Journal Pre-proof

New theropod dinosaur from the late Cretaceous of Brazil improves abelisaurid
diversity

Fabiano Vidoi Iori, Hermínio Ismael de Araújo-Júnior, Sandra A. Simionato Tavares,

Thiago da Silva Marinho, Agustín G. Martinelli

PII: S0895-9811(21)00397-7
DOI: https://doi.org/10.1016/j.jsames.2021.103551
Reference: SAMES 103551

To appear in: Journal of South American Earth Sciences

Received Date: 24 July 2021

Revised Date: 1 September 2021
Accepted Date: 1 September 2021

Please cite this article as: Iori, F.V., Ismael de Araújo-Júnior, Hermí., Simionato Tavares, S.A., da Silva
Marinho, T., Martinelli, Agustí.G., New theropod dinosaur from the late Cretaceous of Brazil improves
abelisaurid diversity, Journal of South American Earth Sciences (2021), doi: https://doi.org/10.1016/
j.jsames.2021.103551.

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Author statements

Conceptualization: FVI; Investigation: all authors; Writing - original draft: all authors; Writing - review
& editing: FVI and AGM.

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 NEW THEROPOD DINOSAUR FROM THE LATE CRETACEOUS OF BRAZIL
IMPROVES ABELISAURID DIVERSITY

Fabiano Vidoi Iori1,2*, Hermínio Ismael de Araújo-Júnior3, Sandra A. Simionato

Tavares2, Thiago da Silva Marinho4,5 and Agustín G. Martinelli6

1
Museu de Paleontologia “Pedro Candolo”, Estação Cultura, Praça Farmacêutico Bruno
Garisto, 15890-000, Uchoa, SP, Brazil.

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2
Museu de Paleontologia “Prof. Antônio Celso de Arruda Campos”, Centro de Artes,

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Praça do Centenário, 15910-000, Monte Alto, SP, Brazil.

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Departamento de Estratigrafia e Paleontologia, Faculdade de Geologia, Universidade
do Estado do Rio de Janeiro, Rua São Francisco Xavier, 524, 20550-013, Maracanã,
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Rio de Janeiro, RJ, Brazil.
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4
Centro de Pesquisas Paleontológicas L. I. Price, Complexo Cultural e Científico
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Peirópolis, Pró-Reitoria de Extensão Universitária, Universidade Federal do Triangulo

Mineiro, Uberaba, Minas Gerais, Brazil.
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5
Instituto de Ciências Exatas, Naturais e Educação (ICENE), Universidade Federal do
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Triângulo Mineiro (UFTM), Av. Randolfo Borges Jr. 1400, Univerdecidade, 38064-
200, Uberaba, MG, Brazil.
6
CONICET-Sección Paleontología de Vertebrados, Museo Argentino de Ciencias
Naturales “Bernardino Rivadavia”, Av. Ángel Gallardo 470, Buenos Aires, C1405DJR,
Argentina.

*Corresponding author: biano.iori@gmail.com (FVI)

Author ORCID’s

Fabiano V. Iori - ORCID: 0000-0003-2211-2276

Hermínio I. Araújo-Júnior - ORCID: 0000-0003-4371-0611

Sandra A. S. Tavares - ORCID: 0000-0001-7262-7548

Thiago S. Marinho - ORCID: 0000-0002-2754-4847

Agustín G. Martinelli - ORCID: 0000-0003-4489-0888

Abstract

The Late Cretaceous Marília Formation (Bauru Group, Bauru Basin) is a

geological unit that occurs on São Paulo, Minas Gerais, Goiás, and Mato Grosso do Sul

states, Brazil. This formation consists predominantly of paleosols developed in a

semiarid/arid environment and recent reappraisal of its formerly known members

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reduced its lithological composition and geographical distribution. Hence, the Marília

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Formation has a very sparse vertebrate fossil record without named species so far. In
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this contribution we describe a new abelisaurid theropod (Dinosauria) from this unit,
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namely Kurupi itaata gen. et sp. nov., discovery in the Municipality of Monte Alto,
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western São Paulo State. The holotype MPMA 27-0001/02 consists of three caudal
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vertebrae and the partial pelvic girdle. Kurupi itaata gen. et sp. nov. shares with other

South American abelisaurids fused ischia and caudal vertebrae with long and
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laterodorsally oriented transverse processes, with fan-shaped distal ends.

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Autapomorphies of the new theropod include the variation of ~15º of the inclination of

the transverse process between the first (C1) and seventh (C7) caudal vertebra; C1 with

anterolaterally projected triangular process placed below the proximodistal mid-portion

of the transverse process, and a notch at the anterodistal portion of the transverse

process, between a shelf-like process and the anterodistal corner of the transverse

process; and anterior caudal vertebrae (present in C1 and C7, inferred in the others) with

a cuneiform process, anterodorsally projected, located on the dorsal surface of the

transverse process. Phylogenetic analysis recovered Kurupi itaata gen. et sp. nov.

among abelisaurid theropods, but was nested in an unresolved massive polytomy of the
entire clade. Taphonomic traits on the studied specimens corroborate previous proposals

for the paleoenvironmental context of the Marília Formation. Kurupi itaata gen. et sp.

nov. was about 5 meters long, with a rigid tail, and cursorial locomotion as indicated by

its a muscles attachment and bones anatomy. This new taxon contributes to the

knowledge of the Maastrichtian continental fauna of Brazil and increases the diversity

of medium-sized abelisaurids in western Gondwana.

Keywords: Theropoda, Abelisauridae, Marília Formation, Maastrichtian.

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1. Introduction

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Saurischian dinosaurs and notosuchian crocodyliforms were the dominant
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tetrapod groups in the Late Cretaceous ecosystems of southeastern Brazil. The Late
Cretaceous record of this region is mainly concentrated in rocks of the Bauru Basin,
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encompassing the Caiuá and Bauru groups (e.g., Menegazzo et al., 2016; Langer et al.,
2019). Although the basin has a relative abundance of titanosaur dinosaurs,
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summarizing up to ten species (e.g., Uberabatitan riberoi, Arrudatitan maximus,

Maxakalisaurus topai) and hundreds of isolated records, theropods are rare, and
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ornithischians are absent. Theropod record includes mostly isolated and fragmentary
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teeth and bones which were assigned to Abelisauroidea, Megaraptora, and

Maniraptoriformes (e.g., Novas et al., 2005, 2008; Machado et al., 2008, 2013; Méndez
et al., 2012, 2014; Martinelli et al., 2013; Tavares et al., 2014; Delcourt and Grilo, 2014;
Brum et al., 2016, 2018; Delcourt and Iori, 2020). Only two species were formally
named: the noasaurid Vespersaurus paranaensis from the Rio Paraná Formation (Caiuá
Group; Langer et al., 2019), and the abelisaurid Thanos simonattoi from the São José do
Rio Preto Formation (Bauru Group; Delcourt and Iori, 2020). Other Cretaceous
abelisauroids from Brazil are Spectrovenator ragei from the mid-Cretaceous Quiricó
Formation (Sanfranciscana Basin; Zaher et al., 2020) and Pycnonemosaurus nevesi
from Mato Grosso state (central Brazil; Kellner and Campos, 2002; Bittencourt and
Langer, 2011), probably correlated to the Bauru Group (Kellner and Campos, 2002;
Sales et al., 2018). Theropods were considered as less abundant in the Bauru rocks due
to competitive factors with some notosuchian groups (e.g., baurusuchids; Riff and
Kellner, 2011); however, taphonomic biases and a still very incomplete, but growing
up, fossil record may hamper the real diversity of theropods in the Late Cretaceous of
southeastern Brazil (Martinelli et al., 2013; Bandeira et al., 2018).

