Available via license: CC BY 3.0
Content may be subject to copyright.
Teeth of Abelisauridae and Carcharodontosauridae cf.
(Theropoda, Dinosauria) from the Campanian-
Maastrichtian Presidente Prudente Formation
(Southwestern São Paulo State, Brazil)
Dientes de Abelisauridae y Carcharodontosauridae cf. (Theropoda,
Dinosauria) del Campaniano-Maastrichtiano Formación Presidente
Prudente (Suroeste Provincia de São Paulo, Brasil)
M.R. Furtado¹, C.R. Candeiro², L.P. Bergqvist¹
ABSTRACT
Isolated theropod teeth referable to Abelisauridae indet., and Carcharodontosauridae cf., from the
Campanian-Maastrichtian Presidente Prudente Formation of the western São Paulo State, Brazil, are des-
cribed. They are compared to the Late Cretaceous Gondwanan theropod dinosaur teeth and their affini-
ties are discussed. These teeth are significant because carnivorous dinosaur remains are poorly known
from the Late Cretaceous beds of Western São Paulo State except for a few isolated elements.
Keywords: Dinosaur, theropod, Brazil, Upper Cretaceous, theropod, Bauru Group.
RESUMEN
Se describen dientes aislados de terópodos referidos a Abelisauridae indet. y Carcharodontosauridae
cf., procedentes de la Formación Presidente Prudente del Campaniano-Maastrichtiano, en el oeste del
estado de San Pablo, Brasil. Los materiales son comparados con dientes de dinosaurios gondwánicos
del Cretácico tardío, y cuyas afinidades son aquí discutidas. Estos dientes de dinosaurios carnívoros son
significativos debido a que su presencia es pobremente conocida en el Cretácico tardío del oeste del
estado del San Pablo, excepto por unos pocos huesos.
Palabras clave: Dinosaurio, teropode, Brasil, Cretacico superior, Grupo Bauru Group.
¹ Programa de Pós-Graduação em Geologia, Departamento de Geologia, Universidade Federal do Rio de Janeiro. Avenida Athos da
Silveira Ramos, 274, Prédio CCMN, UFRJ. Ilha do Fundão, Rio de Janeiro/RJ, Brasil. 21941-916. Email: migueltupa@gmail.com;
bergqvist@geologia.ufrj.br
² Laboratório de Geologia, Universidade Federal de Uberlândia, Campus Pontal, Curso de Geografia. Avenida José João Dib, 2545,
Progresso, Ituiutaba/MG, Brasil. 38300-132. Email: candeiro@pontal.ufu.br
Estudios Geológicos, 69(1)
enero-junio 2013, 105-114
ISSN: 0367-0449
doi:10.3989/egeol.40829.170
Introduction
Theropod teeth are well-documented elements in
the fossil record of dinosaurs. Teeth are the most
common fossils recovered in any vertebrate site due
to their structure - an internal soft pulp inside a denti-
ne which is covered by an outer layer of hard
enamel. This layer is more resistant to intemperism
than the outer layers of bones, the periosteum, allo-
wing a better preservation on the fossil record com-
pared to bones.
Cretaceous theropod record in Brazil is scarce,
being composed of five known species: Irritator
challengeri Martill, Cruickshank, Frey, Small &
Clarke, 1996, Santanaraptor placidus Kellner,
1999; Mirischia asymmetrica Naish & Frey, 2004;
Furtado.qxd:Estudios 24/06/13 12:19 Página 105
Pycnonemosaurus nevesi Kellner & Campos,
2002 and Oxalaia quilombensis Kellner, Azevedo,
Machado, Carvalho & Henriques, 2011.
In order to better understand the species rich-
ness of theropods, the Laboratório de Macrofós-
seis of the Departamento de Geologia at Universi-
dade Federal do Rio de Janeiro, Rio de Janeiro
state, made eight field trips to Alfredo Marcondes
and Flórida Paulista municipalities from 2007 to
2011 and collected remarkable theropod mate-
rials. Among them, carcharodontosaurid and abe-
lisaurid teeth are the most abundant findings.
In this paper, isolated theropod teeth from two
localities are described and compared with other
dinosaur teeth. In the case of isolated theropod
teeth, their systematic value has been demonstra-
ted by Currie et al. (1990), Fiorillo & Currie
(1994), Rauhut (2002), Sankey et al. (2002), Can-
deiro (2007), Fanti & Therrien (2007), and others.
These authors documented a diverse range of the-
ropod taxa in faunas across continents and time
periods using a tentative taxonomic identification
at subfamily (Martill & Hutt, 1996), family (Rau-
hut, 2002; Medeiros, 2006) and superfamily
levels (Candeiro, 2007). In exceptional cases, the
teeth can be identified to even genera or species
(Currie et al., 1990; Sankey et al., 2002).
