Journal of Vertebrate Paleontology

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A sauropodomorph (Dinosauria, Saurischia)

specimen from the Upper Triassic of southern
Brazil and the early increase in size in
Sauropodomorpha

Rodrigo T. Müller & Maurício S. Garcia

To cite this article: Rodrigo T. Müller & Maurício S. Garcia (2022): A sauropodomorph (Dinosauria,
Saurischia) specimen from the Upper Triassic of southern Brazil and the early increase in size in
Sauropodomorpha, Journal of Vertebrate Paleontology, DOI: 10.1080/02724634.2021.2002879

To link to this article: https://doi.org/10.1080/02724634.2021.2002879

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Journal of Vertebrate Paleontology e2002879 (20 pages)
© by the Society of Vertebrate Paleontology
DOI: 10.1080/02724634.2021.2002879

ARTICLE

A SAUROPODOMORPH (DINOSAURIA, SAURISCHIA) SPECIMEN FROM THE UPPER

TRIASSIC OF SOUTHERN BRAZIL AND THE EARLY INCREASE IN SIZE IN
SAUROPODOMORPHA

RODRIGO T. MÜLLER,1,2* and MAURÍCIO S. GARCIA2

1
Centro de Apoio à Pesquisa Paleontológica da Quarta Colônia, Universidade Federal de Santa Maria, Rua Maximiliano Vizzotto,
598, 97230-000, São João do Polêsine, Rio Grande do Sul, Brazil, rodrigotmuller@hotmail.com;
2
Programa de Pós-Graduação em Biodiversidade Animal, Universidade Federal de Santa Maria, 97105-120, Santa Maria, Rio
Grande do Sul, Brazil, mauriciossauro@gmail.com

ABSTRACT—Whereas sauropod dinosaurs from the Jurassic and Cretaceous Periods were the largest land animals that ever
lived, some of their early relatives evolved relatively large bodies during the Triassic Period. The evolutionary pathways
followed by the earliest sauropodomorphs towards the acquisition of massive bodies are poorly understood. However,
new finds from South America and Africa are reshaping our knowledge of this issue. Here, we describe a new early and
relatively large sauropodomorph represented by a partial postcranial skeleton excavated from Carnian-aged beds (Upper
Triassic) of southern Brazil. The new specimen is recovered as a sauropodomorph more closely related to bagualosaurians
than saturnaliids or other early-diverging forms in two phylogenetic analyses. The new specimen is generically
indeterminate but provides important evidence of an early increase in body size in Sauropodomorpha, being significantly
larger than that of coeval or older forms (except Bagualosaurus agudoensis). Furthermore, the specimen is about 3.2 times
heavier than Buriolestes schultzi, the earliest-branching sauropodomorph. The slender hind limbs and typical cursorial
proportions present in the earliest sauropodomorphs are mostly maintained in the new specimen despite its larger body size.

SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

Citation for this article: Müller, R. T. and M. S. Garcia. 2022. A sauropodomorph (Dinosauria, Saurischia) specimen from the
Upper Triassic of southern Brazil and the early increase in size in Sauropodomorpha. Journal of Vertebrate Paleontology.
DOI: 10.1080/02724634.2021.2002879

INTRODUCTION interval, Santa Maria Formation) yield the earliest members of

Dinosauria worldwide (233.23 ± 0.73 Ma; mid-Carnian; Langer
From birds to sauropods, dinosaurs achieved an extreme range
et al., 2018), including sauropodomorphs such as Saturnalia tupi-
of body sizes during their evolutionary history (Benson et al.,
niquim (Langer et al., 1999) and Buriolestes schultzi (Cabreira
2014), with a striking span during the Jurassic and Cretaceous
et al., 2016). These animals were small (femur ∼15 cm long),
Periods (Lacovara et al., 2014; Riga et al., 2016; Carballido
bipedal, and faunivorous (Cabreira et al., 2016; Bronzati et al.,
et al., 2017; Otero et al., 2021). Some non-sauropod sauropodo-
2017, 2019; Müller et al., 2021). The next-youngest stratigraphic
morphs evolved massive and large bodies during the Triassic
horizons (middle, Santa Maria Formation, precise age
Period (Pol and Powell, 2007; Apaldetti et al., 2018; Peyre de
unknown) contain sauropodomorphs displaying dental changes
Fabrègues and Allain, 2019). Towards the end of the Triassic
related to an omnivorous or mostly herbivorous diet, such as
(mid-Norian), quadrupedal posture was adopted for the first
Pampadromaeus barberenai (Cabreira et al., 2011) and Bagual-
time, which evolved rapidly alongside the evolution of increased
osaurus agudoensis (Pretto et al., 2018). The latter species
body mass (Langer et al., 1999; Martinez et al., 2011; Sereno
reveals a slight increase in body size, with femoral length over
et al., 2013; Cabreira et al., 2016; McPhee et al., 2018; Müller
20 cm (Pretto et al., 2018). Finally, the upper stratigraphic interval
et al., 2018a; Novas et al., 2021).
(Caturrita Formation, 225.42 ± 0.37 Ma; early Norian; Langer
Until recently, the evolution of body size in the earliest sauro-
et al., 2018) preserves sauropodomorphs exhibiting the typical
podomorphs has remained obscure. However, increased field-
bauplan of the so-called ‘prosauropods’ (sensu Sereno, 2007),
work has yielded several important finds (e.g., Cabreira et al.,
such as Unaysaurus tolentinoi (Leal et al., 2004) and Macrocollum
2011, 2016; Apaldetti et al., 2018; Müller et al., 2018b; Pretto
itaquii (Müller et al., 2018b). These animals were herbivorous,
et al., 2018) that have reshaped our knowledge of their early evol-
longer-necked, and larger, with femoral lengths over 30 cm
ution (Novas et al., 2021; Pol et al., 2021). Among these deposits,
(Leal et al., 2004; Müller et al., 2018b). Both Unaysaurus tolenti-
those from southern Brazil provide clues to the first steps of the
noi and Macrocollum itaquii lack any evidence of a quadrupedal
sauropodomorph radiation via three distinct horizons (Müller
stance, retaining the plesiomorphic bipedal condition of earliest
and Garcia, 2020) that encompass a ∼8 Ma time interval
sauropodomorphs (McPhee et al., 2018).
(Langer et al., 2018). The oldest horizons (i.e., lower stratigraphic
The fossil record from the middle stratigraphic interval is
scarce in comparison to that of the lower and upper intervals,
and its limited nature hampers the assessment of the macroevo-
*Corresponding author. lutionary trends that took place during this crucial period. For
Color versions of one or more of the figures in the article can be found instance, the anatomical innovations that accompanied the
online at www.tandfonline.com/ujvp.

