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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 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

Proterochampsids are non-archosaurian archosauriforms
that are endemic to South America (Trotteyn et al.
2013; Ezcurra et al. 2015, 2021). Characterized by an
elongated snout, members of the clade were usually
regarded as semi-aquatic predators according to general
similarities with crocodilians (Reig 1959; Romer 1971;
Bonaparte 1978; Trotteyn et al. 2013). However, more
recently, palaeohistological data (Arcucci et al. 2019)
and morphofunctional studies (Ezcurra et al. 2021) have
indicated more terrestrial habits for at least some mem-
bers of this clade. The fossil record of proterochampsids
ranges from the early Carnian to the late Carnian/early
Norian (Ezcurra et al. 2017) and is restricted to
Argentina and Brazil (Ezcurra et al. 2015). The oldest
proterochampsids have been unearthed from the
Massetognathus-Chanaresuchus Assemblage Zone (AZ)
of the Cha~ nares Formation, Argentina (Romer 1971;

Corresponding author. Email: rodrigotmuller@hotmail.com
Santacruzodon AZ (Raugust et al. 2013) of Brazil; how-
ever, the fossil record of proterochampsids from the
early Carnian beds of Brazil is far less abundant than
the Argentinean record. The younger members of the
clade are excavated from the middle to late Carnian/
early Norian beds of the Ischigualasto Formation
(Argentina: Sill 1967; Trotteyn et al. 2012) and the
Candelaria Sequence (Brazil: Huene 1938; Price 1946;
Barberena 1982).
Perhaps one of the most interesting matters regarding
Proterochampsidae is its phylogenetic position. The
most comprehensive studies regarding archosauromorph
affinities have recovered proterochampsids close to
Archosauria (Arcucci 1989; Sereno & Arcucci 1990;
Sereno 1991; Ezcurra 2016; Ezcurra et al. 2017, 2020a;
M€uller et al. 2020a; Trotteyn & Ezcurra 2020; Kellner
et al. 2022). Therefore, Proterochampsidae is a key
clade for investigating the origin and distribution of

# 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
are usually poorly preserved for the group. This portion
of the skeleton holds traits usually regarded as diagnos-
tic for a number of archosaurian clades (Nesbitt 2011;
Ezcurra 2016; Baron et al. 2017). In the present study,
we describe a new proterochampsid from the Late
Triassic of Brazil composed of a complete and articu-
lated hind limb. The new taxon is the first protero-
champsid from the Varzea do Agudo site and provides
insight on the distribution of some classical archosaurian
traits.
Material and methods
Institutional abbreviations
BNM, B€ undner Naturmuseum, Chur, Switzerland; CA,
Colegio Anchieta, Porto Alegre, Rio Grande do Sul,
Brazil; CAPPA/UFSM, Centro de Apoio a Pesquisa
Paleontol ogica da Quarta Col^onia da Universidade
Federal de Santa Maria, Rio Grande do Sul, Brazil;
CPEZ, Paleontology Collection of the Museu
Paleontol ogico e Arqueologico Walter Ilha, S~ao Pedro
do Sul, Brazil; CRILAR-Pv, Paleontologıa de
Vertebrados, Centro Regional de Investigaciones
Cientıficas y Transferencia Tecnologica, Anillaco,
Argentina; FC-DPV, Coleccion de Vertebrados Fosiles,
Departamento de Paleontologıa, Facultad de Ciencias,
Universidad de la Republica, Montevideo, Uruguay;
GR, Ruth Hall Museum of Paleontology at Ghost
Ranch, New Mexico, USA; ISI, Indian Statistical
Institute, Kolkata, India; NHMUK, Natural History
Museum, London, UK; NHMW, Naturhistorisches
Museum Wien, Vienna, Austria; NMQR, National
Museum, Bloemfontein, South Africa; NMT, National
Museum of Tanzania, Dar es Salaam, Tanzania;
PIMUZ, Pal€aontologisches Institut und Museum,
Universit€at Z€urich, Z€urich, Switzerland; PIN, Borissiak
Paleontological Institute of the Russian Academy of
Sciences, Moscow, Russia; PULR, Paleontologıa,
Museo de Ciencias Naturales, Universidad Nacional de
La Rioja, La Rioja, Argentina; PVSJ, Division of
Vertebrate Paleontology of the Museo de Ciencias
Naturales de la Universidade Nacional de San Juan, San
Juan, Argentina; SNSB-BSPG, Staatliche
Naturwissenschaftliche Sammlungen Bayerns-
Bayerische Staatssammlung f€ur Pal€aontologie und
Geologie, Munich, Germany; TMM, Jackson School of
Geosciences Vertebrate Laboratory, University of Texas
at Austin, Austin, Texas, USA; UFRGS,
Paleovertebrate Collection of the Laboratorio de
Paleovertebrados da Universidade Federal do Rio
Grande do Sul, Porto Alegre, Brazil; UFSM,
Laboratorio de Estratigrafia e Paleobiologia,
Universidade Federal de Santa Maria, Brazil; ULBRA,
Universidade Luterana do Brasil, Coleç~ao de
Paleovertebrados, Canoas, Brazil; UWBM, Burke
Museum of Natural History and Culture, Seattle,
Washington, USA.