Previous studies of teeth recovered from the
two mentioned localities of the Bauru Group
revealed the presence of Abelisauroidea, Abeli-
sauridae, Carcharodontosauridae, and Theropoda
indet. (Candeiro et al., 2006 and Bittencourt &
Langer, 2011 and references herein). Other loca-
lities of the same group have yielded a variety of
theropod teeth, showing an unexpected, wides-
pread Gondwanan theropod distribution and
diversity (Candeiro et al., 2004, 2006). The Fló-
rida Paulista and Alfredo Marcondes theropod
record will be accurately described providing
new information on the faunal composition of
theropods in the Late Cretaceous of central
Brazil.
106 M.R. Furtado, C.R. Candeiro, L.P. Bergqvist
Estudios Geológicos, 69(1), 105-114, enero-junio 2013. ISSN: 0367-0449. doi:10.3989/egeol.40829.170
Fig. 1—Bauru Group geological map (abbreviations: AM1, Alfredo Marcondes; FP1, Florida Paulista).
Furtado.qxd:Estudios 24/06/13 12:19 Página 106
Material and methods
Eleven isolated teeth from Alfredo Marcondes
and Flórida Paulista municipalities, both in São
Paulo State, have been studied. Although bone
fragments are quite common in Alfredo Marcondes
(São Paulo State, Brazil), most of them are too
fragmentary for identification down to generic or
specific levels. All specimens described here are
housed in the vertebrate collections of the Departa-
mento de Geologia, Universidade Federal do Rio
de Janeiro, at reptile teeth (Rd) collection (UFRJ-
DG-Rd).
For description of the teeth, the following metrics
were used: FABL (Fore-Aft Basal Length; measu-
red at the level of the basal end of the posterior cari-
na, perpendicular to the longitudinal axis of the
tooth), HIR (Height Relative Index; height of crown
divided by FABL), BW (Basal Width; measured at
the same level as the FABL, and perpendicular
FABL in the horizontal plane), TCH (Tooth Crown
Height; vertical distance from the outer rim of the
tooth socket to the tip, perpendicular to both FABL
and BW; see Farlow et al., 1991), and DSDI (Den-
ticle Size Difference Index; ratio between number
of denticles over a given length on the medial cari-
na, divided by the number of denticles over the
same length on the distal carina; Rauhut & Werner,
1995). For calculating the DSDI, only complete
tooth crowns have been considered to reduce the
Estudios Geológicos, 69(1), 105-114, enero-junio 2013. ISSN: 0367-0449. doi:10.3989/egeol.40829.170
Teeth of Abelisauridae and Carcharodontosauridae cf. from SW Sao Paulo State, Brazil 107
Fig. 2—Abelisauridae teeth. UFRJ-DG-378-Rd – A, labial view; B, posterior carina; C, square-shaped posterior denticles; D, cross-sec-
tion. UFRJ-DG-499-Rd – E, labial view; F, posterior carina; G, posterior square-shaped denticles; H, cross-section. UFRJ-DG-566-Rd –
I, labial view; J, posterior carina; K, cross-section. UFRJ-DG-572-Rd – L, labial view; M, posterior carina; N, cross-section.
Furtado.qxd:Estudios 24/06/13 12:19 Página 107
108 M.R. Furtado, C.R. Candeiro, L.P. Bergqvist
Estudios Geológicos, 69(1), 105-114, enero-junio 2013. ISSN: 0367-0449. doi:10.3989/egeol.40829.170
FABL BW TCH
DSDI CROSS
SECTION
SHAPE
ENAMEL
WRINKLE IAR SHAPE BLOOD
GROOVE PROVENANCE
ANT POST
Abelisauridae indet.
UFRJ DG 378 Rd 8.72 mm 4.07 mm 14.48 mm 32Strongly compressed NO 1.66 X UNMARKED Alfredo Marcondes municipallity
UFRJ DG 499 Rd 7.63 mm 3.92 mm 12.80 mm 33Strongly compressed NO 1.68 SUB-RECTANGULAAR UNMARKED Alfredo Marcondes municipallity
UFRJ DG 559 Rd X 5.89 mm 13.90 mm 2 2 Strongly compressed NO ?? ? Alfredo Marcondes municipallity
UFRJ DG 566 Rd 9.33 mm 4.27 mm 14.21 mm 3 X Strongly compressed NO 1.52 X X Alfredo Marcondes municipallity
Carcharodontosauridae indet.
UFRJ DG 371 Rd 9.72 mm 4.60 mm 15.14 mm 2 2 compressed NO 1.75 RECTANGULAR UNMARKED Florida Paulista municipallity
UFRJ DG 374 Rd 5.72 mm 3.42 mm 8.12 mm X X compressed NO ? RECTANGULAR PRESENT Florida Paulista municipallity
UFRJ DG 557 Rd 9.23 mm 4.88 mm 18.36 mm 32compressed NO 1.99 RECTANGULAR PRESENT Florida Paulista municipallity
UFRJ DG 560 Rd 6.49 mm 3.11 mm 11.97 mm 3 3 compressed NO 1.84 RECTANGULAR PRESENT Alfredo Marcondes municipallity
UFRJ DG 561 Rd 25.86 mm 15.75 mm 45.58 mm 2 2 compressed YES 1.76 RECTANGULAR `PRESENT Alfredo Marcondes municipallity
UFRJ DG 570 Rd 13.92 mm 7.73 mm 16.80 mm X2compressed YES ? RECTANGULAR PRESENT Alfredo Marcondes municipallity
UFRJ DG 593 Rd 9.10 mm 4.80 mm 17.04 mm 33compressed YES 1.87 RECTANGULAR PRESENT Florida Paulista municipallity
Table 1.—Theropod teeth from upper Cretaceous Bauru Group, São Paulo State, Brazil. Abbreviations: FABL (Fore-Aft Basal Length), BW (Basal Width), TCH (Tooth
Crown Height), DSDI (Denticle Size Difference Index), IAR (“Indice de Altura Relativa” = Height Relative Index). All measures in milimeters.
