal portion of the femora of some pan-
archosaurs. Femora of Tanystropheus sp. (Tanystropheidae; PIMUZ A/III 771; Spiekman & Scheyer 2019), Malerisaurus-like
(Allokotosauria; TMM 31025-265; Nesbitt et al. 2022), Hyperodapedon sp. (Rhynchosauria; CAPPA/UFSM 0206), Prolacerta
broomi (Prolacertidae; UWBM 95529; Spiekman 2018), Garjainia prima (Erythrosuchidae; PIN 951/61-1; Maidment et al. 2020),
Stenoscelida aurantiacus (Proterochampsidae; CAPPA/UFSM 0293); Mystriosuchus steinbergeri (Phytosauria; NHMW 1986/0024/
0012; Butler et al. 2019), Dynamosuchus collisensis (Ornithosuchidae; CAPPA/UFSM 0248); Archeopelta arborensis
(Erpetosuchidae; CPEZ-239a; Desojo et al. 2011), Aetosauroides scagliai (Aetosauria; ULBRA-PVT-003; Roberto-da-Silva et al.
2014), Prestosuchus chiniquensis (Loricata; ULBRA-PVT-281; Roberto-da-Silva et al. 2020); Teleocrater rhadinus (Aphanosauria;
NHMUK PV R6795; Nesbitt et al. 2017), Dromomeron romeri (Lagerpetiae; GR 1308; Griffin et al. 2019); Raeticodactylus
filisurensis (Pterosauria; BNM 14524; Ezcurra et al. 2020a); Asilisaurus kongwe (Silesauridae; cast of NMT RB159), Buriolestes
schultzi (Dinosauria; CAPPA/UFSM 0035).

a cranial length of 8.9 cm (Trotteyn et al. 2013). For bonapartei, 16.3 cm and 11.96 cm (Trotteyn & Ezcurra
comparison, the skull and femoral lengths of other pro- 2020); Gualosuchus reigi, 33 cm and 17.2 cm (Ezcurra
terochampsids are, respectively: Chanaresuchus et al. 2020b); Proterochampsa barrionuevoi, 40.45 cm
 A new proterochampsid from Brazil 13

Figure 7. Plot of log10-transformed skull length versus log10-transformed femoral length of proterochampsids depicting (red dots) the
estimated values for Cerritosaurus binsfeldi and Stenoscelida aurantiacus. Red dotted lines represent the 95% confidence intervals.
Skulls modified from Ezcurra et al. (2021). Reconstructed head of Stenoscelida aurantiacus by Caio Fantini.

and 17.9 cm (Trotteyn & Haro 2011); Pseudochampsa 1982; Trotteyn et al. 2013; Sim~ao-Oliveira et al. 2022).
ischigualastensis, 25.4 cm and 15.48 cm (Trotteyn & Although there are no comparable elements between
Ezcurra 2014); and Tropidosuchus romeri, 7.67 cm and both Brazilian specimens, it is possible to compare
6.46 cm (Bestwick et al. 2022). Based on these measure- Stenoscelida aurantiacus with the Argentinian species
ments, the estimated femoral length of Cerritosaurus of the same genus: Proterochampsa barrionuevoi. The
binsfeldi is 7.54 cm (y ¼ 0.6262x þ 0.65702; Fig. 7), latter is known from several specimens, including an
which is approximately two times shorter than the fem- almost complete skeleton (PVSJ 606), and its hind limb
oral length of Stenoscelida aurantiacus (i.e. 14.7 cm). morphology differs substantially from that in
So far, there are no ontogenetic assessments regarding Stenoscelida aurantiacus, not to mention the three local
Cerritosaurus binsfeldi. Therefore, additional studies or apomorphies of the new taxon (see Diagnosis, above),
new fossils are necessary in order to determine if the which will not be repeated here. In relation to the
size discrepancy between both species is an effect of femur, the popliteal fossa of CAPPA/UFSM 0293
ontogeny or another intraspecific factor. Nevertheless, (Stenoscelida aurantiacus) is short, comprising 10% of
geological and stratigraphical evidence provides further the total femoral length, but in PVSJ 606
support for our taxonomic proposal. Cerritosaurus bins- (Proterochampsa barrionuevoi; Trotteyn 2011), it is lon-
feldi was excavated from strata ascribed to the ger in comparison, reaching roughly 23% of the total
Hyperodapedon Acme Zone (Langer et al. 2007), which femoral length. Furthermore, based on femoral measure-
are older than that which yielded Stenoscelida aurantia- ments, CAPPA/UFSM 0293 is 22% smaller than PVSJ
cus (i.e. Exaeretodon sub-AZ; Langer et al. 2007; 606, and although it is possible to argue that the speci-
Martınez et al. 2013; Desojo et al. 2020; M€uller & men probably does not represent an immature individual
Garcia 2020; Schultz et al. 2020). due to the presence of well-developed femoral epiphyses
In relation to the occurrence of Proterochampsa and scars (see above), we cannot determine with the evi-
nodosa in sediments attributable to the Exaeretodon dence at hand if CAPPA/UFSM 0293 reached its max-
sub-AZ, Stenoscelida aurantiacus does not present over- imum size. The proportions of the tibia and femur also
lapping elements with the former, which is known from differ between both specimens, where in CAPPA/UFSM
a single skull with mandible in occlusion (Barberena 0293 the tibia is proportionally longer than in PVSJ