Published online 11 Feb 2022

 Müller and Garcia—An early sauropodomorph from Brazil (e2002879-2)
body size increase during the early evolution of sauropodo-
morphs are poorly understood. Here, we explore this issue with
the description of a new specimen from this transitional stage.
It was excavated from the ‘Várzea do Agudo’ site (southern
Brazil) and is notable for its large size in comparison to the
oldest members of Sauropodomorpha.
Institutional Abbreviations—BP, Evolutionary Studies Insti-
tute, Johannesburg, South Africa (previously the Bernard Price
Institute); BRSUG, University of Bristol, Geology Department,
Bristol, United Kingdom; CAPPA/UFSM, Centro de Apoio à
Pesquisa Paleontológica da Quarta Colônia, Universidade
Federal de Santa Maria, São João do Polêsine, Brazil; GPIT,
Institut und Museum für Geologie und Paläontologie, Universi-
tat Tübingen, Tübingen, Germany; ISI, Geological Studies Unit
of the Indian Statistical Institute, Calcutta, India; MB, Museum
für Naturkunde, Berlin, Germany; MCN, Museu de Ciências
Naturais, Fundaçãoo Zoobotânica do Rio Grande do Sul,
Porto Alegre, Brazil; MCP, Museu de Ciências e Tecnologia Pon-
tifícia Universidade Católica do Rio Grande do Sul, Porto
Alegre, Brazil; MCZ, Museum of Comparative Zoology,
Harvard University, Cambridge, Massachusetts, U.S.A.; MLP,
Museo de La Plata, La Plata, Argentina; MNHN Muséum
National d’Histoire Naturelle, Paris, France; PVL, Instituto
Miguel Lillo, Universidad de Tucumán, San Miguel de
Tucumán, Argentina; PVSJ, Instituto y Museo de Ciencias Nat-
urales, Universidad Nacional de San Juan, San Juan, Argentina;
TMM, Vertebrate Paleontology Laboratory, University of Texas
at Austin, Austin, TX, U.S.A.; UFRGS, Paleovertebrate Collec-
tion of the Laboratório de Paleovertebrados da Universidade
Federal do Rio Grande do Sul, Porto Alegre, Brazil; UFSM,
Laboratório de Estratigrafia e Paleobiologia, Universidade
Federal de Santa Maria, Santa Maria, Brazil; ULBRA, Universi-
dade Luterana do Brasil, Coleção de paleovertebrados, Canoas,
Brazil; UMMP, University of Michigan Museum of Paleontology,
Ann Arbor, MI, U.S.A.
GEOLOGICAL SETTING
‘Várzea do Agudo’ also known as ‘Janner’ site is situated about
2 km west of the urban area of the Agudo municipality (29°39′
10.89″S, 53°17′34.20″W), Rio Grande do Sul, Brazil (Fig. 1A).
It is exposed in ravines of reddish massive to laminated mud-
stones, associated with yellowish cross-bedded sandstones and
sandstones with mudstone interclasts (Fig. 1B). The reddish
mudstones constitute most of the outcrop and are interpreted
as representative of a distal floodplain accumulation, whereas
the yellowish sandstones are mostly present in the top portion
of the ravines and represent a river channel (Da-Rosa, 2015;
Pretto et al., 2018). The mudstones belong to the upper portion
of the Alemoa Member of the Santa Maria Formation, lower
to middle Candelária Sequence, Paraná Basin (Zerfass et al.,
2003; Horn et al., 2014), whereas the overlaying sandstones are
considered part of the Caturrita Formation, upper Candelária
Sequence (Zerfass et al., 2003; Horn et al., 2014).
Based on biostratigraphy, the site is considered Late Triassic
(Langer et al., 2007a, 2018). Its faunal content allows its assign-
ment to the Hyperodapedon Assemblage Zone, comprising the
older Hyperodapedon Acme Zone and the younger Exaeretodon
sub-Assemblage Zone, in which the ‘Várzea do Agudo’ site is
situated (Langer et al., 2007a; Müller and Garcia, 2020; Schultz
et al., 2020). The distinction between both zones is based on
the relative abundance of the index taxa in the corresponding
strata, that is corroborated by biostratigraphic data from Argen-
tina (Martinez et al., 2013; Desojo et al., 2020). These biostrati-
graphic data suggest a latest Carnian age for the site, but
radiometric data are lacking (Müller and Garcia, 2020).
The ‘Várzea do Agudo’ paleofauna is dominated by cynodonts
with a few archosauromorph representatives (Müller et al.,
2020a). This fauna is intriguing because it serves as a bridge
between the land ecosystems with few dinosaurs and the first
environments dominated by dinosaurs (Martinez et al., 2013;
Pol et al., 2021). Regarding cynodonts, the most abundant
taxon is the traversodontid Exaeretodon riograndensis (Oliveira
et al., 2007; Liparini et al., 2013; Müller et al., 2020a), whereas
specimens of the ecteniniid Trucidocynodon riograndensis were
rare (Oliveira et al., 2010; Stefanello et al., 2018). On the other
hand, archosauromorphs are represented by dinosaurs (e.g.,
Pampadromaeus barberenai and Bagualosaurus agudoensis;
Cabreira et al., 2011; Pretto et al., 2018), as well as the ornithosu-
chid Dynamosuchus collisensis (Müller et al., 2020b). Further-
more, rhynchosaur remains attributed to Hyperodapedon sp.
were collected (Langer et al., 2007a), however, with the discov-
ery of the taxon Teyumbaita sulcognathus (Montefeltro et al.,
2010) occurring in stratigraphic levels above Hyperodapedon in
several sites from both Brazil and Argentina (Desojo et al.,
2020), the assignment of these remains to Hyperodapedon
should be considered carefully until they are properly reviewed.
MATERIAL AND METHODS
Material
The specimen here described is housed at the Centro de Apoio
à Pesquisa Paleontológica da Quarta Colônia/Universidade
Federal de Santa Maria (CAPPA/UFSM), under specimen
number CAPPA/UFSM 0276 (Fig. 1C−L). It was prepared
using air scribe tools, scalpels, and needles. Finally, the specimen
was stabilized with Paraloid B-72 (Koob, 1986).
Anatomical and Taxonomic Terminology
This study employs traditional (Romerian) anatomical terms
(i.e., ‘anterior’ and ‘posterior’). The nomenclature for vertebral
laminae follows Wilson (1999) and the nomenclature for ver-
tebral fossae follows Wilson et al. (2011).
The phylogenetic definitions of clades discussed below follow
Langer et al. (2019). Therefore, Bagualosauria comprises the
“minimal clade including Bagualosaurus agudoensis and Salta-
saurus loricatus” (node-based definition; Langer et al., 2019)
and Saturnaliidae represents the “maximal sauropodomorph
clade to encompass Saturnalia tupiniquim, but not Plateosaurus
engelhardti” (branch-based definition; Langer et al., 2019).
Phylogenetic Analysis
The phylogenetic affinities of CAPPA/UFSM 0276 were inves-
tigated with two data matrices. First, the specimen was scored in
the data matrix of Müller (2021), which includes a wide range of
early dinosaurs and related forms. We excluded Nhandumirim
waldsangae given its immature ontogenetic status (Marsola
et al., 2018). The final data matrix includes 277 characters and
63 operational taxonomic units (OTUs). It was analyzed in the
software TNT v.1.1 (Goloboff et al., 2008) following the same par-
ameters as in the former study (i.e., Müller 2021). Therefore,
Euparkeria capensis was used to root the most parsimonious
trees, which were recovered with a ‘Traditional search’ (random
addition sequence + tree bisection reconnection) with 1000 repli-
cates of Wagner trees (with random seed = 0) and using tree
bisection reconnection and branch swapping (holding 10 trees
save per replication). All characters were treated as having
equal weight, and the following characters were treated as
ordered: 4, 13, 18, 25, 63, 82, 84, 87, 89, 109, 142, 166, 174, 175,
184, 186, 190, 201, 203, 205, 209, 212, 225, 235, 236, 239, 250,
and 256. Topologies retained in overflowed replications were
branch-swapped for most parsimonious trees using tree bisection
reconnection. The strict consensus tree was generated using all
 Müller and Garcia—An early sauropodomorph from Brazil (e2002879-3)

FIGURE 1. Location and geological context of the ‘Várzea do Agudo’ site (Agudo, Rio Grande do Sul, Brazil) and specimen CAPPA/UFSM 0276.
A, surface distribution of the geological units in the area (modified from Müller, 2021); B, general view of the site; C, reconstruction of the skeleton of
CAPPA/UFSM 0276 depicting the preserved elements; D, left ilium in lateral view; E, caudal series in left lateral view; F, right ulna in medial view;
G, indeterminate metacarpal in dorsal view; H, indeterminate manual phalanx in dorsal view; I, left femur in anterior view; J, left tibia in lateral view;
K, left fibula in lateral view; L, left metatarsal II in anterior view.
 Müller and Garcia—An early sauropodomorph from Brazil (e2002879-4)