Furtado.qxd:Estudios 24/06/13 12:19 Página 108
risk of misidentification. Mean DSDI values should
be considered with caution because sample size is
rather small. Cross-sections of the specimens were
taken by placing a copper thread around the tooth
crown, near the base of the tooth. The shape resul-
ted was then traced onto a sheet of paper (see Can-
deiro et al., 2004).
Geological Setting
The Bauru Basin, southwestern São Paulo state, is
a tectonic entity developed at the south-Centre of the
South American Platform. It encloses a sedimentary
structure succession with associated volcanic rocks
covering the basaltic Neocomian flows from the
Serra Geral Formation. Its form is approximately
elliptical, with its longer axis directed towards nor-
theast, and it has approximately 370.000 Km² of
exposure and maximum preserved sediment layer of
approximately 300m thickness (Fernandes, 1996).
The basin limits, of essentially tectonic nature, are
drawn by the Rondonópolis anteclise at northwest,
the Alto Parnaíba emergement at Northeast, align-
ments of Moji-Guaçu, São Carlos-Leme e Ibitinga-
Botucatu Rivers at east, Paranapanema, at southeast,
and by the Piquiri River at south. The development
of the Bauru Basin occurred continuously between
the Santonian and the Maastrichtian (Dias-Brito et
al., 2001). Its sediment is essentially sandy in nature,
constituting a unique sequence, deposited under
semi-arid conditions at the borders to desert condi-
tions in the inner basin. At the Maastrichtian, the
increased tectonic activity resulted in an increasing
aportment of rudaceous sediments, mainly associated
with the alignments of the Paranapanema and Ibitin-
ga-Botucatu rivers. In this area alkaline volcanism
occurred along the alignment of the Moji-Guaçu
river tectonic morfisms configuring its recent east
border (Riccominni, 1997). At the Bauru Basin out-
crops, the Presidente Prudente Formation is late
Campanian-early Maastrichtian in age, dated through
lithostratigraphic and biostratigraphic correlation
based on the vertebrae of aelosaurine sauropods
(Simbras, 2009). The Presidente Prudente Formation
differs in its geological context from the Adamantina
Formation for the presence of thick pelitic flood
plain deposits, which can reach 11 m of thickness.
The association of the lithofaciological and architec-
tural elements suggest that the depositional environ-
ment of the Presidente Prudente Formation at the
region was fluvial meandering of high sinuosity, with
an extensive flood plain (model 7 from Miall, 1985).
This interpretation differs from the environment of
the Adamantina Formation, which is sandy fluvial
meandering (Batezelli et al., 2007) or fluvial or brai-
ded fluvial (Fernandes & Coimbra, 1996a, b). The
Adamantina Formation resembles more to the inter-
pretation of the Presidente Prudente Formation:
Sandy fluvial meandering (Fernandes & Coimbra,
2000; Simbras, 2009).
Systematic Paleontology
Order Saurischia Seeley, 1888
Suborder Theropoda Marsh, 1881
Superfamily Abelisauroidea Bonaparte, 1991
Abelisauridade indet.
(Fig. 2)
Referred specimens. Alfredo Marcondes municipality: UFRJ-
DG 378Rd, UFRJ-DG 499Rd; Flórida Paulista municipality:
UFRJ-DG 372Rd, UFRJ-DG 566Rd.
Description. The specimens are strongly compressed labiolin-
gually and curved; TCH equals 12.80 to 14.48 mm in height and
HIR varies from 1.52 to 1.68. These values fall in a typical range
of low crowned teeth. In basal cross-sections (Fig. 2D, H, K, N),
the teeth are strongly labiolingually compressed. FABL varies
from 7.63 to 9.33, BW from 3.92 to 4.27. Both carinae are com-
pletely denticulate. The denticles of the anterior carina are small-
er than the ones of the posterior carina, and DSDI varies between
3 and 2 denticles. The medial denticles are large, rounded and
pointed to the apex of the tooth, whereas medial denticles are
smaller and with apex not pronounced pointed. No marked blood
grooves are present at the base of the denticles. No wrinkles are
present on labial or lingual faces (Fig. 2A, E, I, L). Posterior den-
ticles are subsquared and uniform in size along tooth.
Discussion. These teeth are readily identified as abelisaurid
on the basis of their low crowns, square-shaped denticles (Fig.