14 R. T. M€uller et al.

606; although we recognize that the size difference no records of the anterior trochanter for non-archosaurian
between specimens could possibly influence this trait, archosaurs (to our knowledge). Stenoscelida aurantiacus
there is no evidence to support this sort of allometric (together with Gualosuchus reigi) provides the first evi-
discrepancy in proterochampsids. Therefore, as in the dence of the presence of this feature in a non-archosau-
case of Cerrritosaurus bindfeldi, more specimens are rian archosauriform (Fig. 6B). The occurrence of the
needed in order to test this assumption. Regarding the anterior trochanter in a proterochampsid supports a
tibia, in CAPPA/UFSM 0293 the proximal extremity of deeper origin for this trait than previously thought. It is
the tibia is perpendicular in relation to its long axis, possible that instead of evolving independently in the two
whereas in PVSJ 606 it is oblique. Moreover, in PVSJ main lineages of Archosauria, the anterior trochanter ori-
606, the cnemial crest is smaller than the proximal con- ginated earlier and was retained in the major clades of
dyles (Trotteyn 2011), whereas in CAPPA/UFSM 0293 Pan-Aves and in some members of Pseudosuchia.
it is sub-equal in size to the proximal condyles. The fib- However, the absence of this trait in the proterochamp-
ula of PVSJ 606 maintains its anteroposterior width for sian Vancleavea campi cast doubts on this hypothesis and
most of its proximodistal length, as in other protero- favours a scenario where the anterior trochanter evolved
champsids (although the epiphyses are slightly independently in proterochampsids and archosaurs.
expanded), but in CAPPA/UFSM 0293 the fibula is sig- Above the anterior trochanter, the femur of
nificantly expanded in the proximal third of the element Stenoscelida aurantiacus displays another interesting fea-
and then tapers distally, becoming remarkably narrow, ture. It comprises a raised surface that resembles the
ending in an unexpanded distal epiphysis. In sum, we anterolateral scar (¼ ‘dorsolateral ossification’ of
consider it unlikely that CAPPA/UFSM 0293 represents Piechowski et al. 2014) reported for dinosauriforms and
a new specimen of the genus Proterochampsa. aphanosaurs, both of which are members of Pan-Aves
(Piechowski et al. 2014; Griffin & Nesbitt 2016; M€uller
2022). Conversely, the structure is not recognized in
Distribution of hind limb traits pseudosuchians or non-archosaurian archosauriforms (Fig.
The occurrence of a new species of proterochampsid with 6B). It was hypothesized as the ossification of the iliofe-
a well-preserved hind limb is particularly exciting. moral ligament insertion (Griffin & Nesbitt 2016). The
According to several authors (e.g. Sereno 1991; Ezcurra occurrence of this trait in a non-archosaurian archosauri-
2016; Ezcurra et al. 2020a; Trotteyn & Ezcurra 2020; form implies a similar origin as that of the anterior tro-
Kellner et al. 2022), Proterochampsia is regarded as the chanter, meaning that it could have been secondarily lost
sister group to Archosauria. As a result, the anatomy of in pseudosuchians, whereas in Pan-Aves it was retained.