Material
The specimen is housed in the palaeovertebrates collec-
tion of the Centro de Apoio a Pesquisa Paleontologica
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.
Terminology
This study employs traditional (Romerian) anatomical
terms (i.e. ‘anterior’ and ‘posterior’). The nomenclature
for femoral tubers follows Ezcurra (2016), whereas the
nomenclature for muscle attachment structures follows
Griffin & Nesbitt (2016).
Phylogenetic analysis
The phylogenetic affinities of the new taxon were inves-
tigated employing the data matrix of Ezcurra & Sues
(2021). We rescored character 502, regarding the pres-
ence and shape of the anterior trochanter (¼ lesser or
minor trochanter; iliofemoralis cranialis muscle inser-
tion) of the femur, from the state ‘absent (0)’ to ‘present
and forms a steep margin with the shaft but is com-
pletely connected to it (1)’ in Gualosuchus reigi (see
Description, below, for further information). The ana-
lysis was conducted in TNT v. 1.5 (Goloboff &
Catalano 2016). Following Ezcurra & Sues (2021), 40
operational taxonomic units (ISIR 1132,
Protanystropheus antiquus, Trachelosaurus fischeri,
Tanystropheus haasi, UFRGSPV- 492-T, Malerisaurus
all NA, Arctosaurus osborni, CRILAR-Pv 461,
CRILAR-Pv 462, CRILAR-Pv 497, Cha~nares rhyncho-
saur, PVSJ 2728, Eorasaurus olsoni, Archosaurus com-
plete, FC-DPV 2641, UFSM 11444, UFSM 11394,
Vonhuenia fredericki, C. rossicus combined, C. magnus
combined, Chasmatosuchus vjushkovi, Koilamasuchus
gonzalezdiazi, Kalisuchus rewanensis holotype, NMQR
3570, CRILAR-Pv 499, Shansisuchus kuyeheensis,
Uralosaurus combined, Osmolskina czatkoviensis,
Osmolskina complete, Triopticus primus, Otter
Sandstone archosaur, Dagasuchus santacruzensis,
Ctenosauriscus koeneni, Hypselorhachis mirabilis,
Waldhaus poposauroid, Vytshegdosuchus zbeshartensis,
Bystrowisuchus flerovi, Bromsgroveia walkeri,
Lutungutali sitwensis and Nyasasaurus parringtoni) and
two characters (9 and 119) were set as deactivated. The
characters were equally weighted and characters 1, 2, 7,
 A new proterochampsid from Brazil
10, 17, 19–21, 28, 29, 34, 36, 40, 42, 46, 50, 54, 66,
71, 74–76, 100, 122, 127, 146, 153, 156, 157, 171, 176,
177, 187, 202, 221, 227, 263, 266, 278, 279, 283, 324,
327, 331, 337, 342, 345, 351, 352, 354, 361, 365, 368,
370, 377, 379, 386, 387, 398, 410, 414, 424, 425, 430,
435, 446, 448, 454, 455, 458, 460, 463, 464, 470, 472,
478, 482, 483, 485, 489, 490, 502, 504, 510, 516, 520,
521, 529, 537, 546, 552, 556, 557, 567, 569, 571, 574,
581, 582, 588, 636, 648, 652, 662, 701, 731, 735, 737,
738, 743, 749, 766, 784, 803, 810, 816, 850, 851, 872,
875, 885 and 888 were treated as ordered (additive).
The most parsimonious trees were recovered from an
equally weighted parsimony analysis following the
search protocol by Ezcurra et al. (2020a): search
employing new technology search algorithms until 100
optimal hits are reached, followed by a final round of
TBR branch swapping. Consistency and retention indi-
ces were calculated with the script developed by
Spiekman et al. (2021) that does not take into account a
priori deactivated terminals.
Systematic palaeontology
Archosauromorpha Huene, 1946 sensu Dilkes (1998)
Archosauriformes Gauthier, Kluge & Rowe, 1988
sensu Gauthier et al. (1988)
Proterochampsia Bonaparte, 1971 sensu
Kischlat (2000)
Proterochampsidae Sill, 1967 sensu Trotteyn (2011)
Stenoscelida aurantiacus gen. et sp. nov.
(Figs 1–5)
Holotype. CAPPA/UFSM 0293, a complete and
articulated right hind limb.
Etymology. The genus combines the Greek words
rsemό1 (¼ narrow) and rjeko1 (¼ hind leg), referring
to the slender leg of the creature. The specific epithet
derives from the Latin word aurantiacus (¼ orange), in
allusion to the orange colour of the outcropping sedi-
ments of the Varzea do Agudo site (Fig. 1B).
Type locality. Varzea do Agudo site (¼ Janner site;
29 390 10.8900 S, 53 170 34.2000 W), Agudo, Rio Grande do
Sul, Brazil (Fig. 1).