2C, G) and labiolingually flattened shape in cross section. In
particular, the low crowned teeth from Alfredo Marcondes
(UFRJ DG 378 Rd, UFRJ DG 499 Rd, UFRJ DG 559 Rd,
UFRJ DG 566 Rd), clearly indicate that it represents an
abelisaurid. Bonaparte (1996), Lamanna et al. (2002), Wilson
& Upchurch (2003), and Candeiro et al. (2004) noted a low
crowned teeth in the abelisaurids Aucasaurus garridoi Coria,
Chiappe & Dingus, 2002, Carnotaurus sastrei Bonaparte, 1985
and Rajasarus narmadensis Wilson, Sereno, Srivastava, Bhatt,
Khosla & Sahni, 2003 and regarded it as a diagnostic character
of the family. Candeiro (2007) pointed out that abelisaurid
teeth are characterized by labial and lingual faces that are more
convex in cross-section near the anterior edge and flatter poste-
riorly; the lingual face is also slightly concave mediodistally.
The denticle shape is subsquare. Other characteristic of
abelisaurid teeth include distal margin straight tooth crowns
(Smith & Lamanna, 2006; Smith & Dalla Vecchia, 2006;
Estudios Geológicos, 69(1), 105-114, enero-junio 2013. ISSN: 0367-0449. doi:10.3989/egeol.40829.170
Teeth of Abelisauridae and Carcharodontosauridae cf. from SW Sao Paulo State, Brazil 109
Furtado.qxd:Estudios 24/06/13 12:19 Página 109
Krause et al., 2009; Gianechini et al., 2010). The cross-sections
of the Alfredo Marcondes teeth have these characteristics.
Among theropod dinosaurs, a mean DSDI of 2.0-3.0, as in the
specimens from Alfredo Marcondes, is typical for abelisaurids,
and is only found in Gondwanan theropods of South America
(Abelisauroidea; see Candeiro, 2007). Although within basal
Abelisauroidea (Noasauridae) teeth are poorly known, and
although the original description is far from accurate, the teeth of
Noasaurus neali Bonaparte & Powell, 1980 differ from the spec-
imens from Alfredo Marcondes in their overall shape and rela-
tive size of the denticles (Fanti & Therrien, 2007). The teeth of
Abelisauridae (e.g., Aucasaurus, Carnotaurus) are less markedly
curved than those seen in Noasaurus leali Bonaparte & Powell,
1980 (Candeiro, 2007). Because the teeth from Alfredo Marcon-
des show great similarities to teeth of Abelisauridae (Abelisaurus
comahuensis Bonaparte & Novas, 1985, Aucasaurus, Carnotau-
rus) in their shape, crown height and DSDI (Bonaparte & Novas,
1985; Candeiro et al., 2004; Candeiro, 2007), they are referred to
as Abelisauridae indet.
Order Saurischia Seeley, 1888
Suborder Theropoda Marsh, 1881
Carcharodontosauridae cf.
(Fig. 3)
Referred Specimens. Alfredo Marcondes municipality:
UFRJ-DG 560Rd, UFRJ-DG 561Rd, UFRJ-DG 570Rd, UFRJ-
DG 593Rd. Flórida Paulista municipality: UFRJ-DG 371Rd,
UFRJ-DG 374Rd, UFRJ-DG 557Rd (Tab. 1).
Description. The teeth are medium to large (8.12 to 45.58 mm
in height = TCH) in size and strongly compressed labiolingually
and curved apically. Their HIR varies from 1.75 to 1.99, which
indicates relatively higher crowns. In basal cross sections, the teeth
are compressed labiolingually. FABL ranges between 5.72 and
25.86 mm, and BW 3.11 and 15.75. Both anterior and posterior
carinae are completely denticulate. The denticles are smaller in the
anterior carina than in the posterior carina. DSDI varies from 2 to 3
denticles on both carinae. These teeth develop blood grooves at the
base of the denticles. Strong or smooth wrinkles are present on the
labial or lingual face, slight wrinkles on UFRJ-DG 374-Rd, 557Rd,
560Rd, UFRJ-DG 561Rd, UFRJ-DG 593Rd (Tab. 1). Posterior
denticles are rectangular in shape, uniformly sized along the tooth.
Discussion. These teeth are very similar to teeth from the
Upper Cretaceous carcharodontosaurids Carcharodontosaurus
Stromer, 1931, Giganotosaurus Coria & Salgado, 1995, and
Mapusaurus Coria & Currie, 2002 (Candeiro, 2007). The tall
crowns of the Alfredo Marcondes tooth suggest that they repre-
sent carcharodontosaurids. Candeiro (2007) noted the presence
of a tall crown (HIR ≥ 2.2) in the teeth of the Carcharodon-
tosauridae (e.g. Giganotosaurus, Mapusaurus and Tyrannotitan
Novas, de Valais, Vickers-Rich & T. Rich, 2005) and regarded it
as a diagnostic character of the family. Candeiro (2007) also
pointed out that the South American carcharodontosaurid teeth
are characterized by labial and lingual faces more convex in
cross-section near the anterior edge and flatter posteriorly; the
lingual face is also slightly concave mediodistally. Carcharodon-
tosauridae denticles are rectangular in shape (Fig. 3C, I, M, R,
Y). Cross sections of the specimens are similar in shape to those
of the Carcharodontosauridae.