trees recovered in the analysis and all OTUs. Decay indices
(Bremer support values) were also obtained with TNT v. 1.1.
For the second analysis, CAPPA/UFSM 0276 was scored in the
data matrix of Müller et al. (2021). This data matrix was
employed because of the large sample of sauropodomorphs.
The final data matrix includes 404 characters and 78 OTUs. Fol-
lowing McPhee et al. (2018), characters 199 and 278 were set as
inactive (ambiguous scores and difficulty of interpretation). The
data matrix was subjected to an equally weighted parsimony
analysis in TNT v.1.1 (Goloboff et al., 2008) following the same
parameters as in the former study (i.e., Müller et al., 2021).
Hence, Euparkeria capensis was used to root the most parsimo-
nious trees, which were recovered with a ‘Traditional search’
(random addition sequence + tree bisection reconnection) with
1000 replicates of Wagner trees (with random seed = 1), and
using tree bisection reconnection and branch swapping
(holding 10 trees save per replicate). All characters were
treated as having equal weight, and the following characters
were treated as ordered: 8, 14, 22, 26, 43, 63, 75, 108, 118, 135,
141, 151, 154, 167, 170, 172, 173, 180, 185, 190, 193, 200, 207,
231, 234, 241, 251, 256, 264, 273, 282, 285, 299, 312, 334, 340,
348, 372, 385, 388, 391, 396, and 404. Topologies retained in over-
flowed replications were branch-swapped for most parsimonious
trees using tree bisection reconnection. The strict consensus tree
was generated using all trees recovered in the analysis and all
OTUs. Such as in the first analysis, decay indices (Bremer
support values) were obtained with TNT v. 1.1.
Body Mass Estimation
The body mass (BM) of CAPPA/UFSM 0276 was estimated
based on the femoral circumference (Cfem), according to the
equation by Campione et al. (2014): log10BM = 2.754 × log10
(Cfem) − 0.683. We used this equation, which assumes that
CAPPA/UFSM 0276 was bipedal, because the specimen lacks
the following osteological indicators of quadrupedal stance, a
triradiate proximal ulna (i.e., deep radial fossa); straightening
of the femoral shaft; loss of the anterior trochanter (= lesser tro-
chanter); migration of the fourth trochanter distally and medi-
ally; and a robust fibula (McPhee et al., 2018).
portions. Based on the absence of duplicated bones and their
relative size, the elements are believed to belong to a single indi-
vidual. Most bones are complete, yet some degree of sedimentary
compression is observed (see description for additional com-
ments). Furthermore, most of the elements present a grayish
coloration with orange extremities, as is standard for materials
collected in the ‘Várzea do Agudo’ locality.
Ontogenetic Assessment—The specimen appears to have
reached skeletal maturity based on the absence of open neurocen-
tral sutures (Brochu, 1996) and presence/shape of some muscle
attachment structures (e.g., trochanteric shelf, dorsolateral tro-
chanter; Piechowski et al., 2014; Griffin and Nesbitt, 2016).
DESCRIPTION
Axial Skeleton
Caudal Vertebrae—The centra of caudal vertebrae (Figs. 1E, 2)
are spool-shaped as in other early dinosaurs. The centra of anterior
caudal vertebrae are anteroposteriorly short (Fig. 2C); i.e., the dor-
soventral height of the anterior articular facet is greater than the
anteroposterior length of the centrum (height:length ratio = 1.1;
see Table S1 in Supplemental Data for measurements). The oppo-
site condition is observed in the posterior caudal vertebrae, where
the centrum is longer (height:length ratio of the posteriormost pre-
served centrum = 0.74). Anteroposteriorly compressed anterior
caudal vertebral centra is the typical condition in herrerasaurids,
for example as Herrerasaurus ischigualastensis (PVL 2566; Novas,
1993) and Gnathovorax cabreirai (CAPPA/UFSM 0009; Pacheco
et al., 2019). However, this condition is not unique to herrerasaur-
ids, being observed in some sauropodomorphs (e.g., Coloradi-
saurus brevis, PVL 5904; Eucnemesaurus entaxoni, BP/1/6234;

SYSTEMATIC PALEONTOLOGY
DINOSAURIA Owen, 1842
SAURISCHIA Seeley, 1887
SAUROPODOMORPHA Huene, 1932
Gen. et sp. indet
(Figs. 1–9)

Material—Specimen CAPPA/UFSM 0276 (Fig. 1C–L), partial

postcranial skeleton including 15 caudal vertebrae, the right ulna,
one possible metacarpal, one manual phalanx, the partial left
ilium, the partial left hind limb (including femur, tibia, fibula,
and one partial metatarsal II).
Locality and Horizon—‘Várzea do Agudo’ or ‘Janner’ site (29°
39′10.89″S, 53°17′34.20″W), Agudo, Rio Grande do Sul, Brazil. It
belongs to the Santa Maria Formation, Candelária Sequence,
Paraná Basin (Zerfass et al., 2003; Horn et al., 2014; Fig. 1A).
The site is Carnian in age based on biostratigraphic data
(Langer et al., 2007a; Schultz et al., 2020).
Remarks—The bone elements were accumulated in a small
area of about 1 m2 on the same layer that yielded the holotype FIGURE 2. First caudal vertebra of CAPPA/UFSM 0276 in A, anterior,
B, ventral, and C, left lateral views. The hatched areas are matrix,
of Bagualosaurus agudoensis (Pretto et al., 2018), whereas the whereas the black areas are broken. Abbreviations: acdl, anterior centro-
holotype of Pampadromaeus barberenai comes from a lower diapophyseal lamina; c, centrum; dpr, depression; nc, neural canal; ns,
level (Müller et al., 2016). Most of the caudal series was in articu- neural spine; prcdf, prezygapophyseal centrodiapophyseal fossa; prdl,
lation (a segment with nine vertebrae), as was the hind limb. The prezygodiapophyseal lamina; pz, postzygapophysis; tp, transverse
remaining elements were found scattered around the articulated process.
 Müller and Garcia—An early sauropodomorph from Brazil (e2002879-5)
Apaldetti et al., 2013; McPhee et al., 2015), whereas in the earliest
sauropodomorphs the centrum is proportionally longer (e.g., Bur-
iolestes schultzi, ULBRA-PVT280; Cabreira et al., 2016). There is
a fossa on the lateral surface of the centrum of the anterior
caudal vertebrae (Fig. 2C). It becomes shallow to almost impercep-
tible in the subsequent elements. Unlike Macrocollum itaquii
(CAPPA/UFSM 0001b; Müller et al., 2018b), the ventral surface
of the centra lacks any longitudinal ridges and grooves along the
entire preserved series. There are facets for chevron articulation
in the anterior and posterior ventral corners of the preserved
centra. The only exception is the probable first caudal vertebra,
which lacks facets for chevron articulation on the ventral margin
(Fig. 2B).
The neural arch is poorly preserved in all the caudal vertebral
elements. It is dorsoventrally shorter than the height of its
respective centrum. The height of the neural canal is ∼0.3
times the height of the anterior articular facet of the centrum
in the first caudal vertebra (Fig. 2A). In addition, the neural
canal of the anterior caudal vertebrae is wider than deep. The
prezygapophyses are poorly preserved in anterior elements. In
the posteriormost preserved element, the prezygapophyses are
anteriorly short, barely projecting beyond the anterior articular
facet of their respective centrum. Therefore, this condition differ-
entiates the specimen from herrerasaurids with elongated prezy-
gapophyses (e.g., Herrerasaurus ischigualastensis, PVSJ 53 and
PVSJ 2566; Gnathovorax cabreirai, CAPPA/UFSM 0009;
Novas, 1993; Pacheco et al., 2019). There is a prezygodiapophy-
seal lamina on the first and second caudal vertebrae. This
lamina roofs a prezygapophyseal centrodiapophyseal fossa in
these two vertebrae. Both the lamina and fossa are absent in
the subsequent elements. An anterior centrodiapophyseal
lamina runs from the anterior facet of the centrum to the
ventral portion of the transverse process in the first caudal verte-
bra. In the second caudal vertebra this lamina is poorly devel-
oped, whereas in the subsequent vertebrae it is absent. The
transverse processes of the anterior vertebrae are poorly pre-
served. In the subsequent elements they are laterally oriented,
located in the posterior half of the neural arch, and dorsoven-
trally flat. Finally, the transverse processes are reduced to faint
projections in the posterior elements.
The articular facet of the postzygapophyses faces ventrally in
the anterior caudal vertebrae, whereas in the subsequent
elements it faces ventrolaterally. The postzygapophyses lack
any evidence of projections related to an accessory intervertebral
articulation. The neural spine is incomplete in the anterior
elements and poorly preserved along the series. In the posterior-
most preserved vertebrae, the neural spine is located on the pos-
terior portion of the neural arch and is directed posterodorsally.
It is dorsoventrally short, hardly projecting beyond the dorsal
margin of the postzygapophyses.
Forelimb
Ulna—The shape of the ulna was affected by sedimentary
compression, resulting in a transversely compressed shaft
(Fig. 3). The length of ulna is about 0.42 times that of tibia (see
Table S2 in Supplemental Data for measurements). The ulna
bows medially along its shaft (Fig. 3A, E). The lateral surface
is straight to slightly convex, whereas the medial surface is
concave. The proximal articular surface is sub-triangular in prox-
imal view (Fig. 3C), distinct from the triradiate shape present in
sauropodiforms (e.g., Aardonyx celestae, BP/1/5379c; Leduma-
hadi mafube, BP/1/7120; Yates et al., 2010; McPhee et al.,
2018). The anteromedial process is rounded and poorly devel-
oped. In contrast to most massopodans (e.g., Mussaurus patago-
nicus, MLP 68-II-27-1; Otero and Pol, 2013; Sefapanosaurus
zastronensis, BP/1/7437; Otero et al., 2015), there is no evidence
of a radial fossa between the anterolateral and anteromedial
processes (Fig. 3C). The olecranon process is short (Fig. 3F),
around 7% of the total length of the ulna. Some other dinosaurs
have a proximodistally enlarged olecranon process. For instance,
the olecranon process is about 10.5% in Gnathovorax cabreirai
(CAPPA/UFSM 0009; Pacheco et al., 2019), 10% in Pampadro-
maeus barberenai (ULBRA-PVT016; Langer et al., 2019), and
23% (estimated) in Saturnalia tupiniquim (MCP 3844-PV;
Langer et al., 2007b). In contrast, the process is reduced in Bur-
iolestes schultzi (ULBRA-PVT280; Cabreira et al., 2016:7.8%)
and in Macrocollum itaquii (CAPPA/UFSM 0001a; Müller
et al., 2018b:7.5%). Nevertheless, the shape and size of the ole-
cranon process presents strong intraspecific variation in some
dinosaurs depending upon the ossification degree of cartilagi-
nous structures (e.g., Kentrosaurus aethiopicus, GPIT 1424;
MB.R.4800.33; Mallison, 2010).
The ulnar shaft is quite slender, such as in early sauropodo-
morphs (e.g., Buriolestes schultzi, ULBRA-PVT280), whereas
in sauropodiforms it is usually robust (e.g., Mussaurus patagoni-
cus, MLP 68-II-27-1; Pulanesaura eocollum, BP/1/6210; McPhee
and Choiniere, 2018). The shaft is featureless, whereas in Pampa-
dromaeus barberenai (ULBRA-PVT016; Langer et al., 2019)
there is a suite of longitudinal crests and grooves. In lateral
view (Fig. 3A), the anterior margin is concave, whereas the pos-
terior margin is sigmoidal. The distal portion of the ulna is
slightly rotated with respect to the proximal articular surface.
There is no evidence of a tubercle for the radial-ulnar ligament,
which is well developed in Antetonitrus ingenipes (BP/1/4952;
McPhee et al., 2014). The distal articular surface is transversely
convex. In distal view, it is oval (Fig. 3D).
Indeterminate Metacarpal—The specimen preserves one poss-
ible metacarpal (Fig. 4A–D). However, some portions are
broken away, which hampers a confident identification. It
might also be a phalanx 1, but it is identified as an indeterminate
metacarpal because of the large size, proportions (more than two
times longer than transversely wide), presence of a flat proximal
articular surface (contrasting with the markedly concave surface
of non-proximal phalanges), absence of a dorsal intercondylar
process, and presence of a deep and wide extensor depression
on the dorsal surface of the distal portion. The indeterminate
metacarpal of CAPPA/UFSM 0276 is longer than wide (length:
proximal width = 2.41). All metacarpals are generally longer
than wide in early sauropodomorphs (Sereno et al., 2013; Cab-
reira et al., 2016). The metacarpal I of Eoraptor lunensis (PVSJ
512; Sereno et al., 2013) is longer than wide (length:proximal
width = 1.95; based on the left manus of PVSJ 512), whereas in
some sauropodiforms it is wider than long (e.g., Antetonitrus
ingenipes, BP/1/4952; McPhee et al., 2014; Aardonyx celestae,
BP/1/5379; Otero et al., 2015). However, the ratio of the metacar-
pal I of CAPPA/UFSM 0276 differentiates it from the indetermi-
nate metacarpal of Eoraptor lunensis. The distal asymmetry
typical of metacarpal I in early sauropodomorphs is also not
observed here. Additionally, the ratio of the length of the poss-
ible metacarpal relative to the length of the femur (0.14 in
CAPPA/UFSM 0276) can be compared with that of Eoraptor
lunensis. In the latter, the ratios are 0.09, 0.13, 0.14, and 0.10
for metacarpals I, II, III, and IV, respectively. Hence the
element herein described probably belongs to the manual digit
II or III, assuming a similar proportion when compared with Eor-
aptor lunensis (PVSJ 512; Sereno et al., 2013).
The proximal end is broader than deep and ovoid in proximal
view (Fig. 4C), with a straight palmar margin and a convex dorsal
margin. Moreover, the proximal articular surface is straight to
gently concave. The dorsal surface of the shaft is transversely
convex. As in the metacarpals of Eoraptor lunensis (PVSJ 512,
Sereno et al., 2013) the shaft is straight in lateral view
(Fig. 4B), whereas in Herrerasaurus ischigualastensis (PVSJ
373; Sereno, 1993) the shafts are bowed ventrally. There is a
deep extensor depression on the dorsal surface of the distal
 Müller and Garcia—An early sauropodomorph from Brazil (e2002879-6)