Stratigraphic horizon. Lower portion of the Candelaria
Sequence (Horn et al. 2014) of the Santa Maria
Supersequence (Zerfass et al. 2003), Parana Basin. The
predominance of the cynodont genus Exaeretodon places
the site in the upper part (Exaeretodon sub-Assemblage
Zone; M€ uller & Garcia 2020) of the Hyperodapedon
Assemblage Zone (Schultz et al. 2020), which is
3
biostratigraphically correlated with the Exaeretodon bio-
zone from the Ischigualasto Formation (Martınez et al.
2013). A Bayesian age-model for the profile of the
Ischigualasto Formation at the Hoyada del Cerro Las
Lajas locality in Argentina recovered an age of
227.94 þ 0.83/1.67 for the top of the Hyperodapedon
AZ (Desojo et al. 2020), indicating an age of c. 228 Ma
for the Exaeretodon biozone. A similar age (late
Carnian/early Norian, Late Triassic) for the Exaeretodon
sub-Assemblage Zone is inferred in the
Candelaria Sequence.
Ontogenetic assessment. The presence of some muscle
attachment structures (e.g. anterior trochanter; anterolat-
eral scar) in the femur of CAPPA/UFSM 0293 suggests
some degree of development if the ontogenetic pathways
of other archosauriforms are considered (Griffin &
Nesbitt 2016). However, it is currently not possible to
confirm whether the specimen reached its maximum
size or not.
Diagnosis. Stenoscelida aurantiacus differs from all
other known proterochampsids in (
local autapomor-
phies): possessing a slender femur; presence of an anter-
ior trochanter on the femoral head; presence of a raised
anterolateral scar above the anterior trochanter; fourth
trochanter restricted to the proximal half of the femur;
tibia approximately 84% the total length of the femur;
in proximal view, the cnemial crest of the tibia is sub-
equal in size to the proximal condyles; fibula with an
iliofibularis tubercle on the proximal portion of the
shaft
; proximal third of the fibular shaft is anteropos-
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-
tarsal III is 2.5; and digit V with a phalanx
.
Description and comparison
Overview
The preserved right hind limb of the holotype is entirely
articulated (Fig. 2). Some portions of the specimen are
covered by a thick concretion layer, especially the femoral
midshaft and the ankle. The external surface is well-
preserved in several places, revealing some muscle attach-
ment points. However, the specimen was deformed by
sedimentary compression. Therefore, the elements are lat-
eromedially compressed, affecting the original shape. As
result, the midshaft of the limb bones is collapsed, show-
ing artificial longitudinal grooves or sulci. Part of the fibu-
lar midshaft is not preserved. The same is true for the
distal portion of metatarsal IV, precluding us from deter-
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.
Diagenetic processes might have exaggerated this condi-
tion. The general morphology of the femoral head
resembles that of proterochampsids, whereas it differs
from the anteromedially expanded femoral head of dino-
saurs (Nesbitt 2011) and from the hook-shaped head of
lagerpetids and pterosaurs (Ezcurra et al. 2020a). The
proximal articular surface bears a shallow straight
groove (Fig. 3C), which is usually absent in protero-
champsids, except in an unnamed rhadinosuchine
(Ezcurra et al. 2019; CRILAR-Pv 491) from the
Cha~ nares Formation. There is an anterior tuber (sensu
Ezcurra 2016) on the proximal portion of the femur, and
although the bone lacks a posteromedial tuber, the pos-
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
dorsolateral trochanter on the proximal portion of the
bone. Conversely, the anterolateral surface bears a raised
and rugose area (Fig. 3B) where the anterior (¼ lesser)
trochanter is reported for a number of archosauriforms
(e.g. ornithosuchids, aetosaurs, dinosauriforms).
Whereas the anterior trochanter is absent in several pro-
terochampsids (Trotteyn et al. 2013), the structure
occurs in the holotype of Gualosuchus reigi (PULR-V
05; M. D. Ezcurra, pers. comm.). A trochanteric shelf is
not associated with the anterior trochanter, and the prox-
imal-most portion of this trochanter is completely con-
nected to the shaft. Slightly above the proximal tip of
the anterior trochanter there is an additional scar, which
resembles the anterolateral scar (sensu Griffin & Nesbitt
2016) of aphanosaurs and dinosauriforms. There is a
similar scar in Gualosuchus reigi (PULR-V 05; M. D.
Ezcurra, pers. comm.).
The transition from the femoral head to the shaft is
smooth. The fourth trochanter rests on the posterior surface
of the proximal third of the femur (Fig. 3A). It is crest-like
and symmetrical in shape. Its medial surface is ornamented
with muscle scars. The distal portion of the fourth trochan-
ter does not extend further down along the femoral shaft.
In contrast, the fourth trochanter is strongly proximodistally
developed in Gualosuchus reigi and Chanaresuchus bona-
partei (Ezcurra et al. 2019). The midshaft is slender. The
robustness index (RI, sensu Wilson & Upchurch 2003; i.e.
average of the greatest widths of the proximal end, mid-
shaft and distal end of the element divided by the length
of the element) is 0.14. For comparison, this is slightly
slenderer than that of some early dinosaurs, such as
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.