Typical Carcharodontosauridae enamel wrinkles (sensu Coria
& Currie, 2002; Candeiro et al., 2006) are visible on marginal
crown teeth, and these marginal wrinkles in the teeth from Alfre-
do Marcondes and Flórida Paulista are similar to those of Car-
charodontosaurus, Giganotosaurus, Mapusaurus and carcharo-
dontosaurids from Peirópolis Site (Minas Gerais State). Wrinkles
are present to a lesser degree in other theropods (e.g.,
Aucasaurus garridoi, Abelisauridae, Dromaeosauridae and
Tyrannosauridae; Candeiro et al., 2004), but their teeth are much
more distinct (robust or high crown tooth) than the specimen
studied herein (Benson et al., 2007).
Concluding Remarks
South American sites from Argentina and Brazil
have yielded theropod teeth that belong to abelisau-
roid (Abelisauroidea and Abelisauridae indet.) and
carcharodontosaurid morphotypes (Candeiro et al.,
2006, 2010; Bittencourt & Langer, 2011). These
records show the continuous and sustained compo-
sition of Gondwanan dinosaur fauna in the Bauru
Group during the Late Cretaceous, reinforced with
the Alfredo Marcondes and Flórida Paulista mate-
rials described in this paper. Based on the documen-
ted morphotypes, the fauna from São Paulo State is
strikingly similar to other Late Cretaceous theropod
fauna from South America (Chubut and Neuquen
groups; Leanza et al., 2004; Candeiro, 2007; Casal
et al., 2007; Candeiro & Rich, 2010; Salgado et al.,
2009) and Africa (Kem Kem Formation, Candeiro
& Rich, 2010). Abelisauroid and charcarodontosau-
rid teeth, as well as teeth assignable to African and
South American theropods Giganotosaurus, Car-
charodontosaurus and Abelisaurus and related taxa,
are often found in Turonian-Campanian localities
on those two continents. Although other groups of
theropods such as dromaeosaurids were present in
other Upper Cretaceous localities of South America,
they are absent in the Bauru Group (Lopes & Can-
deiro, 2010; Bittecourt & Langer, 2011). In this res-
pect, the theropod fauna from the post-Turonian of
South America (e.g., Abelisauroidea, Carcharodon-
tosauria), Africa (Abelisauroidea, Carcharodonto-
sauria), India-Madagascar (Abelisauroidea) and
Antarctica (Abelisauroidea) seems, on a broad taxo-
nomic level, to be more similar to Patagonian thero-
pod taxa (e.g., Huene & Matley, 1933; Bonaparte et
al., 1990; Sampson et al., 1998; Case et al., 2001,
2007; Wilson et al., 2003; Novas et al., 2004a,
2005a; Candeiro et al., 2004; Fanti & Therrien,
2007; Sereno & Brussate, 2008; Novas, 2009; Sal-
gado et al., 2009;). Except for the abelisaurid Pyc-
110 M.R. Furtado, C.R. Candeiro, L.P. Bergqvist
Estudios Geológicos, 69(1), 105-114, enero-junio 2013. ISSN: 0367-0449. doi:10.3989/egeol.40829.170
Furtado.qxd:Estudios 24/06/13 12:19 Página 110
nenomosaurus nevesi from the Turonian-Santonian
Adamantina Formation (sensu Fernandes & Coim-
bra, 1996), the premaxilla of an Abelisauridae indet.
(Bertini, 1996) and the occurrences in Western Sao
Paulo State and Triângulo Mineiro region (Candeiro
et al., 2006) - the last ones interpreted as indetermi-
nated Abelisauridae and Carcharodontosauridae the-
ropods (Bittencourt & Langer, 2011) -, Western Sao
Paulo State fossil localities are generally poor in
carnivorous dinosaurs skeletal remains. Moreover,
Estudios Geológicos, 69(1), 105-114, enero-junio 2013. ISSN: 0367-0449. doi:10.3989/egeol.40829.170
Teeth of Abelisauridae and Carcharodontosauridae cf. from SW Sao Paulo State, Brazil 111
Fig. 3—Carcharodontosauridae cf. teeth. UFRJ-DG-371-Rd – A, labial view; B, posterior carina; C, rectangular posterior
denticles; D, cross-section. UFRJ-373-Rd – E, labial view; F, cross-section. UFRJ-DG-557-Rd – G, labial view; H, pos-
terior carina; I, rectangular posterior denticles; J, cross-section. UFRJ-DG-560-Rd – K, labial view; L, posterior carina;
M, rectangular posteiror denticles; N, cross-section. UFRJ-DG-561-Rd – O, labial view; P, posterior Carina; Q, rectan-
gular posterior denticles; R, cross-section, S, lateral wrinkles. UFRJ-DG-570-Rd – T, labial view; U, posterior carina; V,
cross-section. UFRJ-DG-593-Rd – W, labial view; X, posterior Carina; Y, rectangular posterior denticles; Z, cross-sec-
tion; Z’, lateral wrinkles. Scale bar: 10mm (except to C, I, M, R, Y). Arrow show detailed enamel wrinkles.