FIGURE 3. Right ulna of CAPPA/UFSM 0276 in A, lateral, B, anterior, C, proximal, D, distal, E, posterior, and F, medial views. The black areas are
broken. Abbreviations: alp, anterolateral process; amp, anteromedial process; op, olecranon process.
 Müller and Garcia—An early sauropodomorph from Brazil (e2002879-7)

FIGURE 4. Manus of CAPPA/UFSM 0276. Indeterminate metacarpal in A, dorsal, B, lateral or medial, C, proximal, and D, distal views; indetermi-
nate phalanx in E, dorsal, F, proximal, and G, distal views. The hatched areas are matrix, whereas the black areas are broken. Abbreviations: clp,
collateral pit; dc, distal condyle; ded, dorsal extensor depression; ig, intercondylar groove.

portion of the bone (Fig. 4A), which resembles the condition of
herrerasaurids Herrerasaurus ischigualastensis (PVSJ 373;
Sereno, 1993) and Gnathovorax cabreirai (CAPPA/UFSM
0035; Pacheco et al., 2019), whereas in Eoraptor lunensis (PVSJ
512, Sereno et al., 2013) the dorsal extensor depression is
shallow. The distal condyles are not entirely preserved
(Fig. 4D). Nevertheless, the collateral ligament pits are present.
Indeterminate Phalanx—There is only one isolated phalanx
preserved (Fig. 4E–G). It is longer than wide. The proximal
articular surface is concave and slightly broader than deep
(Fig. 4F). The palmar margin of the proximal end is not sym-
metrical; one of the corners extends further proximally
(Fig. 4E). The extensor depression on the dorsal surface of the
distal portion is poorly developed. The distal condyles are asym-
metrical and separated by a shallow intercondylar groove
(Fig. 4D). Its position is uncertain. However, the absence of a
dorsal intercondylar process on the proximal margin (Fig. 4E)
suggests that it is a proximal phalanx. In early sauropodomorphs
the first phalanx of digit I usually bears a well-developed dorsal
intercondylar process (e.g., Martinez et al., 2011; Sereno et al.,
2013; Ballell et al., 2020), and the first phalanx of digits IV and
V are reduced (e.g., Otero and Pol, 2013; Reiss and Mallison,
2014). Therefore, it is plausible that the isolated phalanx rep-
resents the first phalanx of the digit II or III.
Pelvic Girdle
Ilium—The preserved portion of the left ilium (Fig. 5)
resembles the general morphology of other coeval saurischians,
excepting herrerasaurids. There is a preacetabular fossa on the
anterior portion of the bone (Fig. 5A), such as in Pampadro-
maeus barberenai (ULBRA-PVT016; Langer et al., 2019) and
Thecodontosaurus antiquus (BRSUG 23613; Ballell et al.,
2020). Most of the iliac blade is missing, therefore its shape is
uncertain. Posterior to the preacetabular fossa and dorsal to
the supracetabular crest there is a concavity on the ventral
portion of the iliac blade. It corresponds to the attachment
area of m. iliofemoralis (see Hutchinson, 2001; Langer, 2003)
 Müller and Garcia—An early sauropodomorph from Brazil (e2002879-8)