Gnathovorax cabreirai (0.16) and Buriolestes schultzi
(0.15; M€uller & Garcia 2022).
The anterior surface of the distal portion bears an
extensor fossa (Fig. 3D), which results in a concave
anterior margin in distal view. On the opposite side, the
popliteal fossa is well delimited (Fig. 3E). It is proximo-
distally short, approximately 10% of the total length of
the bone. This condition distinguishes the specimen
from silesaurids and aphanosaurs, where the fossa is
considerably longer (Nesbitt et al. 2010, 2017). The dis-
tal condyles are approximately at the same level in
anterior or posterior views. In addition, the lateral sur-
face between the lateral condyle and the crista tibiofibu-
laris is smooth, lacking a deep groove.
Tibia
The tibia (Fig. 4) is 84% the total length of the femur
(Table 1). This resembles the condition in
Pseudochampsa ischigualastensis, where it is approxi-
mately 82% (Trotteyn & Ezcurra 2014), and differs from
both Proterochampsa barrionuevoi (74.5%; Trotteyn
2011) and Tropidosuchus romeri (100%; Arcucci 1990).
The bone is straight in anterior/posterior or medial/lateral
views. The proximal portion is strongly expanded and is
triangular in shape in proximal view (Fig. 4C). The anter-
ior half of the proximal margin is proximally projected in
medial view. In medial or lateral views, the well-devel-
oped cnemial crest extends anteriorly, and in proximal
view it is straight and the anterior margin is rounded. The
cnemial crest is sub-equal in size regarding the proximal
condyles, distinguishing the new species from
Proterochampsa barrionuevoi, where the crest is propor-
tionally smaller. The posterior and lateral condyles are
sub-equal in size. The lateral condyle is offset regarding
the posterior condyle. The posterior condyle does not
taper posteriorly. The presence of any crest on the lateral
surface of the proximal portion of the bone is uncertain.
The midshaft is ovoid in cross-section. The bone wall
is thick, distinct from the thin walls of several pan-avi-
ans and some crocodylomorphs (Kellner et al. 2022).
The distal portion of the bone is moderately expanded.
It lacks a posterolateral process. There is a proximodis-
tally oriented groove on the lateral surface of the distal
portion (Fig. 4F). It is more pronounced in
Proterochampsa barrionuevoi (Trotteyn 2011). The dis-
tal outline of the tibia is elliptical. It is anteroposteriorly
longer than transversely wide.
 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
because a portion of the midshaft is not preserved. The
bone is straight in anterior or lateral views, and the
proximal articular surface is anteroposteriorly expanded
and transversely compressed (Fig. 4C). It differs from
the rounded to elliptical proximal end of some pseudo-
suchians (e.g. Dynamosuchus collisensis, Prestosuchus
chiniquensis). In proximal view, the lateral margin is
convex but the medial margin is concave. In lateral
view, the posterior margin projects posteriorly, whereas
the anterior margin lacks any anterior expansion.
Conversely, in non-proterochampsid proterochampsians
(e.g. Vancleavea campi; Litorosuchus somnii;
Jaxtasuchus salomoni) the proximal portion is symmet-
rical to nearly symmetrical in lateral view. The posterior
margin of the proximal third of the bone is sharp.
expanded, being wider than the tibia (Fig. 4F).
Conversely, the shaft becomes extremely slender for the
next two-thirds, being approximately two times narrower
than the tibia. This is an unusual condition and distin-
guishes the specimen from other proterochampsids.
Furthermore, there is an iliofibularis tubercle on the prox-
imal portion of the shaft (Fig. 4E, F), a feature absent in
other proterochampsids. The condition in Stenoscelida
aurantiacus differs from the more distally located tuber-
cle of several pseudosuchians and rhynchosaurs.
The distal portion of the fibula is gently expanded. It
is approximately 0.45 times the maximum length of the
proximal articular surface (Table 1). There is a faint lon-
gitudinal ridge running on the anterolateral margin. The
distal articular surface is flat to concave. In distal view
(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
2, 5). The astragalus is transversely wider (20.5 mm) than
anteroposteriorly long (11 mm). The anterior margin of
the astragalus is concave in dorsal view (Figs 2B, 5B). A
transverse groove runs on the posterior surface of the
bone. The presence or absence of foramina on this pos-
terior groove is uncertain because it is badly preserved.
As in other proterochampsids, the tibial facet is wider
than the fibular facet. The latter facet is dorsolaterally
oriented, resembling the condition of Chanaresuchus
bonapartei and Proterochampsa barrionuevoi, whereas in
Pseudochampsa ischigualastensis it is dorsally oriented
(Trotteyn & Ezcurra 2014). The ventral surface of the
bone is transversely convex. The calcaneum is 10 mm in
width (approximately half of the astragalar width). There
is a posterolaterally oriented calcaneal tuber. This struc-
ture is sub-rectangular, with a straight lateral margin. In
dorsal view, the posterolateral corner of the calcaneal
tuber is rounded (Fig. 5B).