One relevant place for southeastern Brazilian Late Cretaceous fossils is the
Monte Alto region, located near the oriental border of the Bauru Basin, in the state of
São Paulo. Fossils from this region have been collected in sandstones of the
Adamantina and Marília formations of the Bauru Group, including not only a high
diversity of tetrapods but also mollusks, fishes, coprolites and other ichnofossils (see
summary in Iori, 2019). The tetrapods are the most widely studied records, with several
described species for the Adamantina Formation, including the testudines Yuraramirim

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montealtensis and Roxochelys wanderleyi, the squamate Boipeba tayasuensis, the
crocodilyforms Stratiotosuchus maxhechti, Montealtosuchus arrudacamposi,

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Caipirasuchus montealtensis, Caipirasuchus paulistanus, Morrinhosuchus luzie and
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Barreirosuchus franciscoi, and the titanosaur dinosaur Arrudatitan maximus, plus
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several unnamed tetrapod records (e.g., Carvalho et al., 2007; Andrade and Bertini,
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2008; Pinheiro et al., 2008; Iori and Carvalho, 2009, 2011; Iori and Garcia, 2012;
Santucci and Arruda-Campos, 2011; Ferreira et al., 2018; Fachini et al., 2020, Silva-
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Junior et al., 2021). In contrast, the tetrapod fossil record from the Marília Formation is
sparse (e.g., Bertini et al., 2001; Ragonha and Mezzalira, 1995; Méndez et al., 2014; Iori
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and Arruda-Campos, 2016; Iori, 2019), which limits the taxonomic assignations,
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without named species.

Fossils of theropod dinosaurs are very uncommon in the Monte Alto region.
Roxo (1929) cited the presence of Ceratosauria, however the specimens were not
illustrated and their currently location is unknown; they may be lost. Tavares et al.
(2014) analyzed isolated teeth of abelisaurids and dromeosaurids from the Adamantina
Formation, which were discovered in association to the holotype specimen of the
titanosaur Arrudatitan maximus. Méndez et al. (2014) studied the bones of the pelvis
(specimen MPMA 27-0001/02) of an indeterminate abelisauroid from the Marília
Formation. In this contribution, we study new elements collected in the last years that
belong to the specimen MPMA 27-0001/02 (Méndez et al., 2014) allowing to improve
its taxonomy and phylogenetic relationships. All bone elements referred to this
specimen are fully described and constitute the holotype of a new genus and species of
abelisaurid theropod for the Late Cretaceous of southeastern Brazil.
Institutional Abbreviations — MPMA, Museu de Paleontologia “Prof. Antonio Celso
de Arruda Campos”, Monte Alto, State of São Paulo, Brazil.

2. Geological setting

The region of Monte Alto Municipality (São Paulo State, southeastern Brazil) is
highlighted by the presence of a hill, called “Serra do Jaboticabal”. The hard calciferous
sandstones of the Marília Formation and their slower weathering are responsible for the
existence of the Monte Alto’s plateau and preservation of the lower strata, as was

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pointed out by Paula e Silva et al. (2005) for Marília, Garça and Echaporã highlands,
along the state. The Monte Alto city was building on that plateau and lies over the

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sandstones of the Marília Formation, so the paleontological research sites of this unit
are very close to the urban area (Fig. 1; see also Iori, 2019).
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The “Sítio Paleontológico dos Gaviões” (i.e., “Hawk's paleontological site”) is
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less than one kilometer from the city. It was discovered in 2002 and yielded an isolated
theropod pelvis that was removed and then studied by Méndez et al. (2014). Other four
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excavations carried out during 2006, 2009, 2013 and 2014 in the same quarry (Fig. 1),
an area of approximately 10 m2, resulted in the discovery of more elements that are here
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described.
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The fossil-bearing matrix is predominantly pinkish beige, composed by

sandstones poorly selected and with intense carbonate cementation, common features of
the Echaporã Member of the Marília Formation (see Fernandes, 2004). There are
laminar crystallized layers along the sequence, which are also observed in some of the
fossils (Fig. 3).

The Bauru Basin is composed by Caiuá and Bauru groups. The latter one is
extended over São Paulo, Goiás, Mato Grosso do Sul and Minas Gerais states. The
upper unit of the Bauru Group is the Marília Formation, which was traditionally divided
in the Echaporã, Serra da Galga, and Ponte Alta members (Fernandes, 2004). Recently,
Soares et al. (2021) unified the Serra da Galga and Ponte Alta members as the Serra da
Galga Formation, exclusively exposed in the east portion of the Triângulo Mineiro
region (Minas Gerais State), and consequently, the Marília Formation was reduced to
the area of occurrence of the Echaporã Member. The deposition of the Bauru Group
occurred under semi-arid to arid climatic conditions, between the Coniacian and
Maastrichtian ages (Fernandes and Coimbra, 2000). The exact age of the formations
included within the Bauru Group is not consensual, however in relation to Marília
Formation there seems to be an agreement. Dias-Brito et al. (2001) suggested a
Maastrichtian age based on micropaleontological studies. Menegazzo et al. (2016)
corroborated the age with faunal comparisons, including mollusks, fishes and tetrapods.
However, the majority of species used to formerly establish the relative dating of
Marília Formation come from the deposits now recognized as the new unit Serra da
Galga Formation. Hence, considering the lateral relationships among the three formerly

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known members, a similar age is assumed.

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3. Material and methods -p
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3.1. Specimen. The first element MPMA 27-0001/02 was discovered in 2002,
consisting of an incomplete pelvis, with the almost complete left ischium articulated
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with parts of right ischium, left ilium and an inexpressive portion of the left pubis.
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Subsequent excavations at the quarry permitted to rescue three caudal vertebrae and
other undefined bones. All material was included within the same specimen number and
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is housed at the “Museu de Paleontologia Prof. Antonio Celso de Arruda Campos”

(Monte Alto, São Paulo, Brazil).
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3.2. Taphonomy. The criteria used in the taphonomic analysis follow Hill (1980),
Shipman (1981), Behrensmeyer (1978, 1991), Lyman (1994, 2008), Rogers (1994),
Holz and Simões (2002), Eberth et al. (2007), Simões et al. (2010) and Araújo-Júnior
and Bissaro-Júnior (2017). The following macroscopical features were evaluated: (i)
degree of disarticulation; (ii) types of breaks; (iii) signs of weathering; (iv) marks of
abrasion; (v) trample marks; (vi) tooth marks; (vii) invertebrate modifications; (viii)
signs of rooting; and (ix) diagenetic aspects.

3.3. Phylogenetic analysis. Specimen MPMA 27-0001/02 was included in the data
matrix of Baiano et al. (2021) in order to test its phylogenetic position among
ceratosaurian dinosaurs. This data matrix is based on Rauhut and Carrano (2016), plus
the successive modifications in the characters list, character-scorings and operational
terminal units (OTUs) applied by Langer et al. (2019), Zaher et al. (2020), and Baiano
et al. (2020, 2021). The characters 33, 103, 104, 117, 138, 154, and 165 were
considered as additive (Rauhut and Carrano, 2016; Langer et al., 2019) and the OTUs
Deltadromeus agilis, Spinostropheus gautieri, and USNM-8415 were deactivated before
the analyses, due to its conflictive taxonomic position (see Apesteguía et al., 2016;
Rauhut and Carrano, 2016; Langer et al., 2019). Hence, the data matrix includes 42
OTUs (three are inactive) and 224 morphological characters (Supplementary File 1).

In addition to the inclusion of Kurupi itaata, one character-state was changed for
Pycnonemosaurus nevesi. For character 138 (centrodiapophyseal laminae in anterior
midcaudal vertebrae) we scored state “2” (anterior and posterior centrodiapophyseal
laminae present as low, rounded ridges) instead of state “1” for this taxon, based on

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Delcourt (2017).