FIGURE 5. Left ilium of CAPPA/UFSM 0276 in A, lateral, B, posterior, C, dorsal, and D, medial views. The hatched areas are matrix, whereas the
black areas are broken. Abbreviations: bf, brevis fossa; bs, brevis shelf; ib, iliac blade; ip, ischial peduncle; mb, medial blade; mw, medial wall; pa, post-
acetabular ala; pf, preacetabular fossa; sac, supracetabular crest; sr1, first primordial sacral rib articular surface; sr2, second primordial sacral rib articu-
lar surface.
 Müller and Garcia—An early sauropodomorph from Brazil (e2002879-9)
present in non-dinosaurian dinosauromorphs and plesiomorphic
in dinosaurs such as Saturnalia tupiniquim (MCP 3844-PV, MCP
3846-PV; Langer, 2003; Garcia et al., 2019), Herrerasaurus ischi-
gualastensis (PVL 2566; Novas, 1993), and Caseosaurus crosbyen-
sis (UMMP 8870; Baron and Williams, 2018). The supracetabular
crest is well developed (Fig. 5C), roofing the acetabulum and pro-
jecting ventrolaterally, differing from the laterally oriented crest
described for Bagualosaurus agudoensis (UFRGS-PV-1099-T;
Pretto et al., 2018). The posterior portion of the crest does not
merge with the anterior portion of the brevis shelf. The medial
wall of the acetabulum is extended and, as preserved, it bears a
slightly concave ventral margin (Fig. 5D), which is the condition
observed in similarly aged sauropodomorphs such as Buriolestes
schultzi (ULBRA-PVT280; Cabreira et al., 2016), Saturnalia
tupiniquim (MCP 3844-PV, MCP 3846-PV; Garcia et al., 2019)
and Pampadromaeus barberenai (ULBRA-PVT016; Langer
et al., 2019). It differentiates the specimen from herrerasaurids
and more advanced sauropodomorphs, where the acetabulum
is partially to completely perforated (Novas, 1993; Müller et al.,
2018b; Pacheco et al., 2019; Ballell et al., 2020).
The postacetabular ala is not completely preserved. However,
it seems elongated, distinct from the short ala of some saur-
ischians (Caseosaurus crosbyensis, UMMP 8870; Herrerasaurus
ischigualastensis, PVL 2566; Baron and Williams, 2018; Novas,
1993). The ventral margin of the brevis shelf is deep and covers
the medial blade of the postacetabular ala in lateral view
(Fig. 5A), such as in Panphagia protos (PVSJ 874; Martinez
and Alcober, 2009). Conversely, in Bagualosaurus agudoensis
(UFRGS-PV-1099-T; Pretto et al., 2018) and Thecodontosaurus
antiquus (BRSUG 23613; Ballell et al., 2020), the medial
lamina is deeper than the brevis shelf. The brevis fossa excavates
the ventral portion of the postacetabular ala forming a concave
surface (Fig. 5B). Whereas the fossa is ventrally oriented in the
specimen, in Bagualosaurus agudoensis (UFRGS-PV-1099-T;
Pretto et al., 2018) it is ventrolaterally oriented. Both the
brevis shelf and fossa are considered attachment areas for the
m. caudofemoralis brevis as in other dinosaurs (Hutchinson,
2001). Unlike in Buriolestes schultzi (ULBRA-PVT280; Cabreira
et al., 2016), Saturnalia tupiniquim (MCP 3844-PV; Garcia et al.,
2019) and Chromogisaurus novasi (PVSJ 845; Ezcurra, 2010),
there is no evidence of any raised scar (i.e., rugosity) on the
lateral surface of the preserved portion of the postacetabular
ala. Nevertheless, the postacetabular ala and the iliac blade are
not entirely preserved, hence, the absence of scars is ambiguous.
The ischial peduncle extends posteroventrally, being ovoid in
cross-section, and its convex anterior surface merges anteriorly
with the acetabulum wall. This region corresponds to the antitro-
chanter and does not bear the raised process seen in Guaiba-
saurus candelariensis (MCN PV2355, UFRGS PV0725T;
Langer et al., 2011) and Ixalerpeton polesinensis (ULBRA-
PVT059; Cabreira et al., 2016). Unlike Coloradisaurus brevis
(PVL 5904; Apaldetti et al., 2013) and Antetonitrus ingenipes
(NM QR1545; McPhee et al., 2014), the specimen lacks a pos-
terior projection (‘heel’) at the distal end of the ischial peduncle.
Likewise, there is no evidence of a small foramen (or pair of for-
amina) between the supracetabular crest and the brevis shelf
(Garcia et al., 2019). Medially, the bone preserves the scars
from the attachment of the two primordial sacral vertebrae
(Fig. 5D); however, the total number of sacral vertebrae is
uncertain.
Hind Limb
Femur—The femur (Fig. 6) is mainly straight in lateral/medial
and markedly sigmoid in anterior/posterior views, such as in
other early sauropodomorphs (e.g., Saturnalia tupiniquim,
MCP 3844-PV; Buriolestes schultzi, ULBRA-PVT280; Langer,
2003; Cabreira et al., 2016). On the other hand, the femora are
usually straight in anterior/posterior views in larger sauropodo-
morphs (e.g., Coloradisaurus brevis, PVL 5904; Meroktenos tha-
banensis, MNHN.F.LES16c; Apaldetti et al., 2013; Peyre de
Fabrègues and Allain, 2016). The proximal articular surface is
poorly preserved. Thus, the presence or absence of a proximal
groove is uncertain. The posteromedial and anteromedial tuber-
cles are subequal in size (Fig. 6C). In Macrocollum itaquii
(CAPPA/UFSM 0001b; Müller et al., 2018b), the posteromedial
tubercle is the largest one, whereas in the herrerasaurid Gnatho-
vorax cabreirai (CAPPA/UFSM 0009; Pacheco et al., 2019), the
posteromedial tubercle is absent.
The femoral head is expanded and bears a marked concave
surface where it merges with the femoral shaft (Fig. 6E). This
concave surface is limited to a restricted area on the posterome-
dial aspect of the femur. The dorsolateral trochanter is low and
rounded (Fig. 6A, B), like that of Saturnalia tupiniquim (MCP
3844-PV; Langer, 2003), Bagualosaurus agudoensis (UFRGS-
PV-1099-T; Pretto et al., 2018), and Pampadromaeus barberenai
(ULBRA-PVT016; Langer et al., 2019). A scar apparently
related to ontogeny (see Griffin and Nesbitt, 2016) and associ-
ated with the dorsolateral trochanter in some specimens of Bur-
iolestes schultzi (e.g., CAPPA/UFSM 0035; Müller et al., 2018a)
and Gnathovorax cabreirai (CAPPA/UFSM 0009; Pacheco
et al., 2019) is absent in CAPPA/UFSM 0276. The anterior tro-
chanter is small and lacks a cleft between the proximal tip and
the femoral shaft (Fig. 6F), contrasting with the basally branching
theropod Erythrovenator jacuiensis (CAPPA/UFSM 0157;
Müller, 2021). There is a trochanteric shelf associated with the
anterior trochanter (Fig. 6B). The shelf occurs in several early
saurischians (e.g., Gnathovorax cabreirai, CAPPA/UFSM 0009;
Pacheco et al., 2019; Buriolestes schultzi, ULBRA-PVT280; Cab-
reira et al., 2016); however, it is absent in post-Carnian sauropo-
domorphs (e.g., Macrocollum itaquii, CAPPA/UFSM 0001b;
Müller et al., 2018b; Thecodontosaurus antiquus, BRSUG
23615; Ballell et al., 2020). The fourth trochanter is crest-like
and slightly asymmetrical. If differs from the strongly symmetri-
cal fourth trochanter of theropods (Langer and Benton, 2006).
Moreover, the fourth trochanter is located at the posteromedial
surface and in the proximal half of the femoral shaft, whereas in
sauropodiforms it is located more distally (e.g., Meroktenos tha-
banensis, MNHN.F.LES16c; Peyre de Fabrègues and Allain,
2016; Antetonitrus ingenipes, BP/1/4952; McPhee et al., 2014).
The distal portion of the femur is transversely expanded
(Fig. 6D). Unlike theropods (Pinheiro, 2016) and several post-
Carnian sauropodomorphs (e.g., Coloradisaurus brevis, PVL
5904; Apaldetti et al., 2013), the anterior surface of the distal
portion of the bone lacks an extensor groove/depression. On
the opposite side, the popliteal fossa extends longitudinally and
separates the distal condyles (Fig. 6E). The fossa is much
shorter proximodistally than that of silesaurids and aphanosaurs
(Nesbitt et al., 2010, 2017). Conversely, the fossa is transversely
wide, such as in Pampadromaeus barberenai (ULBRA-
PVT016; Langer et al., 2019), whereas it is usually narrow in saur-
opodomorphs. The lateral and medial condyles are subequal in
size, whereas the crista tibiofibularis is slightly smaller than the
condyles. This condition differs from the poorly defined crista
tibiofibularis of Staurikosaurus pricei (MCZ 1669; Bittencourt
and Kellner, 2009).
Tibia—The tibia (Fig. 7) is slightly longer than the femur (see
Table S3 in Supplemental Data for measurements), such as in
other early sauropodomorphs. The reverse condition (i.e.,
femur longer than the tibia) occurs in most herrerasaurids and
post-Carnian sauropodomorphs, (e.g., Gnathovorax cabreirai,
CAPPA/UFSM 0009; Pacheco et al., 2019; Macrocollum itaquii,
CAPPA/UFSM 0001b; Müller et al., 2018b). The tibia is straight
(Fig. 7A, B) with an anteroposteriorly expanded proximal
portion. In proximal view (Fig. 7C), the bone is subtriangular
and the cnemial crest is directed anteriorly. The latter extends
 Müller and Garcia—An early sauropodomorph from Brazil (e2002879-10)