As in other proterochampsids, the metatarsals (Fig. 5)
overlap each other. Metatarsal I is shorter than metatar-
sals II–IV (Fig. 5C). It is almost three times shorter
than metatarsal III (Table 1). The proximal articular sur-
face is transversely expanded. There is a longitudinal
smooth crest on the dorsomedial corner of the proximal
half of the bone. The distal end is moderately expanded.
The lateral condyle is more pronounced dorsally than
the medial one. There is a shallow extensor fossa on the
dorsal surface of the distal portion. The presence of col-
lateral ligament pits on the condylar sides is uncertain.
The distal articular surface is not strongly ginglymoid.
Metatarsal II is the stoutest metatarsal. The proximal
articular surface is strongly expanded (Table 1). The
ratio between the proximal articular surface and the total
length is 0.39. It is 0.36 in Rhadinosuchus gracilis
(Ezcurra et al. 2015). Conversely, the shaft is stouter in
Rhadinosuchus gracilis, where the ratio between the
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
clearly the largest ungual. It is transversely compressed
and tapers to a sharp point. The ungual is not ventrally
recurved and lacks a well-developed flexor tubercle.
Digit III is the longest (Fig. 5B). The first phalanx from
digit III is sub-equal in length to the first phalanx of
digit II (Table 2). On the other hand, the shaft of this
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
remarkably smaller than that of digit II. The number of
phalanges in digit IV is unknown and the distal portion
of metatarsal IV is not preserved. Therefore, the absence
of phalanges is ambiguous. The phalanx of digit V (Fig.
5A) is reduced to an elongated structure with a concave
proximal articular surface. No condyles or other struc-
tures are present in this vestigial element.
Phylogenetic results
The phylogenetic analysis recovered 39,200 most parsi-
monious trees (MPTs) of 6287 steps (consistency index
¼ 0.18375; retention index ¼ 0.63457). Stenoscelida aur-
antiacus nests within Proterochampsidae in all MPTs
(Fig. 6A). The clade is supported by 19 synapomorphies,
of which three are coded for Stenoscelida aurantiacus: 1)
midshaft diameter of metatarsal II is more than the mid-
shaft diameter of metatarsal I (ch. 572: 0!1); 2) mid-
shaft diameter of metatarsal IV is lower than that of
metatarsal III (ch. 573: 0!1); and 3) crest-like fourth tro-
chanter, dorsoventrally/anteroposteriorly taller than or
equal to the shaft at its minimum depth (ch. 803: 1!2).
Proterochampsidae is the sister group to Doswelliidae,
which is composed of Rugarhynchus sixmilensis,
Sphodrosaurus pennsylvanicus, Doswellia kaltenbachi
and Jaxtasuchus salomoni. The clade supporting
Proterochampsidae þ Doswelliidae is in a trichotomy with
Polymorphodon adorfi and an unnamed clade composed
of Vancleavea campi and Litorosuchus somnii. These
clades are included within Proterochampsia.
Regarding the internal relationships of
Proterochampsidae, there is a polytomy composed of
Stenoscelida aurantiacus, Tropidosuchus romeri,
Cerritosaurus binsfeldi, Gualosuchus reigi, both species
of Proterochampsa, and Rhadinosuchinae in the strict
consensus tree (Fig. 6A). Given the absence of overlap-
ping elements between the new taxon and several other
proterochampsids (e.g. Proterochampsa nodosa,
Cerritosaurus binsfeldi), these unresolved affinities are
unsurprising. The new taxon, however, does not nest
within Rhadinosuchinae in any of the MPTs, thus
Stenoscelida aurantiacus is a non-rhadinosuchine proter-
ochampsid. The clade Rhadinosuchinae is composed of
11
a trichotomy including Rhadinosuchus gracilis,
Pseudochampsa ischigualastensis and Chanaresuchus
bonapartei. Except for the polytomy at the base of
Proterochampsidae, the topology of the strict consensus
tree is identical to that recovered by Ezcurra &
Sues (2021).
Discussion
Taxonomic status
In addition to a series of cranial features, proterochamp-
sids are characterized by a metatarsal II with a robust
midshaft, whereas metatarsal IV has a slender profile
(Trotteyn et al. 2013). The new taxon possesses this
particular combination of traits. Therefore, Stenoscelida
aurantiacus is unequivocally assigned to
Proterochampsidae due to the shape of metatarsals.
Moreover, this is supported by our phylogenetic ana-
lysis. Unfortunately, the absence of overlapping ele-
ments between Stenoscelida aurantiacus and several
other proterochampsids (especially the Brazilian forms)
precludes a clear assessment of its specific phylogenetic
relationships. Rhadinosuchus gracilis is the only
Brazilian proterochampsid from the Hyperodapedon AZ
that preserves any hind limb elements. Actually, the
holotype preserves a left metatarsal II (Ezcurra et al.