The data matrix was analyzed under equally weighted maximum parsimony

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using TNT v.1.5 (Goloboff et al., 2008; Goloboff and Catalano, 2016). We performed a
heuristic search of 10,000 replications of Wagner trees followed by TBR branch-
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swapping algorithm (holding 10 trees per replication) and, subsequently, a final round
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of TBR branch swapping to the best trees obtained. Zero-length branches in any of the
recovered most parsimonious trees were collapsed. Branch support was quantified using
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decay indices (Bremer support values; Bremer, 1994) and a bootstrap resampling
analysis, using 1,000 pseudoreplicates and reporting both absolute and GC frequencies
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(Goloboff et al., 2008).

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4. Taphonomic remarks

Desiccation marks were observed on the lateral surfaces of the caudal vertebrae
and ischium (Fig. 2) and are referred to the stages 1 and 2 of Behensmeyer (1978).
Taken together with the degree of disarticulation, these features suggest that the
material remained for a short time span of subaerial exposure prior to its burial
(Behrensmeyer, 1978; Hill and Behrensmeyer, 1984).

Signs of abrasion (e.g., wear marks) are absent in the sample, suggesting the
bones did not experience significant transport from the death place to the final burial
site (Shipman, 1981; Fiorillo, 1988; Lyman, 1994). The presence of low-transportable
specimens (pelvic girdle) and high-transportable ones (vertebrae) and the absence of
other heavy elements (e.g., articulated skull and jaws) reinforce the idea of short-
distance transportation (Voorhies, 1969; Frison and Todd, 1986).

Some vertebrae were fractured or deformed during the diagenetic processes,

probably by lithostatic compression (Medeiros, 2010; Sinibaldi, 2010).
Permineralization by calcite (CaCO3) is macroscopically observed on the cancellous
bone of fragmented elements (Fig. 3). The fully permineralization of the pores indicates
that the diagenetic environment was abundant in water, probably in a context of a
poorly-drained floodplain (Behrensmeyer and Hook, 1992). In some cases, fissures in
vertebrae, generated by weathering or lithostatic compression, are filled by calcite. Such
a filling seems to be the continuity of the permineralization which affected the

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cancellous bone. In this case, the overfilling by calcite can be related to the evolution of
a paleosol after the end of the flooded phase of the floodplain.

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In summary, the taphonomic history of the holotype of Kurupi itaata gen. et sp.
nov. can be interpreted as follow: (1) The individual died near its final burial place and
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experienced a short time span of subaerial exposure prior to burial. (2) After, some
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elements were transported by a short distance and then buried on a poorly-drained

floodplain. (3) During the fossildiagenesis, the bones experienced multiple events of
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lithostatic compression and permineralization by calcite in a moment of drying

floodplain. The taphonomic homogeneity of material described in this paper suggests
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that all specimens seem to belong to the same individual.

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5. Results

5.1. Systematic Paleontology

Theropoda Marsh, 1881

Ceratosauria Marsh, 1884

Abelisauroidea Bonaparte and Novas, 1985

Abelisauridae Bonaparte and Novas, 1985

Kurupi gen. nov.

ZooBank registration: urn:lsid:zoobank.org:act:0E8EECDD-ADB9-40DB-84D8-

BFEDDBEC492E
5.1.1. Type species: Kurupi itaata sp. nov.

5.1.2. Diagnosis: As for type and only known species.

5.1.3. Etymology: The generic name Kurupi refers to a legendary monster of the
Guarani indigenous culture, God of fertility and sexuality. The choice of the name is
due to the fact that the fossils were found in the region of “Motel Paraíso” (“Paradise
Motel”), a place intended for intimate encounters.

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Kurupi itaata sp. nov.

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ZooBank
89C70CCE9DA0
registration: -p
urn:lsid:zoobank.org:act:12D65CF3-A92E-424E-97BF-
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5.1.4. Holotype: MPMA 27-0001/02, three incomplete caudal vertebrae, part of pelvic
girdle and two undefined bones (The pelvic girdle was described previously by Méndez
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et al., 2014).
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5.1.5. Etymology: Specific name itaata comes from the indigenous language Tupi and
has two roots: ita for rock and atã for hard, in allusion to the very cemented rocks of the
Marília Formation at the Monte Alto region.

5.1.6. Diagnosis: Mid-sized (~5 meter long) abelisaurid theropod with the following
combination of characters not known in any other theropod (autapomorphies are marked
with an asterisk): first caudal vertebra with anterolaterally projected triangular process
placed below and at mid-length of the anterior edge of the transverse process*, first
caudal vertebra with a notch at the anterodistal portion of the transverse process
(possibly to receive the iliac blade), between a shelf-like process and the anterodistal
corner of the transverse process*; first caudal vertebra with a very concave and broad
anterior (=prezygapophyseal) centrodiapophyseal fossa; anterior caudal vertebrae
(present in C1 and C7, inferred in the others) with a cuneiform process, anterodorsally
projected, located on the dorsal surface of the transverse process, near the anterior base
of the neural spine*; increased inclination (from ~20° to ~35°) of the transverse
processes along the first third of the vertebral caudal region*; lack of medial
centrodiapophyseal fossa (=infradiapophyseal fossa) in the first caudal; presence of
reduced anterior and posterior centrodiapophyseal laminae in the seventh caudal; lack of
ischial tuberosity.

5.1.7. Locality and Horizon: “Gaviões’s Paleontological Site” (21º17’38.9” S/

48º31’09.9” W), located about 1 km southwest of the urban area of the Monte Alto City,

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São Paulo State, Brazil. Marília Formation, Bauru Group, Bauru Basin, Upper

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Cretaceous (Maastrichtian).

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5.2. Description and comparison
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The specimen MPMA 27-0001/02 is a somatically mature individual on the basis of
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fused bones of the pelvic, also supported by the absence of neurocentral suture between
the centrum and neural arch in the caudal vertebrae. One neural arch was found
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separated of its respective centrum by means of a fracture that occurred ventral and
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slightly oblique to centrum-neural arch suture, allowing to joint both pieces after
preparation.
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5.2.1. Caudal vertebrae. Three vertebrae of the anterior portion of the caudal series
are preserved. They are considered as C1, C5 and C7 vertebrae (Figs. 4, 5, 6, 7, 8, 9, 10
and 11).

5.2.1.1. C1. This element was broken by means of an oblique fracture that goes from
the one third dorsal portion of anterior articular surface of the centrum to the base of the
hyposphene, fossilizing separately both portions. It also lacks the distal portion of both
transverse processes. The centrum is anteroposteriorly long (88mm) and transversely
wide (81mm at its posterior articular surface); it is spool-shaped, lacking pleurocels,
facets for the chevron, and a ventral groove (Fig. 5A). The anterior articular facet is
slightly concave and sub-quadrangular in outline due to the slightly straight lower
border, while the caudal articular surface is almost flat and circular in outline (Figs. 2A
and 2B). The neural arch has robust and transversely wide bases (Fig. 2C), delimiting a
small neural canal. The transverse processes lack their distal portion; they are
anteroposteriorly broad and well expanded laterodorsally. In dorsal view, the preserved
portions of the transverse process have a rectangular outline and are caudally oriented.
The posterodistal corner of the right process is placed slightly posterior to the level of
the centrum border. In relation to dorsal inclination, the transverse processes are almost
plane, forming an angle of ~20º with the horizontal line, in posterior view. The ventral
surface of the transverse processes is not well preserved; however, it is possible to
observe that the first proximal third is flat and the medial centrodiapophyseal fossa
(=infradiapophyseal fossa) is absent in the C1 of Kurupi.