proterochampsians is crucial in order to optimize the ori- The alternative scenario implies the independent origin of
gin and distribution of morphological traits usually pre- the scar in Pan-Aves and proterochampsids (considering
sent in the earliest diverging clades of archosaurs. the presence in both Stenoscelida aurantiacus and
Several of these traits rely on the pelvic and hind limb Gualosuchus reigi). It has been demonstrated that the
anatomy (Nesbitt 2011; Ezcurra 2016; Ezcurra et al. presence and shape of the anterolateral scar is affected by
2020a). Ankle anatomy has been historically used to clas- intraspecific variation (Piechowski et al. 2014; Griffin &
sify Pseudosuchia and Pan-Aves (Cruickshank 1979), and Nesbitt 2016; M€uller 2022). Therefore, the presence/
the presence and shape of trochanters, condyles and other absence of this trait must be carefully considered.
features from the zeugopodium and stylopodium have The occurrence of the iliofibularis tubercle on the fib-
supported internal relationships among these clades ula of Stenoscelida aurantiacus is another unusual trait
(Nesbitt 2011; Cabreira et al. 2016; Baron et al. 2017; for Proterochampsia. Nevertheless, this feature is widely
Ezcurra et al. 2020a). For instance, the anterior (¼ lesser) distributed within crocopodans (Ezcurra et al. 2020a),
trochanter is exhaustively reported for the femur of dino- and it is not regarded as a traditional synapomorphy of
saurs and related groups (Hutchinson 2001; Pintore et al. any exclusive clade (e.g. Nesbitt 2011; Ezcurra 2016).
2021). It is treated as the scar of the M. iliofemoralis Similarly, the presence of a vestigial phalanx in digit V
(Hutchinson 2001). The shape of the anterior trochanter comprises another peculiar condition among the mem-
varies within the main lineages of dinosaurs (Pintore bers of the clade. Yet, the presence/absence of pha-
et al. 2021), providing data for phylogenetic analyses langes in digit V, which is often vestigial, is highly
(Nesbitt 2011; Langer & Ferigolo 2013; Baron et al. affected by taphonomic biases.
2017). Yet, the occurrence of the structure is not
restricted to the pan-avian lineage, since the anterior tro-
chanter is reported for crocodylomorphs and ornithosu- Local palaeofauna
chids (Clark & Sues 2002; Baczko et al. 2019; M€uller The Varzea do Agudo (¼ Janner) site is one of the most
et al. 2020a), which are pseudosuchians. So far, there are prolific fossiliferous localities from southern Brazil,
 A new proterochampsid from Brazil 15

from which year-after-year numerous new specimens proterochampsids probably had a mode of life related to
have been exhumed. This fauna is dominated by traver- water bodies, other were more likely terrestrial in habit
sodontid cynodonts of the genus Exaeretodon (M€uller (e.g. Chanaresuchus bonapartei; Arcucci et al. 2019;
et al. 2020b). This remarkable abundance has been used Ezcurra et al. 2021).
to correlate these beds with those from the upper portion
of the Ischigualasto Formation (Langer et al. 2007;
Ezcurra et al. 2015; M€uller & Garcia 2020; Schultz Conclusions
et al. 2020). As a consequence, the younger Brazilian
proterochampsids are divided into two distinct horizons Stenoscelida aurantiacus was erected based on the new
within the Hyperodapedon AZ. Rhadinosuchus gracilis specimen CAPPA/UFSM 0293, composed of a nearly
and Cerritosaurus binsfeldi were unearthed from out- complete right hind limb. Its phylogenetic position
crops of the Alemoa complex (Garcia et al. 2019), within Proterochampsidae is uncertain; however, our
which belong to the Hyperodapedon Acme Zone (lower results support the new species as a non-rhadinosuchine
portion; Ezcurra et al. 2015; Schultz et al. 2020),
proterochampsid. The discovery of a complete and rela-
whereas Proterochampsa nodosa comes from
tively well-preserved hind limb of a proterochampsid is
Exaeretodon-dominated beds (upper portion; Schultz
of great interest because the fossil record of this skeletal
et al. 2020). In this context, Stenoscelida aurantiacus
portion is poorly preserved, especially from the mid-
expands the abundance and diversity of proterochamp-
dle–late Carnian to early Norian. Although the new
sids in the Exaeretodon sub-AZ.