2015). It is the only comparable element between both
taxa. The latter is a member of Rhadinosuchinae
(Ezcurra et al. 2015), a less inclusive clade within
Proterochampsidae, a nesting reinforced by our results
(see above). On the other hand, Stenoscelida aurantia-
cus is not recovered as a rhadinosuchine in our analysis,
and despite the few comparable elements, there are
some differences between the metatarsal II of CAPPA/
UFSM 0293 (Stenoscelida aurantiacus) and SNSB-
BSPG AS XXV 50 (Rhadinosuchus gracilis). In SNSB-
BSPG AS XXV 50 the midshaft is stouter and the prox-
imal articular surface is proportionally smaller than in
CAPPA/UFSM 0293. The medial and lateral condyles
of SNSB-BSPG AS XXV 50 are uneven distally,
whereas in CAPPA/UFSM 0293 both condyles are in
the same plane. A more detailed comparison demands
additional overlapping elements. In sum, CAPPA/UFSM
0293 cannot be ascribed to Rhadinosuchus gracilis
according to the lack of anatomical correspondence and
distinct phylogenetic positions.
Although the holotype (CA unnumbered) of
Cerritosaurus binsfeldi preserves some postcranial ele-
ments, there are no hind limb bones (Price 1946;
Trotteyn et al. 2013). Despite the absence of overlap-
ping bones between Cerritosaurus binsfeldi and
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
ischigualastensis, 25.4 cm and 15.48 cm (Trotteyn &
Ezcurra 2014); and Tropidosuchus romeri, 7.67 cm and
6.46 cm (Bestwick et al. 2022). Based on these measure-
ments, the estimated femoral length of Cerritosaurus
binsfeldi is 7.54 cm (y ¼ 0.6262x þ 0.65702; Fig. 7),
which is approximately two times shorter than the fem-
oral length of Stenoscelida aurantiacus (i.e. 14.7 cm).
So far, there are no ontogenetic assessments regarding
Cerritosaurus binsfeldi. Therefore, additional studies or
new fossils are necessary in order to determine if the
size discrepancy between both species is an effect of
ontogeny or another intraspecific factor. Nevertheless,
geological and stratigraphical evidence provides further
support for our taxonomic proposal. Cerritosaurus bins-
feldi was excavated from strata ascribed to the
Hyperodapedon Acme Zone (Langer et al. 2007), which
are older than that which yielded Stenoscelida aurantia-
cus (i.e. Exaeretodon sub-AZ; Langer et al. 2007;
Martınez et al. 2013; Desojo et al. 2020; M€uller &
Garcia 2020; Schultz et al. 2020).
In relation to the occurrence of Proterochampsa
nodosa in sediments attributable to the Exaeretodon
sub-AZ, Stenoscelida aurantiacus does not present over-
lapping elements with the former, which is known from
a single skull with mandible in occlusion (Barberena
1982; Trotteyn et al. 2013; Sim~ao-Oliveira et al. 2022).
Although there are no comparable elements between
both Brazilian specimens, it is possible to compare
Stenoscelida aurantiacus with the Argentinian species
of the same genus: Proterochampsa barrionuevoi. The
latter is known from several specimens, including an
almost complete skeleton (PVSJ 606), and its hind limb
morphology differs substantially from that in
Stenoscelida aurantiacus, not to mention the three local
apomorphies of the new taxon (see Diagnosis, above),
which will not be repeated here. In relation to the
femur, the popliteal fossa of CAPPA/UFSM 0293
(Stenoscelida aurantiacus) is short, comprising 10% of
the total femoral length, but in PVSJ 606
(Proterochampsa barrionuevoi; Trotteyn 2011), it is lon-
ger in comparison, reaching roughly 23% of the total
femoral length. Furthermore, based on femoral measure-
ments, CAPPA/UFSM 0293 is 22% smaller than PVSJ
606, and although it is possible to argue that the speci-
men probably does not represent an immature individual
due to the presence of well-developed femoral epiphyses
and scars (see above), we cannot determine with the evi-
dence at hand if CAPPA/UFSM 0293 reached its max-
imum size. The proportions of the tibia and femur also
differ between both specimens, where in CAPPA/UFSM
0293 the tibia is proportionally longer than in PVSJ
14 R. T. M€uller et al.
606; although we recognize that the size difference
between specimens could possibly influence this trait,
there is no evidence to support this sort of allometric
discrepancy in proterochampsids. Therefore, as in the
case of Cerrritosaurus bindfeldi, more specimens are
needed in order to test this assumption. Regarding the
tibia, in CAPPA/UFSM 0293 the proximal extremity of
the tibia is perpendicular in relation to its long axis,
whereas in PVSJ 606 it is oblique. Moreover, in PVSJ
606, the cnemial crest is smaller than the proximal con-
dyles (Trotteyn 2011), whereas in CAPPA/UFSM 0293
it is sub-equal in size to the proximal condyles. The fib-
ula of PVSJ 606 maintains its anteroposterior width for
most of its proximodistal length, as in other protero-
champsids (although the epiphyses are slightly
expanded), but in CAPPA/UFSM 0293 the fibula is sig-
nificantly expanded in the proximal third of the element
and then tapers distally, becoming remarkably narrow,
ending in an unexpanded distal epiphysis. In sum, we
consider it unlikely that CAPPA/UFSM 0293 represents
a new specimen of the genus Proterochampsa.