The anterior centrodiapophyseal lamina delimits a large fossa (i.e., anterior or

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prezygapophyseal centrodiapophyseal fossa, or infraprezygaphophyseal fossa) which is
rimed by the anterolaterally projected triangular process (altp) of the transverse process,

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the prezygadiapophyseal lamina and the hypantrum. The fossa is oval-shaped and very
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deep, with ~50 mm in its major axis, facing anterolaterally (Fig. 6). The posterior
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centrodiapophyseal lamina faces posteriorly and defines an elliptical fossa (i.e.,
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posterior centrodiapophyseal fossa or infrapostzygapophyseal fossa), vertically oriented

and with ~10 mm in its major axis (Fig. 6).
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In the anteroventral portion of each transverse process there is a conspicuous

anterolaterally projected triangular process (altp), which is clearly seen in anterior view
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(Figs. 4, 7). These processes can be related to attachment areas for connection with the
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sacrum and/or iliac blade. The ventral angle of this process corresponds to the point
where the distal portion of the anterior centrodiapophyseal lamina anchors.

Between the anterolaterally projected triangular process and the body of the
transverse process there is a shelf-like process, roughly rectangular in dorsal view,
which defines a notch with the anterior margin of the transverse process. The shelf-like
process protrudes beyond the anterior margin of the transverse process and probably
overlapped the medial portion of the caudal iliac margin. Certainly, the anterior process
of the distal portion of the transverse processes of Aucasaurus and Carnotaurus
laterally embrace the ilia (Coria et al., 2002), but, differing from Kurupi, they are more
strongly laterodorsally projected. As a consequence, the notch seen in Kuripi can be an
intermediate condition to that present in Aucasaurus and Carnotaurus.
 The dorsal surface of the transverse processes also bears over its anteromedial
portion a conspicuous cuneiform process, which is anterodorsally projected (Fig. 4A,
C). This process is well preserved on the right side of the vertebra (Fig. 4A) and can be
seen also in C7.

The prezygapophyses and postzygapophyses are incomplete, only preserving

their bases. Both prezygapophyseal bases are connected by a faint and short crest. In
addition, a crest connects to the base of each prezygapophysis to the anterior edge of the
transverse process. These crests form the anterior limit of the prespinal fossa. The
hypantrum-hyposphene articulation is well-developed. The lateral surface of the
hyposphene is slightly concave and bears a tiny depression near its anterior edge.

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The neural spine preserves only its proximal portion; it is a laminar structure,
transversely narrow and anteroposteriorly long. The prespinal fossa is relative wide and

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divided by a septum which is formed by the base of the neural spine. The left fossa is
better preserved, being diamond-shaped. The postspinal fossa is deep but transversely
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narrower than the prespinal one. It is twice high than wide, slight oval-shaped and faces
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posterodorsally. This fossa is ventrally rimed by the Y-shaped

hiposphene+postzygapophysis complex.
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5.2.1.2. C5 and C7. The fifth caudal vertebra is partially preserved, including the
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posterior half of the centrum and the neural arch with most of the transverse processes
and neural spine broken off (Fig. 8), whereas the seventh caudal only lacks the left
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transverse process (Fig. 9). The morphology of the fifth caudal vertebra (Fig. 8) is
similar to the seventh vertebra (C7, Fig. 9), but having a more conspicuous medial
ventral groove in the region for articulation with the chevron (Fig. 5B and 8C), larger
hyposphene and postzygapophyses, and less developed posterior centrodiapophyseal
fossa. Although incomplete, the bases of the transverse processes seem to be more
laterally than laterodorsally projected, with a wider angle than in C7, similar to C1. The
posterior articular surface of the centrum of C5 is subcircular, slightly taller than wide.
Its maximum width is 63 mm, whereas the anterior articular facet of the inferred C7 is
60 mm. The inclination of the transverse processes and its measurements suggest that
these vertebrae were not adjacent elements and at least one vertebra should be present
between them. Méndez (2014) stated that, in lateral view, the first four caudal centra in
Abelisauridae exhibit a convex posterior outline. In the element here described of
Kurupi the outline is more straight than convex, thus it is interpreted as the probably 5th
vertebra of the caudal series, and the other one as the probably 7th vertebra.

Both vertebral centra are similar to that of other abelisaurids (e.g., Bonaparte et
al., 1990; Kellner and Campos, 2002; O’Connor, 2007; Méndez, 2014, Filippi et al.,
2016); they are spool-shaped, amphicoelous, medially compressed and without
pneumatic foramina over the lateral surfaces. The articular facets are rounded; however,
the posterior facet of C7 is slight oval-shaped (probably by its more posterior position in
the series), as can be seen in the most complete caudal series of Pycnonemosaurus and
Aucasaurus but very distinct from those of Arcovenator, Majungasaurus and
Rajasaurus, which have an elliptical outline, well-defined in anterior and posterior

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surfaces (Wilson et al., 2003, O’Connor, 2007; Méndez, 2014; Tortosa et al., 2014;
Delcourt, 2017). Similar to the C1, the anterior facet of C7 is deeper (more concave)

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than the posterior one. -p
The median groove in the ventral surface of the centra is very subtle, except
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between the two pedicels for the articulation of the chevron (Fig. 5B). A median ventral
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groove is also observed in Aucasaurus, Majungasaurus, Ilokelesia, Viavenator, whereas

Ekrixinatosaurus and Rajasaurus have a median keel (Coria and Salgado, 1998;
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Méndez, 2014; Fillipi et al., 2018).

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The angle of inclination of the transverse process in C7 is ~35º with respect to

the horizontal plane and the ratio TPL/CL (length of transverse process/length of
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centrum) is higher than 1.3, typical of brachyrostran species (see Méndez, 2014). The
major angle (transverse process/horizontal plane) among abelisaurids occur in
Carnotaurus, Aucasaurus and Ekrixinatosaurus, with angle reaching more than 40°
(Méndez, 2014). In Pycnonemosaurus, Skorpiovenator, Viavenator, and also in Kurupi,
the inclination ranges between 30º and 40º (Méndez, 2014; Fillipi et al., 2016; Delcourt,
2017). The transverse process of the caudals of Ilokelesia is laterally projected, while in
Arcovenator, Majungasaurus and Rahiolisaurus the inclination is about 20º (Coria and
Salgado, 1998; O’Connor, 2007; Novas et al., 2010).

The ventral surface of the transverse process of C7 is convex, except in its distal
portion (insertion area of M. ilio-ischiocaudalis), which precedes the bone ridge of the
fan-shaped distal projection, where is almost plane (Fig. 11). The anterior and posterior
centrodiapophyseal laminae are low, forming rounded ridges that are most evident in
the proximal third of the transverse process, where the laminae delimit the three fossae
on the lateral surface (Fig. 6B), as observed in Pycnonemosaurus (Delcourt, 2017), but
differing from the robust laminae of Ekrixinatosaurus (Calvo et al., 2004). The
infradiapophyseal fossa (=medial centrodiapophyseal fossa) is triangular shaped and
relatively shallow, with its deepest point at its posterodorsal surface. The anterior
centrodiapophyseal fossa (=infraprezygapophyseal fossa) in C7 is shallower than in C1
(Fig. 6). The posterior centrodiapophyseal fossa (=infrapostzygapophyseal fossa) is
placed ventromedial to the base of the postzygapophyses and faces almost posteriorly
(Fig. 6).

The fan-shaped projection of traverse process is about twice longer than the base

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of the neural arch. In dorsal view, the anterior process of the distal end of the transverse
process is acute, with the lateral edge gently convex, as in Viavenator and Carnotaurus

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(Bonaparte et al., 1990; Filippi et al., 2017). In contrast, the lateral edge of the
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transverse process is less convex in the anterior portion of the tail of Pycnonemosaurus,
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based on the preserved large and medium-sized fragments of transverse processes of
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this taxon (Delcourt, 2017). In Kurupi a subtle scar forms a septum which separate two
muscle areas that would be for the M. ilio-ischiocaudalis and the M. longissimus (see
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Fig. 11B).