Furtado.qxd:Estudios 24/06/13 12:19 Página 111
Cretaceous Patagonian dinosaurs have been repor-
ted from the largely coeval Chubut and Neuquen
groups (e.g., Leanza et al., 2004; Casal et al.,
2009; Agnolin et al., 2012; Ibiricu et al. 2012) and
from the basal part of the Presidente
Prudente/Adamantina Formation (Fernandes &
Coimbra, 2000; Simbras, 2009). These have been
attributed to unknown theropods, as the skeletal
remains are yet to be recovered. Hence, instead of
considering all isolated teeth from this beds as
Gondwanan theropods the recovery of P. nevesi
and Abelisauridae indet. premaxilla (Bertini, 1996;
Kellner & Campos, 2002) and our current finding
show that in the Late Cretaceous sites where dino-
saur remains are poorly known, isolated theropod
teeth deserve more attention (Candeiro et al.,
2006; Bittencourt and Langer, 2011).
ACKNOWLEDGEMENTS
The authors would like to thank the Laboratório de Metaliza-
ção do Centro de Tecnologia from UFRJ for the microphotogra-
phies of the teeth, CAPES and Geologia Department for Miguel
Rodrigues Furtado schollarship. The manuscript was substantial-
ly improved after the revision of Federico Agnolin (Museo
Argentino de Ciencias Naturales/Argentina) and José I. Canudo
(Universidad de Zaragoza/Spain). Tetsuto Miyashita (University
of Alberta/Canada) made the final revision of the article. We are
grateful to Aline Ghilardi (Universidade Federal do Rio de
Janeiro for preparing Figures 3S and 3Z’. R. Candeiro and L.
Bergqvist were supported by Conselho Nacional de Desenvolvi-
mento Científico e Tecnológico (CNPq)/Bolsista de Produtivi-
dade em Pesquisa fellow.
References
Batezelli, A.; Saad, A.R.; Etchebehere, M.L.C.; Perinotto,
J.A.J. & Fulfaro, V.J., (2003). Análise Estratigráfica
Aplicada à Formação Araçatuba (Grupo Bauru – KS)
no Centro-Oeste do Estado de São Paulo. Geociências,
22: 5-19.
Batezelli, A. (2003). Análise de Sedimentação Cretácea
no Triângulo Mineiro e sua Correlação com Áreas
Adjacentes Adjacentes. Curso de Pós-Graduação em
Geociências, Área de Concentração em Geologia Regio-
nal, Universidade Estadual Paulista, Tese de Doutora-
mento, 183 pp.
Batezelli, A. (2007). Arquitetura Deposicional e Evolução
da Seqüência Aluvial Neocretácea da Porção Setentrio-
nal da Bacia Bauru, no Sudeste Brasileiro. Revista Bra-
sileira de Geociências, 37(1): 163-181.
Bittencourt, J.S. & Langer, M.C. (2011). Brazilian dino-
saurs and their biogeographical implications. Anais da
Academia Brasileira de Ciências 83: 23-60.
doi:10.1590/S0001-37652011000100003
Bonaparte, J.F.; Novas, F.E. & Coria, R.A. (1990). Carno-
taurus sastrei Bonaparte, the horned, lightly built carno-
saur from the Middle Cretaceous of Patagonia. Contri-
butions in Science, Natural History Museum of Los
Angeles County 416: 1–41.
Canale, J.I.; Agnolín, F.L.; Ezcurra, M.D. & Novas, F.E.
(2007). Notas sobre el registro de dientes aislados de
dinosaurios terópodos cretácicos de América del Sur.
Ameghiniana, 44 (4): 8R.
Canale, J.I.; Scanferla, C.A.; Agnolín, F.L. & Novas, F.E.
(2009). New carnivorous dinosaur from the Late Creta-
ceous of NW Patagonia and the evolution of abelisaurid
theropods. Naturwissenschaften, 96: 409-414.
doi:10.1007/s00114-008-0487-4
Casal, G.; Candeiro, C.R.A.; Martínez, R.; Inavy, E. &
Ibiricu, L. (2009). Dientes de teropoda (Dinosauria:
Saurischia) de la Formación Bajo Barreal, Cretácico
Superior, Provincia del Chubut, Argentina. Geobios
(Lyon), 42: 553-560. doi:10.1016/j.geobios.2009.03.002
Case, J.A.; Martin, J.E. & Reguero, M. (2007). A dromae-
osaur from the Maastrichtian of James Ross Island and
the Late Cretaceous Antarctic dinosaur fauna. U.S. Geo-
logical Survey and the National Academies, Short
Research Paper, 83: 1–4.
Fanti, F. & Therrien, F. (2007). Theropod tooth assembla-
ges from the Late Cretaceous Maevarano Formation and
the possible presence of dromaeosaurids in Madagascar.