Distribution of hind limb traits
The occurrence of a new species of proterochampsid with
a well-preserved hind limb is particularly exciting.
According to several authors (e.g. Sereno 1991; Ezcurra
2016; Ezcurra et al. 2020a; Trotteyn & Ezcurra 2020;
Kellner et al. 2022), Proterochampsia is regarded as the
sister group to Archosauria. As a result, the anatomy of
proterochampsians is crucial in order to optimize the ori-
gin and distribution of morphological traits usually pre-
sent in the earliest diverging clades of archosaurs.
Several of these traits rely on the pelvic and hind limb
anatomy (Nesbitt 2011; Ezcurra 2016; Ezcurra et al.
2020a). Ankle anatomy has been historically used to clas-
sify Pseudosuchia and Pan-Aves (Cruickshank 1979), and
the presence and shape of trochanters, condyles and other
features from the zeugopodium and stylopodium have
supported internal relationships among these clades
(Nesbitt 2011; Cabreira et al. 2016; Baron et al. 2017;
Ezcurra et al. 2020a). For instance, the anterior (¼ lesser)
trochanter is exhaustively reported for the femur of dino-
saurs and related groups (Hutchinson 2001; Pintore et al.
2021). It is treated as the scar of the M. iliofemoralis
(Hutchinson 2001). The shape of the anterior trochanter
varies within the main lineages of dinosaurs (Pintore
et al. 2021), providing data for phylogenetic analyses
(Nesbitt 2011; Langer & Ferigolo 2013; Baron et al.
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-
chids (Clark & Sues 2002; Baczko et al. 2019; M€uller
et al. 2020a), which are pseudosuchians. So far, there are
no records of the anterior trochanter for non-archosaurian
archosaurs (to our knowledge). Stenoscelida aurantiacus
(together with Gualosuchus reigi) provides the first evi-
dence of the presence of this feature in a non-archosau-
rian archosauriform (Fig. 6B). The occurrence of the
anterior trochanter in a proterochampsid supports a
deeper origin for this trait than previously thought. It is
possible that instead of evolving independently in the two
main lineages of Archosauria, the anterior trochanter ori-
ginated earlier and was retained in the major clades of
Pan-Aves and in some members of Pseudosuchia.
However, the absence of this trait in the proterochamp-
sian Vancleavea campi cast doubts on this hypothesis and
favours a scenario where the anterior trochanter evolved
independently in proterochampsids and archosaurs.
Above the anterior trochanter, the femur of
Stenoscelida aurantiacus displays another interesting fea-
ture. It comprises a raised surface that resembles the
anterolateral scar (¼ ‘dorsolateral ossification’ of
Piechowski et al. 2014) reported for dinosauriforms and
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
pseudosuchians or non-archosaurian archosauriforms (Fig.
6B). It was hypothesized as the ossification of the iliofe-
moral ligament insertion (Griffin & Nesbitt 2016). The
occurrence of this trait in a non-archosaurian archosauri-
form implies a similar origin as that of the anterior tro-
chanter, meaning that it could have been secondarily lost
in pseudosuchians, whereas in Pan-Aves it was retained.
The alternative scenario implies the independent origin of
the scar in Pan-Aves and proterochampsids (considering
the presence in both Stenoscelida aurantiacus and
Gualosuchus reigi). It has been demonstrated that the
presence and shape of the anterolateral scar is affected by
intraspecific variation (Piechowski et al. 2014; Griffin &
Nesbitt 2016; M€uller 2022). Therefore, the presence/
absence of this trait must be carefully considered.
The occurrence of the iliofibularis tubercle on the fib-
ula of Stenoscelida aurantiacus is another unusual trait
for Proterochampsia. Nevertheless, this feature is widely
distributed within crocopodans (Ezcurra et al. 2020a),
and it is not regarded as a traditional synapomorphy of
any exclusive clade (e.g. Nesbitt 2011; Ezcurra 2016).
Similarly, the presence of a vestigial phalanx in digit V
comprises another peculiar condition among the mem-
bers of the clade. Yet, the presence/absence of pha-
langes in digit V, which is often vestigial, is highly
affected by taphonomic biases.
Local palaeofauna
The Varzea do Agudo (¼ Janner) site is one of the most
prolific fossiliferous localities from southern Brazil,
 A new proterochampsid from Brazil
from which year-after-year numerous new specimens
have been exhumed. This fauna is dominated by traver-
sodontid cynodonts of the genus Exaeretodon (M€uller
et al. 2020b). This remarkable abundance has been used
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
et al. 2020). As a consequence, the younger Brazilian
proterochampsids are divided into two distinct horizons
within the Hyperodapedon AZ. Rhadinosuchus gracilis
and Cerritosaurus binsfeldi were unearthed from out-
crops of the Alemoa complex (Garcia et al. 2019),
which belong to the Hyperodapedon Acme Zone (lower
portion; Ezcurra et al. 2015; Schultz et al. 2020),
whereas Proterochampsa nodosa comes from
Exaeretodon-dominated beds (upper portion; Schultz
et al. 2020). In this context, Stenoscelida aurantiacus
expands the abundance and diversity of proterochamp-
sids in the Exaeretodon sub-AZ.