The hyposphene-hypantrum articulation in the caudals of Kurupi is well-

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developed. In posterior view of C5 and C7, the hyposphene and postzygapophyses have
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a Y-shaped. As in C1, the lateral surface of the hyposphene is subtle concave with a
circular depression on the central of the lamina (Fig. 6). The articular facets of the
prezygapophysis and postzygapophysis are oriented about 40º with the horizontal plane
and are oval-shaped. In C7, the articular surfaces of the prezygapophyses face
posteromedially and are continuous with the walls of the prespinal fossa; medially they
are separated by the hypantrum. The hyposphene-hypantrum complex does not occur in
caudal vertebrae of Majungasaurus and Arcovenator (O’Connor, 2007; Tortosa et al.,
2014), but is present in Eoabelisaurus, Pycnonemosaurus, Aucasaurus, Carnotaurus,
Viavenator and Masiakasaurus (e.g., Bonaparte et al., 1990; Pol and Rauhut, 2012;
Méndez, 2014; Delcourt, 2017; Filippi et al., 2018).

A conspicuous feature seen in the C7 of Kurupi is the presence of one small

finger-like pedicle (i.e., cuneiform process) in each side of the dorsal surface of neural
arch, laterally to the prespinal fossa (Fig. 10), as in C1.
5.2.2. Pelvic Girdle. The pelvic girdle is represented by both ischia and the right
acetabular region including small portions of the right pubis and ilium (Fig. 12). The
detailed description of this element was performed by Méndez et al. (2014); thus, it is
not repeated here. The authors compared the pelvis amongst different groups of
theropods and found numerous features typical of abelisauroids (e.g., Coria et al., 2006;
Carrano and Sampson, 2008). They include the fusion of the pelvic bones (highlighted
by the ischia), the well-developed distal foot expansion of ischium, the supraacetabular
crest continuous with the brevis shelf, and the peg-and-socket morphology of the
contact between ilium and ischium.

Méndez et al. (2014) also pointed out several traits related to muscle insertion in

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the pelvic girdle of Kurupi. They include a well-developed ischiadic antitrochanter that
continues within the ilium and the obturator process continues distally into a narrow

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ridge, which extends to the middle of the ischial shaft (Figs. 12 and 13). These features
-p
together with the anatomy of the caudal vertebrae suggest a robust body plan for this
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taxon.
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5.3. Results of the phylogenetic analysis

The first phylogenetic analysis recovered 172,224 most parsimonious trees

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(MPTs) of 477 steps, with a consistency index (CI) of 0.503 and a retention index (RI)
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of 0.731. The strict consensus tree (SCT) resulted in two massive polytomies within
Abelisauroidea, one representing noasaurids and the other abelisaurids, with Kurupi
itaata nested in the latter one (see Supplementary File 2). The majority rule (50%) tree
better resolves noasaurids and recovered Eoabelisaurus and Spectrovenator as
successive sister taxa of a massive polytomy of abelisaurids, in which only the clade
Ilokelesia plus Ekrixinatosaurus is formed. The frequency of group tree shows a value
of 13 for a polytomic clade that include Kurupi, Pycnonemosaurus, MPCN-PV-69,
Rajasaurus, Majungasaurus and Indosaurus, with Abelisaurus as sister taxon (see
Supplementary File 2).

The iterPCR protocol recognized Rahiolisaurus, Aucasaurus, MPCN-PV-69 and

Kurupi as rouge taxa. Therefore, the data matrix was reanalyzed excluding the first
three taxa and it resulted in 5,724 MPTs of 460 steps (Ci = 0.530; Ri = 0.759). The SCT
recovered Eoabelisaurus and Spectrovenator as successive sister taxa of a massive
polytomy of more inclusive abelisaurids (Fig. 14A). Bremer supports values are low
(=1) for Abelisauroidea, Noasauridae, Abelisauridae and its two more inclusive clades.
The majority rule (50%) resolved two more clade within abelisaurids (Fig. 14B), with a
polytomy of Rugops, Genusaurus, Dahalokely and a more inclusive unresolved clade.
The inclusion of Kurupi in this analysis supports its abelisaurid affinities rather than
noasaurid ones, but a better resolution is not obtained. Seven common synapomorphies
support the more inclusive abelisaurid clade (the large polytomy) in the strict consensus
tree [characters 45(0→1), 112(0→1), 136(0→1), 138 (1→2), 141(0→1), 167(0→1),
223 (0→1)] and only one (Character 138: 2, anterior and posterior centrodiapophyseal
laminae present as low, rounded ridges, in anterior mid-caudal vertebrae) is known for

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Kurupi (see Supplementary File 2).

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6. Discussion -p
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The Bauru Basin has been an important source to understand the Late
Cretaceous biota of southwestern Brazil for over one century. In particular, the Bauru
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Group revealed dozens of new species of tetrapods, including mainly testudines,

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squamates, crocodyliforms and dinosaurs (see Menegazzo et al., 2016). The hierarchical
organization of the units of the Bauru Group is not yet consensual, and the Marília
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Formation, described by Almeida and Barbosa (1953) and formalized by Soares et al.,
(1980), has been recently emended, solely enclosing its formerly Echaporã Member
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whereas the Serra da Galga and Ponte Alta members are now considered as the Serra da
Galga Formation, restricted to Minas Gerais state (see Soares et al., 2021). Thus, most
of the formerly known tetrapod content of the Serra da Galga Member of the Marília
Formation is now part of the Serra da Galga Formation, reducing considerably the
faunal diversity of the Marília Formation as it is now defined.

With the new proposal of the Marília and Serra da Galga formations (see Table
1), the former unit lacks any vertebrate species described up to now. Kurupi itaata
represents the first dinosaur formally nominated for the Marília Formation and the first
non-avian theropod for São Paulo State. The first paleontological prospection in the
region occurred in 1918 in deposits of the Marília Formation. The first discovered
fossils included sauropod remains from the “Tabarana” Paleontological Site (Iori,
2019). Other regional paleontological sites (PS) of the Marília Formation revealed
important materials. In the “Campestre” PS, the excavations lasted for almost a decade
(1985 to 1994) and revealed almost exclusively titanosaur bones, besides Bivalvia
indet., peirosaurid teeth, and one ctenoid scale (Bertini et al., 2001; Iori, 2019). From
“Água Limpa” PS, Bivalvia specimens are reported and assigned to the genera Iridina
and Anodontites, and a cranial fragment attributed to a peirosaurid crocodyliform
(Ragonha and Mezzalira, 1995; Iori and Arruda-Campos, 2016).