FIGURE 6. Left femur of CAPPA/UFSM 0276 in A, lateral, B, anterior, C, proximal, D, distal, E, posterior, and F, medial views. The black areas are
broken. Abbreviations: 4t, fourth trochanter; alt, anterolateral tubercle; amt, anteromedial tubercle; at, anterior trochanter; ctf, crista tibiofibularis;
dlt, dorsolateral trochanter; faa, facies articularis antitrochanterica; lc, lateral condyle; mc, medial condyle; pf, popliteal fossa; pmt, posteromedial
tubercle; ts, trochanteric shelf.
 Müller and Garcia—An early sauropodomorph from Brazil (e2002879-11)

FIGURE 7. Left tibia of CAPPA/UFSM 0276 in A, lateral, B, anterior, C, proximal, D, distal, E, posterior, and F, medial views. The black areas are
broken. Abbreviations: cc, cnemial crest; fta, foramen for the nutritive tibial artery; it, insisura tibialis; lc, lateral condyle; ln, lateral notch; mc, medial
condyle; plf, posterolateral flange; rdg, ridge.
 Müller and Garcia—An early sauropodomorph from Brazil (e2002879-12)
for approximately one fourth of the tibial length. There is a prox-
imodistal sulcus (= insisura tibialis sensu Butler, 2010) posterior
to the cnemial crest in the lateral surface of the proximal region
of the bone. Immediately posterior to the sulcus, there is a faint
ridge, which differs from the well-developed fibular crest of
neotheropods (e.g., Powellvenator podocitus, PVL 4414;
Ezcurra, 2017). The medial surface is gently concave. On the
proximal articular surface, the medial condyle is set posterior
to the lateral condyle, whereas in Jaklapallisaurus asymmetrica
(ISI R274; Novas et al., 2011) and Staurikosaurus pricei (MCZ
1669; Bittencourt and Kellner, 2009) both condyles lie near the
posterior margin. Distinct from Eoraptor lunensis (PVSJ 559;
Sereno et al., 2013), the specimen lacks a notch between the pos-
terior portion of the condyles, and both condyles are poorly
delimited. There is no projection from the medial condyle, as
in Buriolestes schultzi (ULBRA-PVT280; Cabreira et al., 2016).
The midshaft is ovoid in cross-section. The foramen for the
nutritive tibial artery lies on the proximal third of the lateral
portion of the bone (Fig. 7A). The distal end of the tibia is sub-
quadrangular in distal view (Fig. 7D), resembling the condition
of other early sauropodomorphs (e.g., Bagualosaurus agudoensis,
UFRGS-PV-1099-T; Pretto et al., 2018; Macrocollum itaquii,
CAPPA/UFSM 0001c; Müller et al., 2018b) and some herrera-
saurids (e.g., Herrerasaurus ischigualastensis, PVSJ 373; Novas,
1993). In contrast, some other herrerasaurids have a rounded/
ovoid outline in distal view (Garcia et al., 2021). The lateral
notch is longitudinally short. The posterior margin of the distal
end is deeper than the anterior margin. In addition, the posterior
margin is straight in distal view, whereas in Panphagia protos
(PVSJ 874; Martinez and Alcober, 2009) it is concave. In contrast
with Macrocollum itaquii (CAPPA/UFSM 0001c; Müller et al.,
2018b) and Panphagia protos (PVSJ 874; Martinez and
Alcober, 2009), the posterolateral flange of the new specimen
does not exceed the lateral margin of the bone in distal view.
This condition is usually seen in herrerasaurids (e.g., Herrera-
saurus ischigualastensis, PVSJ 373; Novas, 1993; Staurikosaurus
pricei, MCZ 1669; Bittencourt and Kellner, 2009). The postero-
lateral flange projects beyond the distal limit of the astragalar
articular surface, a trait common in post-Carnian sauropodo-
morphs (Langer, 2003; Garcia et al., 2021), but also present inci-
piently in Carnian forms such as Eoraptor lunensis (PVSJ 512;
Sereno et al., 2013) and Saturnalia tupiniquim (MCP 3844-PV;
Langer, 2003). There is no evidence of a posteromedial notch
(Fig. 7E, F), which occurs in Guaibasaurus candelariensis
(MCN-PV2356; Langer et al., 2011) and Bagualosaurus agudoen-
sis (UFRGS-PV-1099-T; Pretto et al., 2018).
Fibula—The fibula (Fig. 8) lacks part of the proximal surface.
The bone is slender and expands in the extremities (Fig. 8A). In
proximal view (Fig. 8C), the lateral margin of the proximal
articular surface is convex, whereas the medial margin is straight.
The latter margin is concave in Mussaurus patagonicus (MLP 68-
II-27-1; Otero and Pol, 2013) and Powellvenator podocitus (PVL
4414-4; Ezcurra, 2017). The proximal portion of the bone
expands anteroposteriorly and is transversely compressed.
There is a gently sloping ridge on the medial surface of the prox-
imal portion (Fig. 8F), similar to that of Saturnalia tupiniquim
(MCP 3844-PV; Langer, 2003) and Buriolestes schultzi
(CAPPA/UFSM 0035; Müller et al., 2018a). In the proximal
third of the bone, there is a proximodistally oriented ridge
(∼20 mm in length, see Fig. 8A, E) for the iliofibularis muscle
insertion (Langer, 2003), such as in Buriolestes schultzi
(CAPPA/UFSM 0035; Müller et al., 2018a), Chromogisaurus
novasi (PVSJ 845; Ezcurra, 2010), and Bagualosaurus agudoensis
(UFRGS-PV-1099-T; Pretto et al., 2018). This ridge is absent in
Macrocollum itaquii (CAPPA/UFSM 0001b; Müller et al.,
2018b). The specimen lacks any evidence of a tubercle on the
anteromedial margin of the shaft (Fig. 8B). A protruding tuber-
cle occurs in the proximal third of the fibula of Buriolestes
schultzi (CAPPA/UFSM 0035; Müller et al., 2018a), Pampadro-
maeus barberenai (ULBRA-PVT016; Langer et al., 2019), and
Gnathovorax cabreirai (CAPPA/UFSM 0009; Pacheco et al.,
2019). Also, in the proximal third of the fibula, there is no evi-
dence of a longitudinally oriented medial sulcus, which is
present in Staurikosaurus pricei (MCZ 1669, Bittencourt and
Kellner, 2009) and Gnathovorax cabreirai (Garcia et al., 2021).
The midshaft is ‘D’-shaped in cross-section, where the lateral
margin is convex and the medial is flat. The transverse width of
the midshaft is 0.56 times the width of the tibia. In lateral view,
the long axis of the shaft is not entirely straight, and the distal
third projects anteriorly (Fig. 8A). This condition resembles
that of Pampadromaeus barberenai (ULBRA-PVT016; Langer
et al., 2019); however, it is probably related to taphonomic defor-
mation in both specimens (Langer et al., 2019). The distal portion
of the bone expands anteroposteriorly, resulting in an oval extre-
mity in distal view (Fig. 8D). The anterior and posterior margins
of the distal portion are sharp, forming a ridge on each side. In
lateral/medial view, the posterior corner of the distal end
extends more distally than the anterior corner. Furthermore,
the distal margin is convex, whereas it is sigmoid in the theropod
Lepidus praecisio (TMM41936-1.3; Nesbitt and Ezcurra, 2015).
In contrast to Sefapanosaurus zastronensis (BP/1/7447; Otero
et al., 2015), the distal portion lacks any evidence of an enlarged
anteromedial projection.
Metatarsal II—The element preserves its proximal portion and
part of the shaft (Fig. 9). The general anatomy resembles that of
the metatarsal II of other early sauropodomorphs, such as Satur-
nalia tupiniquim (MCP 3844-PV; Langer, 2003) and Macrocol-
lum itaquii (CAPPA/UFSM 0001a; Müller et al., 2018b), where
the proximal articular surface is flat, dorsoplantarly expanded
(Fig. 9B, D), and subrectangular in proximal view (Fig. 9C).
This subrectangular surface differentiates the specimen from
some sauropodomorphs with an hourglass-shaped proximal
surface (e.g., Coloradisaurus brevis, PVL 5904; Apaldetti et al.,
2013). Moreover, the transversely compressed condition of the
proximal articular surface (Fig. 9A) differs from the transversely
expanded proximal articular surface of Pampadromaeus barber-
enai (ULBRA-PVT016; Langer et al., 2019). However, the latter
was strongly affected by sedimentary compression, therefore, its
anatomy should be carefully considered. The lateral margin of
the proximal surface is concave, whereas in early sauropodo-
morphs it is usually straight (e.g., Saturnalia tupiniquim, MCP
3844-PV; Langer, 2003). The medial margin is straight to slightly
sigmoid, such as in Saturnalia tupiniquim (MCP 3844-PV;
Langer, 2003) and Macrocollum itaquii (CAPPA/UFSM 0001a;
Müller et al., 2018b), whereas in Unaysaurus tolentinoi
(UFSM11069; Leal et al., 2004) and Thecodontosaurus antiquus
(BRSUG 26627; Ballell et al., 2020) it is concave. There are no
ventral flanges projecting from the proximal articular surface.
The shaft is slender and transversely compressed (Fig. 9E).
RESULTS
Phylogenetic Analysis
The first analysis (i.e., employing the data matrix of Müller,
2021) recovered 72 most parsimonious trees (MPTs) of 981
steps (consistency index = 0.321; retention index = 0.668). Near
the base of the tree, Saturnaliidae is composed of Chromogi-
saurus novasi and Saturnalia tupiniquim. The clade is supported
by an enlarged olecranon process of ulna (149 [0→1]) and the
pubic peduncle of the ilium forming an angle of less than 45
degrees to the long axis of the iliac blade (186 [1→2]).
CAPPA/UFSM 0276 nests within Bagualosauria in all MPTs
(Fig. 10A). This clade is supported by three unambiguous syna-
pomorphies: (1) ratio between the dorsoventral height and the
anteroposterior length of the dentary is greater than 0.2 (73
 Müller and Garcia—An early sauropodomorph from Brazil (e2002879-13)