The presence of a proterochampsid in the Varzea do
Agudo site is not particularly helpful regarding a bio-
stratigraphic approach because proterochampsids occur
across the entire Ischigualasto Formation (Martınez
et al. 2013). Nevertheless, the new taxon comprises an
additional predatorial taxon for the palaeofauna of this
site. The absence of cranial remains precludes an unam-
biguous interpretation regarding the diet of Stenoscelida
aurantiacus, but since all proterochampsids are consid-
ered carnivorous animals (Trotteyn et al. 2013), the null
hypothesis is that Stenoscelida aurantiacus exhibited a
similar diet. So far, Stenoscelida aurantiacus and
Dynamosuchus collisensis comprise the only carnivorous
archosauriforms from this site, with the ectetniniid cyno-
dont Trucidocynodon riograndensis (Oliveira et al.
2010) adding to this context. Other occurrences at the
site belong to herbivorous/omnivorous animals: the
archosauromorph Hyperodapedon sp., the traversodontid
cynodont Exaeretodon, and the sauropodomorph dino-
saurs Pampadromaeus barberenai (Cabreira et al. 2011)
and Bagualosaurus agudoensis (Pretto et al. 2019).
The putative mode of life related to water bodies
often attributed to proterochampsids are an interesting
theme added to the Varzea do Agudo palaeofauna,
which previously recorded only fully terrestrial taxa (see
above). Currently, the site lacks other organisms that
possibly preyed within aquatic environments, such as
temnospondyls and phytosaurs. Therefore, if protero-
champsids (especially Stenoscelida aurantiacus) were
semi-aquatic predators (following the general morph-
ology of extant crocodilians; Trotteyn et al. 2013), the
new taxon expands the ecological diversity within the
Varzea do Agudo site. On the other hand, some methods
have cast doubts on these habits. Therefore, while some
15
proterochampsids probably had a mode of life related to
water bodies, other were more likely terrestrial in habit
(e.g. Chanaresuchus bonapartei; Arcucci et al. 2019;
Ezcurra et al. 2021).
Conclusions
Stenoscelida aurantiacus was erected based on the new
specimen CAPPA/UFSM 0293, composed of a nearly
complete right hind limb. Its phylogenetic position
within Proterochampsidae is uncertain; however, our
results support the new species as a non-rhadinosuchine
proterochampsid. The discovery of a complete and rela-
tively well-preserved hind limb of a proterochampsid is
of great interest because the fossil record of this skeletal
portion is poorly preserved, especially from the mid-
dle–late Carnian to early Norian. Although the new
taxon displays traits that unequivocally place it in
Proterochampsidae, Stenoscelida aurantiacus carries a
set of unusual hind limb traits within proterochampsids,
such as the presence of an anterior trochanter and a
dorsolateral scar on the femoral head, an iliofibularis
tubercle on the anterior margin of the fibular shaft, and
a vestigial phalanx in digit V.
Proterochampsids are rare in the Late Triassic sedi-
ments of Brazil when compared to those of Argentina.
Stenoscelida aurantiacus is the first proterochampsid
from the Varzea do Agudo site, expanding the taxo-
nomic and ecological diversity of this iconic fossilifer-
ous locality. The investigation and understanding of the
Carnian biota is crucial because this moment witnessed
the rise of dinosaurs (Novas et al. 2021) and the dawn
of modern ecosystems (Dal Corso et al. 2020).
Acknowledgements
We thank Martin D. Ezcurra for photographs of some
proterochampsids. We thank the Willi Hennig Society
for supporting the free use of TNT software. We extend
our gratitude to Martin D. Ezcurra and an anonymous
reviewer for corrections and comments that greatly
improved this manuscript. We are grateful to Caio
Fantini for the digital reconstruction of Stenoscelida
aurantiacus depicted in Figure 7. This work was carried
out with aid of the Fundaç~ao de Amparo a Pesquisa do
Estado do Rio Grande do Sul (FAPERGS 21/2551-
0000680-3) and the Conselho Nacional de
Desenvolvimento Cientıfico e Tecnologico (CNPq
404095/2021-6) to Rodrigo Temp M€uller, and a
Coordenaç~ao de Aperfeiçoamento de Pessoal de Nıvel
16 R. T. M€uller et al.
Superior (CAPES 88887.608076/2021-00) scholarship
for Mauricio Silva Garcia.
Supplemental material
Supplemental material for this article can be accessed
here: https://doi.org/10.1080/14772019.2022.2128913.
ORCID
Rodrigo Temp M€uller http://orcid.org/0000-0001-
8894-9875
Mauricio Silva Garcia http://orcid.org/0000-0001-
5317-0140
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