In addition to these reports, the vertebrate record from the Marília Formation
mostly includes briefly descriptions presented in abstract meetings (see Table 1). They
include isolated theropod teeth, a partial skeleton of a titanosaur sauropod and cranial
remains of a sphagesaurid crocodyliform discovered near the city of Marília, São Paulo

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State (Dezordi et al., 2013; Nava, pers. comm.), remains of a titanosaur sauropod
discovered in the municipality of Campina Verde, Minas Gerais State (Riff et al., 2013),

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and fragmentary fossils of testudinate, crocodyliforms and titanosaur from Rio Verde
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and Quirinópolis municipalities, Goiás State (Candeiro et al., 2018). As summarized in
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Table 1, the Marília Formation has a numerically low fossil record when compared with
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the Serra da Galga Formation, although the fossiliferous potential is seen in both units.
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6.1. Kurupi itaata, a new Maastrichtian Abelisauridae

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The knowledge about theropod dinosaurs from the Bauru Basin is extremely
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discrete when compared with coeval Late Cretaceous units from Patagonia Argentina
(e.g., Bonaparte et al., 1990; Coria et al., 2002; Canale et al., 2009; Novas et al., 2013).
Most brazilian records are based on isolated specimens that lack diagnostic traits to
allow more accurate taxonomic classifications. With the exception of the noasaurid
Vespersaurus paranaensis (Langer et al., 2019), the occurrence of abelisauroids in the
Basin is restricted to isolated bones and teeth (Novas et al., 2008; Machado et al., 2008,
2013; Méndez et al., 2014; Tavares et al., 2014; Brum et al., 2016, 2018; Delcourt and
Iori, 2020; Martinelli et al., 2019); even the holotype of Thanos simonattoi consists of
only an isolated axis and intercentrum (Delcourt and Iori, 2020). Better known brazilian
abelisauroids are the Late Cretaceous Pycnonemosaurus nevesi from Mato Grosso
(Kellner and Campos, 2002; Delcourt, 2017) and Early Cretaceous Spectrovenator
ragei from Minas Gerais (Zaher et al., 2020).
 The pelvic bones of Kurupi, described in detail by Méndez et al. (2014), have
many diagnostic traits of Abelisauroidea, and indicate that it may belong to a medium-
sized Abelisauridae. They also mentioned the possibility that the pelvic girdle was of a
large noasaurid abelisauroid, based on the fact that the ischia are 37.5% smaller than
that of Carnotaurus, with anteroposteriorly narrow shaft that resembles the early
ceratosaurian Ceratosaurus and the noasaurid Masiakasaurus. However, the relative
small size of the pelvis of Kurupi in comparison to that of Carnotaurus is not indicative
of noasaurid affinities and recent findings have shown a considerable size variation
among abelisaurids (e.g., Canale et al., 2016; Filippi et al., 2018; Zaher et al., 2020;
Cerroni et al., 2020; Aranciaga Rolando et al., 2021; Gianechini et al., 2021). In

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addition, the pelvis of Kurupi is more robust with strongly fused pelvic bones than in

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noasaurids (e.g., Carrano et al., 2011; Langer et al. 2019). The supraacetabular crest and

r
the base of the lateral brevis shelf are stouter when compared with noasaurids (e.g.,
-p
Carrano et al., 2011; Langer et al. 2019). The ischia have also subparallel lateral
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margins, differing from the slightly concave profile seen in the ischia of noasaurids
(e.g., Masiakasaurus; Carrano et al., 2011). The referral of caudal vertebrae to this
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pelvis adds support for abelisaurid affinities of Kurupi. The three available caudal
vertebrae of Kurupi are typical of brachyrostran abelisaurids rather than of noasaurids or
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other non-abelisaurid ceratosaurs. They have the transverse processes with the distal
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portion bearing a conspicuous anterior process and a posteriorly expanded margin.

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In Kurupi the angle of the transverse processes of the C1 is about 20°, less than
in Carnotaurus and Aucasaurus (Bonaparte et al., 1990; Coria et al., 2002), but larger
than in Skorpiovenator, which possesses nearly horizontal transverse processes in the
caudals (Canale et al., 2009). As such, the angle in the C1 of Kurupi exhibits an
intermediate state between the laterally and strongly dorsolaterally projection of the
transverse processes of abelisaurids. Also, in Kurupi the angle of the transverse
processes of C7 is more elevated than in C1, having an orientation similar to the
condition of Carnotaurus and Aucasaurus. This change in the elevation of the
transverse processes in Kuripi shows that the tendency toward higher angle of the
transverse processes does not start at the beginning of the caudal vertebrae and the
vertebra C1 of Kurupi seems to have a function more related to the establishment of the
rigid tail than to gain in muscle mass of the M. caudofemoralis.
 The available material of Kurupi is quite incomplete compared to some other
South American abelisaurids (e.g., Carnotaurus, Skorpiovenator) but the combination
of traits supports the recognition of a new genus and species. Autapomorphies for
Kuripi are recognized in the first caudal vertebra, including an anterolaterally projected
triangular process placed below the proximodistal mid-length of the transverse process,
a notch at the anterodistal portion of the transverse process (possibly to receive the iliac
blade), a very concave and broad anterior centrodiapophyseal fossa
(=infraprezygaphophyseal fossa), and cuneiform process, anterodorsally projected, that
raises over the anteroproximal part of the transverse process, which is also observed in
the C7. Also, the increased inclination (from ~20° to ~35°) of the transverse processes

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along the first third of the vertebral caudal region can be added as an autapomorphy of

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this taxon. These features have not been reported for any other abelisarids, including the

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ones recovered in the Cretaceous units of Brazil.
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The phylogenetic analysis does not permit a more accurate result for Kurupi but
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support its inclusion within Abelisauridae, based on the aforementioned traits. Only
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~5% of the used characters were scored for Kuripi, highlighting the incompleteness of
the holotype specimen and the IterPCR protocol recognized it as a rouge taxon. The
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discoveries of new abelisaurids in the last decades have shown the great diversification
of this clade by the Late Cretaceous, but several taxa are still poorly known and more
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specimens will be mandatory to improve their phylogenetic relationships.

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6.2. Musculature of the pelvic and anterior caudal regions of Kurupi

The most derived abelisaurids are characterized by the morphology of the

proximal and middle caudal vertebrae, with strongly dorsolaterally inclined transverse
processes, as can be seen in the South American Carnotaurus, Aucasaurus,
Skorpiovenator, Pycnonemosaurus, and Kurupi. Méndez (2014) proposed that several
of the morphological features observed in derived abelisaurids would be related to a
considerable increase in the stiffness of the proximal and middle portion of the tail, a
condition supported with other axial regions of the body. Persons and Currie (2011)
pointed out that besides rigidity, these vertebral traits reveal a progressive increasing of
the caudofemoral mass in South American species. Accordingly, Méndez (2014)
suggested that this trait would have been acquired in isolation of other non-South
American forms, considering that the inclination of the transverse processes is reduced
to absent in Arcovenator, Majungasaurus and Rahiolisaurus (Coria and Salgado, 1998;
O’Connor, 2007; Novas et al., 2010).

According to the model of Persons and Currie (2011), the dorsal elevation of the
transverse processes in South American abelisaurids would result in a smaller area of
the M. longissimus and M. spinalis (epaxial muscles evolved in the motility and support
of the tail). The bone structural framework (mainly the fan distal end of the transverse
process) giving rigidity would support the hypertrophied M. caudofemoralis actions
involved in locomotion, attributing to these species gains in cursoriality and speed, an
advantage to escape from larger predators such as hunting for quick prey; corroborating

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the proposal by Bonaparte et al. (1990) for Carnotaurus sastrei. In Kurupi, the
transverse processes in C1 have two triangular processes, namely anterolaterally

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projected triangular process (altp), that may contact the last sacral vertebra, and the
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anterolateral projected shelf-like processes, which define a notch where may articulate
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the ilium. As a consequence, these structures could provide rigidity to the tail, but
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unlike the condition in Carnotaurus sastrei, the more horizontal position of the
transverse processes suggests more developed epaxial muscles in Kurupi. The
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inclination of the transverse process in C7 of Kurupi was not as pronounced as in

Carnotaurus or Aucasaurus, however it exceeded 30º, similar to Skorpiovenator and
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Pycnonemosaursus, and represents a mass gain of hypaxial muscles. In Kurupi, there is

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a variation (incremental) of approximately 15º in the inclination of the transverse

process between the C1 and C7, a peculiar feature of the species. In other abelisaurids
(e.g., Aucasaurus, Carnotaurus, and Majungasaurus) the inclination of the transverse
process is nearly equal among the first caudal vertebrae. The ischium symphyses are
marked by a long continuous crest with a plug. The crest as well as a large part of the
groove is marked by a well-marked muscle insertion scar (Fig. 13). Méndez et al.,
(2014) associated such regions to attachment of the M. pubosichiotibialis, a muscle
involved in adducting the leg (Hutchinson & Gatesy, 2000). The broad areas of muscle
insertion in the ischium and the mass gain of hypaxial muscles in Kurupi corroborate
previous studies that indicate the development of the musculature associated with
cursoriality among the abelisaurids (e.g., Bonaparte et al., 1990; Persons and Currie,
2011; Méndez et al., 2014).