FIGURE 8. Left fibula of CAPPA/UFSM 0276 in A, lateral, B, anterior, C, proximal, D, distal, E, posterior, and F, medial views. The black areas are
broken. Abbreviations: ifi, m. iliofibularis insertion; rdg, ridge.
 Müller and Garcia—An early sauropodomorph from Brazil (e2002879-14)

FIGURE 9. Left metatarsal II of CAPPA/UFSM 0276 in A, anterior, B, lateral, C, proximal, D, medial, and E, posterior views. The black areas are
broken. Abbreviations: mt I cs, metatarsal I contact surface; mt III cs, metatarsal III contact surface.

[0→1]); (2) short caudoventral process of dentary (75 [0→1]);
and (3) serrations of maxillary/dentary teeth forming oblique
angles with the margin of the tooth (88 [1→2]). Given the
absence of cranial elements, none of these features are pre-
served in CAPPA/UFSM 0276. The specimen is positioned as
the sister taxon of Bagualosaurus agudoensis, and the clade
they form is supported by two unambiguous synapomorphies:
(1) vertically oriented ischial peduncle of the ilium (179
[1→0]); and (2) absence of a distinct proximodistally oriented
ridge on the posteromedial surface of the distal portion of
the tibia (242 [1→0]). The node joining CAPPA/UFSM 0276
and Bagualosaurus agudoensis is the sister taxon of a clade
composed of post-Carnian sauropodomorphs. The remaining
topology of the strict consensus tree is equivalent to that of
Müller (2021). The Bremer support value is 1 for all the
nodes within Sauropodomorpha. CAPPA/UFSM 0276 nests
basal to Bagualosauria with one extra step. A possible sister-
taxon relationship between CAPPA/UFSM 0276 and Pampa-
dromaeus barberenai requires at least one extra step, as does
placement of CAPPA/UFSM 0276 within Saturnaliidae.
 Müller and Garcia—An early sauropodomorph from Brazil (e2002879-15)

FIGURE 10. Result of the phylogenetic analyses. A, abbreviated strict consensus tree of the first analysis (employing the data matrix of Müller, 2021)
depicting the phylogenetic position of CAPPA/UFSM 0276; B, abbreviated strict consensus tree of the second analysis (employing the data matrix of
Müller et al., 2021) depicting the phylogenetic position of CAPPA/UFSM 0276. Circles and black arrows indicate node- and branch-based clade
names, respectively.

Placement within Theropoda or Herrerasauridae requires at
least five extra steps.
The second heuristic search (i.e., employing the data matrix of
Müller et al., 2021) recovered 24 MPTs of 1555 steps (consistency
index = 0.304; retention index = 0.673). Saturnaliidae, which
includes Saturnalia tupiniquim, Chromogisaurus novasi, Pampa-
dromaeus barberenai, and Panphagia protos, is supported by
three synapomorphies: (1) a bone sheet between the anterior
and ventral processes of the prefrontal (47 [0→1]); (2) posterior
margin of the acromion process rises from the scapular blade at
an angle that is greater than 65° from the long axis of the scapula,
at its steepest point (226 [0→1]); and (3) a greatly enlarged ole-
cranon process of the ulna (243 [0→2]). In the strict consensus
tree, the node supporting CAPPA/UFSM 0276 plus
Bagualosauria nests as the sister taxon of an unresolved Saturna-
liidae. CAPPA/UFSM 0276 nests as the sister taxon of Bagual-
osauria in all MPTs (Fig. 10B), and this node preceding
Bagualosauria is supported by two unambiguous synapomor-
phies: (1) concave lateral margin of the proximal surface of the
metatarsal II (377 [0→1]); and (2) femoral length between 200
and 399 mm (404 [0→1]). Bagualosauria is supported by a ventro-
laterally oriented brevis fossa (287 [1→2]). The Bremer support
value is 1 for all the nodes within Sauropodomorpha that are
basal to Plateosauria. A possible sister-taxon relationship
between CAPPA/UFSM 0276 and Bagualosaurus agudoensis
requires one extra step, whereas a sister taxon relationship
between CAPPA/UFSM 0276 and Pampadromaeus barberenai
requires two extra steps. Placement within Saturnaliidae or
 Müller and Garcia—An early sauropodomorph from Brazil (e2002879-16)

FIGURE 11. Chrono- and biostratigraphy of the Candelária Sequence from southern Brazil, showing the pelvic girdle and hind limb of sauropodo-
morphs from each Assemblage Zone (AZ). A, Macrocollum itaquii (CAPPA/UFSM 0001b); B, indeterminate sauropodomorph (CAPPA/UFSM
0276; pes from the holotype of Bagualosaurus agudoensis); C, Buriolestes schultzi (ULBRA-PVT280; CAPPA/UFSM 0035). The radiometric ages
of 233.2 and 225.4 Ma are according to Langer et al. (2018). Silhouettes are not to scale.