taxon displays traits that unequivocally place it in
The presence of a proterochampsid in the Varzea do
Proterochampsidae, Stenoscelida aurantiacus carries a
Agudo site is not particularly helpful regarding a bio-
stratigraphic approach because proterochampsids occur set of unusual hind limb traits within proterochampsids,
across the entire Ischigualasto Formation (Martınez such as the presence of an anterior trochanter and a
et al. 2013). Nevertheless, the new taxon comprises an dorsolateral scar on the femoral head, an iliofibularis
additional predatorial taxon for the palaeofauna of this tubercle on the anterior margin of the fibular shaft, and
site. The absence of cranial remains precludes an unam- a vestigial phalanx in digit V.
biguous interpretation regarding the diet of Stenoscelida Proterochampsids are rare in the Late Triassic sedi-
aurantiacus, but since all proterochampsids are consid- ments of Brazil when compared to those of Argentina.
ered carnivorous animals (Trotteyn et al. 2013), the null Stenoscelida aurantiacus is the first proterochampsid
hypothesis is that Stenoscelida aurantiacus exhibited a from the Varzea do Agudo site, expanding the taxo-
similar diet. So far, Stenoscelida aurantiacus and nomic and ecological diversity of this iconic fossilifer-
Dynamosuchus collisensis comprise the only carnivorous ous locality. The investigation and understanding of the
archosauriforms from this site, with the ectetniniid cyno- Carnian biota is crucial because this moment witnessed
dont Trucidocynodon riograndensis (Oliveira et al. the rise of dinosaurs (Novas et al. 2021) and the dawn
2010) adding to this context. Other occurrences at the of modern ecosystems (Dal Corso et al. 2020).
site belong to herbivorous/omnivorous animals: the
archosauromorph Hyperodapedon sp., the traversodontid
cynodont Exaeretodon, and the sauropodomorph dino- Acknowledgements
saurs Pampadromaeus barberenai (Cabreira et al. 2011)
and Bagualosaurus agudoensis (Pretto et al. 2019). We thank Martin D. Ezcurra for photographs of some
The putative mode of life related to water bodies proterochampsids. We thank the Willi Hennig Society
often attributed to proterochampsids are an interesting for supporting the free use of TNT software. We extend
theme added to the Varzea do Agudo palaeofauna, our gratitude to Martin D. Ezcurra and an anonymous
which previously recorded only fully terrestrial taxa (see reviewer for corrections and comments that greatly
above). Currently, the site lacks other organisms that improved this manuscript. We are grateful to Caio
possibly preyed within aquatic environments, such as Fantini for the digital reconstruction of Stenoscelida
temnospondyls and phytosaurs. Therefore, if protero- aurantiacus depicted in Figure 7. This work was carried
champsids (especially Stenoscelida aurantiacus) were out with aid of the Fundaç~ao de Amparo a Pesquisa do
semi-aquatic predators (following the general morph- Estado do Rio Grande do Sul (FAPERGS 21/2551-
ology of extant crocodilians; Trotteyn et al. 2013), the 0000680-3) and the Conselho Nacional de
new taxon expands the ecological diversity within the Desenvolvimento Cientıfico e Tecnologico (CNPq
Varzea do Agudo site. On the other hand, some methods 404095/2021-6) to Rodrigo Temp M€uller, and a
have cast doubts on these habits. Therefore, while some Coordenaç~ao de Aperfeiçoamento de Pessoal de Nıvel
16 R. T. M€uller et al.

Superior (CAPES 88887.608076/2021-00) scholarship Triassic of Austria. Zoological Journal of the Linnean
for Mauricio Silva Garcia. Society, 187, 198–228.
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)
Supplemental material from the Triassic of Brazil. Naturwissenschaften, 98,
1035–1040.
Supplemental material for this article can be accessed Cabreira, S. F., Kellner, A. W. A., Dias-da-Silva, S.,
Roberto-da-Silva, L., Bronzati, M., Marsola, J. C.,
here: https://doi.org/10.1080/14772019.2022.2128913. M€ uller, R. T., Bittencourt, J. S., Batista, B. J.,
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Rodrigo Temp M€uller http://orcid.org/0000-0001- crocodylomorph archosaurs from the Lower Jurassic and
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