Acta Palaeontologica Polonica, 52: 155-166.
Farlow, J.O.; Brinkman, D.L.; Abler, W.L. & Currie, P.J.
(1991). Size, shape and serration density of theropod
dinosaur lateral teeth. Modern Geology, 16: 161-198.
Fernandes, L.A. & Coimbra, A.M. (1996a). A Bacia
Bauru (Cretáceo Superior, Brasil). Anais da Academia
Brasileira de Ciências, 68(2): 195-205.
Fernandes, L.A. & Coimbra, A.M., (1996b). Estratigrafia
y Ambientes Deposicionales de La Cuenca Bauru (Cre-
tácico Superior, Brasil). Acta Geologica Hispanica,
30(4): 11-30.
Fernandes, L.A. & Coimbra, A.M. (2000). Revisão Estrá-
tigráfica da Parte Oriental da Bacia Bauru (Neocretá-
ceo). Revista Brasileira de Geociencias, 30(4): 717-728.
Fiorillo, A.R. & Currie, P.J. (1984). "Theropod teeth from
the Judith River Formation (Upper Cretaceous) of south-
central Montana." Journal of Vertebrate Paleontology,
14: 74-80. doi:10.1080/02724634.1994.10011539
Gianechini, F.A.; Lio, G.L. & Apesteguía, S. (2010). Iso-
lated archosaurian teeth from “La Bonita” (Late Creta-
ceous, Santonian-Campanian). Río Negro province,
Argentina. Historia Natural, 1: 3-14.
Huene, F. & Matley, C.A. (1933). The Cretaceous Sauris-
chia and Ornithischia of the central provinces of India.
Memoirs of the Geological Survey of India, 21: 1–74.
Kellner, A.W.A. & Campos, D.A. (2002). On a theropod
dinosaur (Abelisauria) from the continental Cretaceous
of Brazil. Arquivos do Museu Nacional Rio de Janeiro,
60 (3): 163-170.
Kellner, A.W.A. (1999). Short note on a new dinosaur
(Theropoda, Coelurosauria) from the Santana Formation
(Romualdo Member, Albian), Northeastern Brazil. Bole-
tim do Museu Nacional (Rio de Janeiro, Brasil), 49: 8.
112 M.R. Furtado, C.R. Candeiro, L.P. Bergqvist
Estudios Geológicos, 69(1), 105-114, enero-junio 2013. ISSN: 0367-0449. doi:10.3989/egeol.40829.170
Furtado.qxd:Estudios 24/06/13 12:19 Página 112
Kellner, A.W.A.; Azevedo, S.A.K.; Machado, E. B.; Car-
valho, L.B. & Henriques, D.D.R. (2011). A new dino-
saur (Theropoda, Spinosauridae) from the Cretaceous
(Cenomanian) Alcântara Formation, Cajual Island, Bra-
zil. Anais da Academia Brasileira de Ciências, 83 (1):
99–108. doi:10.1590/S0001-37652011000100006
Lopes, L.N.F. & Candeiro, C.R.A. (2010). The Gondwa-
nan Unenlagiinae (Dinosauria, Theropoda) paleobiogeo-
graphy. Boletim de Resumos do VII Simpósio Brasileiro
de Paleontologia de Vertebrados (Rio de Janeiro), Bole-
tim de Resumos, 1:6.
Martill, D.M.; Cruickshank, A. R. I.; Frey, E.; Small, P. G.
& Clarke, M. (1996). "A new crested maniraptoran
dinosaur from the Santana Formation (Lower Cretace-
ous) of Brazil". Journal of the Geological Society, 153:
5. doi:10.1144/gsjgs.153.1.0005
Martill, D.M. & Hutt, S. (1996). Possible baryonychid
dinosaur teeth from the Wessex Formation (Lower Cre-
taceous, Barremian) of the Isle of Wight, England. Pro-
ceedings of the Geologists’ Association, 107: 81-84.
Miall, A.D. (1985). Architectural-Element Analysis: A
new Method of Facies Analysis Applied to Fluvial
Deposits. Earth Science Reviews, 22: 261-308.
doi:10.1016/0012-8252(85)90001-7
Naish, D.; Martill, D.M. & Frey, E. (2004). Ecology, Syste-
matics and Biogeographical Relationships of Dinosaurs,
Including a New Theropod, from the Santana Formation
(?Albian, Early Cretaceous) of Brazil. Historical Biology,
16 (2-4): 57-70. doi:10.1080/08912960410001674200
Novas, F.E. (2009). The Age of Dinosaurs in South Ame-
rica. Indiana University Press, Bloomington and India-
napolis, Indiana, USA. 458 p.
Novas, F.E.; Cambiaso, A.V. & Ambrosio, A. (2004a). A
new basal iguanodontian (Dinosauria, Ornithischia)
from the Upper Cretaceous of Patagonia. Ameghiniana
41, 75–82.
Novas, F.E.; Dalla Vecchia, F. & Pais, D.F. (2005a). The-
ropod pedal unguals from the Late Cretaceous (Ceno-
manian) of Morocco, Africa. Revista del Museo Argenti-
no de Ciencias Naturales, 7 (2): 167–175.