6.3. Size and ecology of Kurupi

 Grillo and Delcourt (2017) estimated the size of abelisaurids, indicating that the
holotype specimen of Pycnonemosaurus would represent the largest known abelisaurids
with about 9 meters-long while Carnotaurus could reaches 8 meters. Méndez et al.
(2014) mentioned that the ischia of Kurupi is 37.5% smaller than that of Carnotaurus
sastrei; if this proportion were maintained for the body length, Kurupi would reach
about 5 meters-long. The length of the centrum of the C1 and C7 are 89 and 85 mm
long, respectively, similar to those indicated for Majungasaurus, for which an
approximate size of 5 meters in length was proposed (Grillo and Delcourt, 2017). In
relation to the South American species, Kurupi would have a body size close to that
estimated for Quilmesaurus (5.3 m) or Llukalkan (~5 m), and intermediated between

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Niebla (4-4.5 m) and Skorpiovenator, Aucasaurus, Viavenator and Tralkasaurus (≥6 m)

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(e.g., Grillo and Delcourt, 2017; Cerroni et al., 2020; Aranciaga et al., 2021; Gianechini

r
et al., 2021).
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Kurupi was a medium-sized abelisaurid with a muscular caudal apparatus that
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suggested cursoriality. The relationship between femur/tibia and fibula in abelisaurids
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calls into question the high speed of the abelisaurids (Persons and Currie, 2011),
however the development of the muscles involved with walking is conspicuous. It is
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possible to infer that in Kurupi such muscles were robust, and could give to the species
the ability to cover a large area in search of prey and water in the hostile scenario of the
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Maastrichtian in the Marília Formation (Fig. 15). Kurupi lived in an arid environment
with little rainfall (annual precipitation around 300 to 200 mm; Dal’Bó et al., 2009);
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however, the presence of remains of bivalve mollusks, fishes and peirosaurid

crocodilyforms in the Maastrichtian paleofauna of the region reinforce the lacustrine-
fluvial deposits with bodies of water that would concentrate the fauna (Fig. 15).

6.4. Refining depositional aspects from the discovery of Kurupi

The deposits of Marília Formation were the result of progradation of alluvial

fans at the margins north and east of the basin resulting of tectonic rejuvenation in its
margins N-NE (Fernandes and Basilici, 2009). The unit is characterized by a vertical
succession of deposits (ephemeral fluvial and eolian) and paleosols. During the
sedimentation pause, the floodplain was occupied by vegetation and the soils developed
(Dal’Bó et al., 2009; Silva et al., 2015). Dal’bó et al. (2009) recognized aridisols
(including the geological profiles of Monte Alto region) with calcic horizon, based on
macro- and microfeatures and mineralogical and geochemical analyses. The presence of
aridisols suggests a semi-arid regime with scarce and periodical precipitation. Silva et
al. (2015) corroborated the evolution of calcic paleosols in semi-arid conditions and its
slowest evolution. Basilici et al. (2012) suggested a time for developed of aridisols of
about 175.000 years, a long time attributed to hydric scarcity. The taphonomic history
here interpreted for the holotype of K. itaata agrees with the idea of a semi-arid climate
(Fig. 15), because some of the taphonomic attributes (e.g. high degree of
permineralization) can be assigned to relatively humid periods (rainy seasons), while
other ones (e.g. presence of calcite on the cancellous bone) can be related to relatively

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drier moments (drought periods).

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7. Conclusions -p
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Kurupi itaata represents the first formally named vertebrate of the Marília
Formation (Bauru Group, Bauru Basin) and one of the few theropod records for the
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Maastrichtian of the Bauru Basin. Its abelisaurid affinities are well-established based on
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the anatomy of the pelvis and anterior caudal vertebrae; however, closer relationships
with other abelisaurids are still unclear. The holotype specimen indicates a somatically
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mature individual with an estimated size of around 5 meter long that bolsters the
presence of mid-sized abelisaurids and a wider spectrum of sizes among the clade, as
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has been shown by recent findings of abelisaurids (e.g., Canale et al., 2016; Grillo and
Delcourt, 2017; Filippi et al., 2018; Cerroni et al., 2020; Zaher et al., 2020; Aranciaga
Rolando et al., 2021; Gianechini et al., 2021). Features in the pelvis and caudal
vertebrae indicates muscle mass gain involved with cursoriality, as observed in other
South American abelisaurids, but with more plesiomorphic traits (such as least inclined
transverse processes) as in early diverging South American brachyrostrans. Kurupi
itaata enlarges the diversity of South American abelisaurids and highlights that the
meager theropod fossil record of the Late Cretaceous of Brazil would be biases and
systematic field works would improve it.

Acknowledgements
 We thank all the people involved in field activities, especially to Antonio C.
Arruda-Campos (in memoriam), Cledinei A. Francisco and José A. Bugarelli, to the
artist Júlia D’Oliveira for the in vivo reconstruction of Kurupi itaata, and to Mattia A.
Baiano for sharing bibliographic sources. The reviewers Federico Agnolín and Ariel
Méndez are thanked for the comments and suggestions that notably improved the
manuscript.

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Brazil), and a revision of the Sphagesauridae. Historical Biology, 20, 101–136.
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Apesteguía, S., Smith, N. D., Juárez Valieri, R., Makovicky, P. J., 2016. An unusual
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Paulo, Brazil. Cretaceous Research, 50, 59–71.
Tortosa, T., Buffetaut, E., Vialle, N., Dutour, Y., Turini, E., Cheylan, G., 2014. A new
abelisaurid dinosaur from the Late Cretaceous of Southern France:
Palaeobiogeographical implications. Annales de Paleontologie, 100(1), 63–86.
doi:10.1016/j.annpal.2013.10.003.

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new abelisaurid (Dinosauria, Theropoda) from the Lameta Formation

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(Cretaceous, Maastrichtian) of India. Contributions from the Museum of

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Paleontology, University of Michigan, 31(1), 1–42.

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Zaher, H., Pol, D., Navarro, B.A., Delcourt, R., Carvalho, A.B., 2020. An Early
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Cretaceous theropod dinosaur from Brazil sheds light on the cranial evolution of
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Figure captions
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Figure 1. Map of the Bauru Basin in the São Paulo State, Brazil (A) and the place
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where the holotype was found. The photographs show details of the specimen in situ,
discovered in 2002 (B) and prospective excavations in 2009 (C) and 2014 (D).

Figure 2. Kurupi itaata gen. et sp. nov., holotype MPMA 27-0001/02. Desiccation
marks on the surfaces of the ischium.

Figure 3. Kurupi itaata gen. et sp. nov., holotype MPMA 27-0001/02. Taphonomic
signals in the seventh caudal vertebra, in lateroventral view. Black arrows indicate the
calcite sheets and grey arrows indicate diagenetic breaks.
Figure 4. Kurupi itaata gen. et sp. nov., holotype MPMA 27-0001/02. First caudal
vertebra (C1) in and anterior (A), caudal (B) and dorsal (C) views, and schematic
drawing of the restored vertebra in dorsal view (D). Abbreviations: acf, anterior
centrodiapophyseal fossa (=prezygapophyseal centrodiapophyseal fossa,
=infraprezygaphophyseal fossa); altp: anterolaterally projected triangular process; cen,
centrum; cptv, cuneiforme projection of transverse process; ha, hypantrum; ho,
hyposphene; nc, neural canal; ns, neural spine; pcf, posterior centrodiapophyseal fossa
(=infrapostzygapophyseal fossa); posf, postspinal fossa; poz, postzygapophysis; prsf,
prespinal fossa; prz, prezygapophysis; stp, shelf-like process of transverse process; tvp,
transverse process.