Theropoda requires at least two extra steps. Placement within DISCUSSION

Herrerasauridae requires at least three extra steps.
Body Mass Estimation
Following the equation of Campione et al. (2014), which is
based on the relation between body mass and femoral circumfer-
ence ( = 66 mm in CAPPA/UFSM 0276), the resulting estimated
body mass for the specimen described here is 21 kg, with a range
of 15–26 kg applying a 25% prediction error (Campione et al.,
2014).
Taxonomic Status
CAPPA/UFSM 0276 adds to the paleofauna of the ‘Várzea do
Agudo’ site and expands the knowledge of uncommon portions
of the skeleton of Carnian dinosaurs, such as the forelimb.
Prior to CAPPA/UFSM 0276, only Eoraptor lunensis (Sereno
et al., 2013) and Buriolestes schultzi (Cabreira et al., 2016) pre-
served manual elements among Carnian sauropodomorphs, the
latter being limited to a single metacarpal. It also expands the
record of relatively large sauropodomorphs (femoral length >
200 mm) from the late Carnian. It is the second Carnian-aged
specimen known to date, the first being the holotype of Bagual-
osaurus agudoensis (Pretto et al., 2018; Table S4). Another
 Müller and Garcia—An early sauropodomorph from Brazil (e2002879-17)
potential large specimen was recently reassessed and is now con-
sidered a herrerasaurid instead of a sauropodomorph (Garcia
et al., 2021). This raises some relevant taxonomic questions
necessary for a better understanding of the diversity of this ‘tran-
sitional’ paleofauna of the ‘Várzea do Agudo’ site.
The new specimen comes from the same fossiliferous site that
yielded two other early sauropodomorphs: Pampadromaeus bar-
berenai and Bagualosaurus agudoensis. CAPPA/UFSM 0276 may
be referrable to one of these taxa, or pertain to a new taxon. Ana-
tomical and size differences exist among CAPPA/UFSM 0276,
Pampadromaeus, and Bagualosaurus, but taphonomic distortion
and ontogenetic uncertainty surrounding the specimens of
concern mean that more material and histological analyses are
necessary to further refine the taxonomic affinities of the new
skeleton.
The Early Increase in Size in Sauropodomorpha
The holotype of Bagualosaurus agudoensis is particularly
interesting because it is substantially larger than coeval or puta-
tive older sauropodomorphs. Its estimated femoral length is 22.4
cm (Pretto et al., 2018), similar to the femoral length of CAPPA/
UFSM 0276 (20.4 cm). Indeed, the femoral lengths of both speci-
mens are 40 and 30% (respectively) larger than the next-largest
sauropodomorphs of similar age (excluding skeletally immature
specimens; see Table S4), though some of those may not have
died at their asymptotic size. Therefore, both represent the
oldest evidence of body size increase within the sauropodomorph
lineage.
The earliest branching sauropodomorph Buriolestes schultzi
(Cabreira et al., 2016) is estimated to weigh ∼7 kg based on the
femoral circumference (Müller et al., 2021). Applying the same
approach, the estimated mass of CAPPA/UFSM 0276 is three
times heavier at ∼21 kg. Below, we explore how this mass
increase did or did not affect various aspects of skeletal
anatomy within Sauropodomorpha.
The hindlimb zeugopodium/stylopodium length ratio has been
traditionally employed in phylogenetic studies of early dinosaurs
(e.g., Gauthier, 1986; Nesbitt, 2011; Cabreira et al., 2016; Baron
et al., 2017). The typical condition in the earliest sauropodo-
morphs is to have hindlimb stylopodia and zeugopodia similar
in length (Fig. 11C; Table S6). Conversely, more derived sauropo-
domorphs usually have hindlimb zeugopodia shorter than stylo-
podia (Fig. 11A). Curiously, despite its relatively larger size,
CAPPA/UFSM 0276 retains the plesiomorphic condition,
where the hindlimb zeugopodium is subequal in length to the sty-
lopodium (Fig. 11B; Table S6). The ratio is unknown in Bagual-
osaurus. In large herrerasaurids, the hindlimb stylopodium is
longer than the zeugopodium (Novas, 1993; Pacheco et al.,
2019; Table S6), whereas in the smaller herrerasaurid Stauriko-
saurus pricei (MCZ 1669), the hindlimb stylopodium is shorter
than the zeugopodium. The latter condition is frequently
regarded as indicative of cursoriality (Persons and Currie, 2016).
The presence or absence of a trochanteric shelf on the femur is
another trait that has commonly been employed in phylogenetic
studies of early dinosaurs (e.g., Gauthier, 1986; Nesbitt, 2011;
Cabreira et al., 2016). This structure is usually regarded as the
insertion point of m. iliofemoralis externus (Hutchinson, 2001;
Langer, 2003; Piechowski and Tałanda, 2020) and occurs in
several non-dinosaur dinosauriforms and saurischian dinosaurs
(Nesbitt, 2011). In Saurischia, it is present in some herrerasaurids
(Novas, 1993; Alcober and Martinez, 2010; Pacheco et al., 2019),
some theropods (Griffin, 2018), and sauropodomorphs from the
Carnian (Langer, 2003; Ezcurra, 2010; Cabreira et al., 2016).
Conversely, the trochanteric shelf is absent in post-Carnian saur-
opodomorphs, such as Macrocollum itaquii (Fig. 11) and Theco-
dontosaurus antiquus (Müller et al., 2018b; Ballell et al., 2020).
Therefore, its presence is regarded as a plesiomorphic feature
in CAPPA/UFSM 0276. The relationship between the absence
of the trochanteric shelf and the body size increase in sauropodo-
morphs is clear; i.e., there are no large sauropodomorphs with
trochanteric shelf. However, as observed in CAPPA/UFSM
0276, the loss of this feature did not occur.
Another signature of increased body size that might be
expected to occur in the hind limb of CAPPA/UFSM 0276 is
an increase in limb robustness, which is observed in sauropodo-
morphs from younger Upper Triassic beds (Ezcurra and Apal-
detti, 2012). However, the Robustness Index (RI, sensu Wilson
and Upchurch, 2003; i.e., average of the greatest widths of the
proximal end, midshaft, and distal end of the element divided
by the length of the element) obtained for CAPPA/UFSM 0276
does not differ from those of smaller sauropodomorphs (Table
S6). Both the tibia and fibula are quite slender with indices com-
parable to those of Eoraptor lunensis. Even the indices obtained
for Macrocollum itaquii are not distinct from those of smaller
forms. Conversely, the RI of the tibia of PULR 136, a large
(tibia length = 40.5 cm) indeterminate sauropodomorph from
upper Norian–Rhaetian beds of Argentina (Ezcurra and Apal-
detti, 2012), is 0.33, which is roughly twice as great as the RI
for any Carnian–early Norian sauropodomorph. Hence, body
size and hind limb robustness do not appear to have increased
in a coordinated fashion in the early evolutionary history of saur-
opodomorphs; only at larger body size thresholds is the hindlimb
robustness of early sauropodomorphs markedly increased above
plesiomorphic values. The gracile construction of the pes (pha-
langes much longer than wide) of both Bagualosaurus agudoensis
(Pretto et al., 2018) and Macrocollum itaquii (Müller et al.,
2018b) reinforces this point (Fig. 11). In larger sauropodo-
morphs, such as Eucnemesaurus entaxonis (BP/1/6234; McPhee
et al., 2015) and Coloradisaurus brevis (PVL 5904; Apaldetti
et al., 2013), the phalanges are stouter.
CONCLUSIONS
Despite its indeterminate taxonomic status, the new early saur-
opodomorph specimen described herein (CAPPA/UFSM 0276)
expands the fossil record between the mid-to-late Carnian, fauni-
vorous, small-bodied, and short-necked members of the clade
(e.g., Buriolestes schultzi) and the early Norian sauropodo-
morphs with herbivorous/omnivorous adaptations, relatively
large body sizes, and elongated necks (e.g., Macrocollum
itaquii). The new specimen represents one of the oldest examples
of body increase size within Sauropodomorpha. Whereas the
skeleton of CAPPA/UFSM 0276 is significantly larger than that
of nearly all coeval or older forms, its proportions (hindlimb zeu-
gopodium subequal to stylopodium in length; low robustness
indices) and anatomy (presence of a trochanteric shelf on the
femur) resemble that of these earliest forms. Hence, despite its
proportionally large body size, the new specimen retains a con-
servative postcranial anatomy. Alternatively, CAPPA/UFSM
0276 may exist at the large extreme of its ontogenetic trajectory,
whereas other known specimens of coeval sauropodomorphs
may represent not fully grown individuals. Such a scenario
would indicate a larger ancestral body size for Sauropodomorpha
than is presently hypothesized.
ACKNOWLEDGMENTS
We thank C. Peyre de Fabrègues and an anonymous reviewer
for corrections and comments that greatly improved this manu-
script. We also extend our gratitude to the editor
M. D. D’Emic for corrections and suggestions. We thank the
Coordenação de Aperfeiçoamento de Pessoal de Nível Superior
(CAPES) scholarship for M.S.G. We also thank the Willi
Henning Society, for the gratuity of TNT software. This work
 Müller and Garcia—An early sauropodomorph from Brazil (e2002879-18)
was carried out with the aid of the Fundação de Amparo à Pes-
quisa do Estado do Rio Grande do Sul (FAPERGS 21/2551-
0000680-3) to R. T. Müller.
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Submitted April 15, 2021; revisions received October 14, 2021;
accepted October 17, 2021.
Handling Editor: Michael D’Emic.