Oliveira, E.F.; Rangel, C.C.; Alves, D.S.J.; Carvalho, A.A.
& Candeiro, C.R.A. (2011). Short communication on
the geology and vertebrate paleontology of the Adaman-
tina Formation (Bauru Group, Upper Cretaceous), in the
Campina Verde municipality, Minas Gerais State, Bra-
zil. Earth Sciences Research Journal, 15: 41-45.
Sereno, P.C. & Brusatte, S.L. (2008). Basal abelisaurid
and carcharodontosaurid theropods from the Lower Cre-
taceous Elrhaz Formation of Niger. Acta Palaeontologi-
ca Polonica, 53 (1): 15-46. doi:10.4202/app.2008.0102
Rauhut, O.W.M. (2002). Dinosaur teeth from the Barre-
mian of Una, Province of Cuenca, Spain. Cretaceous
Research, 23: 255-263. doi:10.1006/cres.2002.1003
Riccominni, C. (1997). Arcabouço estrutural e aspectos do
tectonismo gerador e deformador da Bacia Bauru no
estado de São Paulo. Revista Brasileira de Geociências,
27(2): 153-162.
Salgado, L.; Coria, R.A.; Arcucci, A.B. & Chiappe, L.M.
(2009). Restos de Alvarezsauridae (Theropoda, Coelu-
rosauria) en la Formación Allen (Campaniano-Maas-
trichtiano), en Salitral Ojo de Agua, Provincia de Río
Negro, Argentina. Andean Geology, 36: 67–80.
doi:10.4067/S0718-71062009000100006
Sampson, S.D.; Carrano, M.T. & Forster, C.A. (2001). A
bizarre predatory dinosaur from the Late Cretaceous of
Madagascar. Nature, 409: 504–506.
doi:10.1038/35054046
Sampson, S.D.; Krause, D.W. (eds.) 2007. Majungasaurus
crenatissimus (Theropoda: Abelisauridae) from the Late
Cretaceous of Madagascar. Society of Vertebrate Pale-
ontology Memoir 8.
Sampson, S.D.; Witmer, L.M.; Forster, C.A.; Krause,
D.W.; O´ Connor, P.M.; Dodson, P. & Ravoavy, F.
(1998). Predatory dinosaur remains from Madagascar:
implications for the Cretaceous biogeography of Gond-
wana. Science, 280: 1048–1051. doi:10.1126/scien-
ce.280.5366.1048
Sankey, J.T.; Brinkman, D.B.; Guenther, M. & Currie, PJ.
(2002). Small theropod and bird teeth from the Judith
River Group (Late Campanian), Alberta. Journal of
Paleontology, 76: 751-763. doi:10.1666/0022-
3360(2002)076<0751:STABTF>2.0.CO;2
Simbras, F.M. (2009). Análise Estratigráfica do Intervalo
Superior do Grupo Bauru (Cretáceo Superior) na
Região de Alfredo Marcondes - SP. Monografia, Depar-
tamento de Geologia, Universidade Federal do Rio de
Janeiro, 78 pp.
Smith, J.B. & Dalla Vecchia, F.M. (2006). An abelisaurid
(Dinosauria: Theropoda) tooth from the Lower Cretace-
ous Chicla Formation of Lybia. Journal of African
Earth Sciences, 46: 240-244. doi:10.1016/j.jafrears-
ci.2006.05.007
Smith, J.B. & Lamanna, M.D. (2006). An abelisaurid from
the Upper Cretaceous of Egypt: implications for thero-
pod biogeography. Naturwissenchaften, 93: 242-245.
doi:10.1007/s00114-006-0092-3
Soares, P.C.; Landim, P.M.B.; Fúlfaro, V.J.; Amaral, G.;
Suguio, K.; Coimbra, A.M.; Sobreiro Neto, A.F.;
Correa, W.A.G.; Castro, C.G.J. (1979) Geologia da
Região Sudoeste do Estado de São Paulo. Simpósio
Regional de Geologia, Núcleo São Paulo, SBG, 2, São
Paulo. Atas do 2º Simpósio Regional de Geologia,
Núcleo de São Paulo, SBG. São Paulo, 307-319.
Wilson, J.A., Sereno, P.C., Srivastava, S., Bhatt, D.K.,
Khosla, A., Sahni, A., 2003. A new abelisaurid (Dino-
sauria, Theropoda) from the Lameta Formation (Creta-
ceous, Maastrichtian) of India. Contribution to the
Museum of Paleontology, University of Michigan, 31:
1–42.
Recibido el 13 de octubre de 2011
Aceptado el 21 de junio de 2012
Publicado online el 10 de enero de 2013
Estudios Geológicos, 69(1), 105-114, enero-junio 2013. ISSN: 0367-0449. doi:10.3989/egeol.40829.170
Teeth of Abelisauridae and Carcharodontosauridae cf. from SW Sao Paulo State, Brazil 113
Furtado.qxd:Estudios 24/06/13 12:19 Página 113