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Figure 5. Kurupi itaata gen. et sp. nov., holotype MPMA 27-0001/02. First (A) and

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seventh (B) caudal vertebrae in ventral view, with details of the median groove and
pedicles for the chevron in C7. Abbreviation: cf, chevron facet.
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Figure 6. Kurupi itaata gen. et sp. nov., holotype MPMA 27-0001/02. Comparison of
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pneumaticity between the C1 (A) and C7 (B) caudal vertebrae, in right lateroventral
view. Abbreviations: acf, anterior centrodiapophyseal fossa (=prezygapophyseal
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centrodiapophyseal fossa, =infraprezygaphophyseal fossa); mcf, medial

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centrodiapophyseal fossa (=infradiapophyseal fossa); pcf, posterior centrodiapophyseal

fossa (=infrapostzygapophyseal fossa). Images not to scale.

Figure 7. Kurupi itaata gen. et sp. nov., holotype MPMA 27-0001/02. Detail of the
first caudal vertebra (C1) in anterior view. Abbreviations: acf, anterior
centrodiapophyseal fossa (=prezygapophyseal centrodiapophyseal fossa,
=infraprezygaphophyseal fossa); altp: anterolaterally projected triangular process; stp,
shelf-like process of transverse process.

Figure 8. Kurupi itaata gen. et sp. nov., holotype MPMA 27-0001/02. Fifth caudal
vertebra (C5), posterior half in lateral (A), posterior (B) and ventral (C) views.
Abbreviations: cf, chevron facet; ho, hyposphene; mcf, medial centrodiapophyseal
fossa (=infradiapophyseal fossa); mg, median groove; nc, neural canal; ns, neural spine;
pcr, posterior centrodiapophyseal fossa (=infrapostzygapophyseal fossa); posf,
postspinal fossa; poz, postzygapophysis; tvp, transverse process.

Figure 9. Kurupi itaata gen. et sp. nov., holotype MPMA 27-0001/02. Seventh caudal
vertebra (C7) in anterior (A), caudal (B), lateral (C) and dorsal (D) views.
Abbreviations: acf, anterior centrodiapophyseal fossa (=prezygapophyseal
centrodiapophyseal fossa, =infraprezygaphophyseal fossa); aptp, anterior projection of
transverse process; cen, centrum; ha, hypantrum; ho, hyposphene; mcf, medial
centrodiapophyseal fossa (=infradiapophyseal fossa); nc, neural canal; ns, neural spine;

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pcf, posterior centrodiapophyseal fossa (=infrapostzygapophyseal fossa); posf,
postspinal fossa; poz, postzygapophysis; prsf, prespinal fossa; prz, prezygapophysis;

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tvp, transverse process. -p
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Figure 10. Kurupi itaata gen. et sp. nov., holotype MPMA 27-0001/02. Details of the
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caudal vertebra (C7) in dorsal view. Squematic draw (A) and fossil (B). Abbreviations:
cptv, cuneiforme projection of transverse process; ns, neural spine; posf, postspinal
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fossa; poz, postzygapophysis; prsf, prespinal fossa; prz, prezygapophysis; tvp,

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transverse process.
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Figure 11. Kurupi itaata gen. et sp. nov., holotype MPMA 27-0001/02. Details of the
caudal vertebra (C7). Fan-shaped distal projection of the right transverse process in
lateral (A) and lateroventral (B) views.

Figure 12. Kurupi itaata gen. et sp. nov., holotype MPMA 27-0001/02. Pelvic girdle
(fused ischia and a portion of right acetabular region, including part of ilium and pubis)
in anterior (A), lateral (B, right; C, left) and posterior (D) views. Abbreviations: ac,
acetabulum; bf, brevis fossa; gr, groove; ib; ischial boot; il, ilium; ip, ischial peduncle;
it, ischidiac trochanter; k, kell; op, obturador process; pp, pubic peduncle; sac,
supraacetabular crest.
Figure 13. Kurupi itaata gen. et sp. nov., holotype MPMA 27-0001/02. Pelvic girdle in
anterior views with detail of the scars for muscle insertions on the keel and groove of
the ischia.

Figure 14. Strict consensus tree (A) and majority rule tree (50%) (B) of 5,724 most
parsimonious trees (see text for details of the analysis) showing the position of Kurupi
itaata among abelisaurid theropods. The numbers in (A) are Bremer support values (left
side) and absolute frequencies of bootstrap resampling analysis (right side).

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Figure 15. Life reconstruction of the abelisaurid Kurupi itaata by Júlia D’Oliveira.

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 1

Table 1. List of main vertebrate taxa recovered in the Serra da Galga and Marília formations.
Abbreviations: GO, Goiás State; MG, Minas Gerais State; SP, São Paulo State. Fossil record of the
Serra da Galga Formation modified from Martinelli and Teixeira (2015) plus the record of Brum et
al. (2021).

Serra da Galga Formation (MG) Marília Formation (SP, MG, GO)

OSTEICHTHYES TESTUDINATA
Sarcopterygii Podocnemidoidae indet. (Rio Verde, GO)
Ceratodontidae indet.
Ceratodus sp. CROCODYLIFORMES

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Actinopterygii

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Sphagesauridae indet. (Marília, SP)
Actinopterygii indet. Peirosauridae indet. (Monte Alto, SP)
Amiidae indet. Notosuchia indet. (Rio Verde, GO)

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Lepisosteidae indet. Crocodyliformes indet. (Quirinópolis, GO)
Teleostei indet.
Siluriformes indet.
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Characiformes indet. Titanosauria indet. (Campina Verde, MG)
Perciformes indet. Titanosauria indet. (Rio Verde, GO)
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Titanosauria indet. (Marília, SP)

ANURA Titanosauria indet. (Monte Alto, SP)
Abelisauridae indet. (Marília, SP)
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Baurubatrachus pricei Báez and Perí, 1989

Uberabatrachus carvalhoi Báez et al., 2012 Kurupi itaata gen. et sp. nov. (Monte Alto, SP)
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TESTUDINATA
Cambaramys langertoni França and Langer,
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2005
Peiropemis mezzalirai Gaffney et al., 2011
Pricemys caiera Gaffney et al., 2011
Podocnemididae indet.

LEPIDOSAURIA
Pristiguana brasiliensis Estes and Price, 1973
CROCODYLIFORMES
Itasuchus jesuinoi Price, 1955
Labidiosuchus amicum Kellner et al., 2011
Peirosaurus torminni Price, 1955
Uberabasuchus terrificus Carvalho et al., 2004

DINOSAURIA
Trigonosaurus pricei Campos et al., 2005
Baurutitan brittoi Kellner et al., 2005
Uberabatitan ribeiroi Salgado and Carvalho,
2008
Ypupiara lopai Brum et al., 2021
 2

Titanosauria indet.
Aeolosaurini indet.
Abelisauroidea indet.
Abelisauridae indet.
Maniraptora indet.
Avialae indet.
Enantiornithes indet.

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Highlights
- A new abelisaurid theropod (Dinosauria) is described for the Cretaceous of southeastern
Brazil.

- It represents the first named tetrapod for the Late Cretaceous Marília Formation (Bauru
Group).

- The new taxon represents a mid-sized abelisaurid (~5 meters long).

- The new species increase the abelisauroid diversity for western Gondwana.

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Declaration of interests

☒ The authors declare that they have no known competing financial interests or personal relationships
that could have appeared to influence the work reported in this paper.

☐The authors declare the following financial interests/personal relationships which may be considered
as potential competing interests:

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