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First fossil record of amiid fishes (Halecomorphi, Amiiformes, Amiidae) from

the Late Cretaceous of Uberaba, Minas Gerais State, Brazil

Article in Alcheringa An Australasian Journal of Palaeontology · March 2013

DOI: 10.1080/03115518.2012.709444

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 First fossil record of amiid fishes (Halecomorphi, Amiiformes,
Amiidae) from the Late Cretaceous of Uberaba, Minas Gerais
State, Brazil
AGUSTÍN G. MARTINELLI, SERGIO BOGAN, FEDERICO L. AGNOLIN, LUIZ C. B. RIBEIRO,
CAMILA L. CAVELLANI, MARA L. F. FERRAZ and VICENTE P. A. TEIXEIRA
Downloaded by [American Museum of Natural History] at 06:08 25 October 2012

MARTINELLI, A.G., BOGAN, S., AGNOLIN, F.L., RIBEIRO, L.C.B., CAVELLANI, C.L., FERRAZ, M.L.F. & TEIXEIRA, V.P.A., iFirst article. First fossil
record of amiid fishes (Halecomorphi, Amiiformes, Amiidae) from the Late Cretaceous of Uberaba, Minas Gerais State, Brazil. Alcheringa, 1–
9. ISSN 0311-5518.

The first fossil amiid fishes (Halecomorphi, Amiiformes) from the Late Cretaceous Marı́lia Formation (Bauru Group) at Uberaba County,
Triângulo Mineiro region (Minas Gerais State, Brazil), are described. The material includes some partial maxillae, a dermopterotic, a
cleithrum, several vertebral centra and teeth. Features such as the absence of a supramaxillary notch on the dorsal edge of the maxilla, a wide
and deep pit on the maxilla for the articulation of the premaxilla, anterior portion of the maxilla with a sub-circular cross-section, teeth with
acrodine cup with strong mesial and distal keels, among others, permit confident referral of the material to the Subfamily Vidalamiine
(Amiidae), previously recognized in Lower Cretaceous strata of northeasthern Brazil. These specimens constitute the first Late Cretaceous
record of this group in Brazil and one of the few in South America.

Agustı´n G. Martinelli [agustı´n_martinelli@yahoo.com.ar], Centro de Pesquisas Paleontológicas Llewellyn Ivor Price, Complexo Cultural e Cientı´fico
Peirópolis (CCCP/UFTM), BR-262, Km 784, Bairro Peirópolis, Uberaba, Minas Gerais, Brazil; Sergio Bogan [sergiobogan@yahoo.com.ar],
Fundación de Historia Natural ‘Félix de Azara’, Departamento de Ciencias Naturales y Antropologı´a, CEBBAD—Universidad Maimónides, Hidalgo
775 piso 7 (1405BDB), Buenos Aires, Argentina. Federico Agnolı´n* [fedeagnolin@yahoo.com.ar], Sección Paleontologı´a de Vertebrados, Museo
Argentino de Ciencias Naturales ‘Bernardino Rivadavia’, Av. Ángel Gallardo 470 (C1405BDB), Buenos Aires, Argentina. Luiz Carlos Borges Ribeiro
[lcbrmg@terra.com.br], Camila Lourencini Cavellani [camila@patge.uftm.edu.br], Mara Lúcia da Fonseca Ferraz [mara@patge.uftm.edu.br] and
Vicente de Paula Antunes Teixeira [vicente@patge.uftm.edu.br], Centro de Pesquisas Paleontológicas Llewellyn Ivor Price, Complexo Cultural e
Cientı´fico Peirópolis (CCCP/UFTM), BR-262, Km 784, Bairro Peirópolis, Uberaba, Minas Gerais, Brazil. *Also affiliated with: Fundación de
Historia Natural ‘Fe´lix de Azara’, Departamento de Ciencias Naturales y Antropologı´a, CEBBAD—Universidad Maimónides, Valentı´n Virasoro 732
(C1405BDB), Buenos Aires, Argentina. Received 15.4.2012; revised 11.6.2012; accepted 20.6.2012.

Key words: Vidalamiinae, Amiidae, Fossil fishes, Bauru Group, Triângulo Mineiro.

THE TRIÂNGULO MINEIRO region (west trian-
gular portion of Minas Gerais State, southeastern
Brazil) is known globally for its Late Cretaceous fossils
collected from several palaeontological sites since the
beginning of the 20th century. The vertebrate record of
this region includes taxonomically diverse fishes (Gayet
& Brito 1989, Bertini et al. 1993, Brito et al. 2006),
anurans (Báez & Perı́ 1989, Báez et al. 2012), turtles
(França & Langer 2005, Gaffney et al. 2011), lizards
(Estes & Price 1973), crocodyliforms (Price 1955,
Carvalho et al. 2004, 2011, Montefeltro et al. 2011)
and dinosaurs, including birds (Price 1951, Santucci &
Bertini 2001, Campos et al. 2005, Kellner et al. 2005,
Novas et al. 2005, 2008, Salgado & Carvalho 2008,
Candeiro et al. in press). The most intensely studied
sites are located in Uberaba County, especially near the
town of Peirópolis, east of Uberaba (Price 1955,
ISSN 0311-5518 (print)/ISSN 1752-0754 (online)
Ó 2012 Association of Australasian Palaeontologists
http://dx.doi.org/10.1080/03115518.2012.709444
Campos & Kellner 1999), and north of Uberaba, in
road cuts of highway BR–050 at km 153 (Salgado &
Carvalho 2008, Ribeiro & Carvalho 2009) and km
153.5 (equivalent to locality km 24 of Bertini et al. 1993
and the Uberaba locality of Gayet & Brito 1989); both
sites have been referred to the ‘Serra da Galga region’.
Other sites in Triângulo Mineiro region to have yielded
Late Cretaceous fossils are located in the counties of
Monte Alegre de Minas (Huene 1931), Iturama
(Kellner et al. 1995), Prata (Goldberg et al. 1995,
Candeiro et al. 2006, Kellner et al. 2006), Verı́ssimo
(Lopes & Buchmann 2008), Campina Verde (Carvalho
et al. 2011) and Gurinhatã (Montefeltro et al. 2011). All
these fossils derive from outcrops of the Bauru Group
(Fernandes & Coimbra 1996, Fernandes 2004). This
group is represented by an Upper Cretaceous con-
tinental sedimentary succession that roughly covers the
northwest of Paraná, west of São Paulo, northeast of
Mato Grosso do Sul, west of Minas Gerais state, and
south of Goiás states (southeastern Brazil; Fernandes &
Coimbra 1996). There have been numerous studies of
 2 AGUSTÍN G. MARTINELLI et al.
its geology, stratigraphy and sedimentology (e.g.,
Fernandes & Coimbra 1996, Dias-Brito et al. 2001,
Fernandes 2004, Batezelli et al. 2007, Paula e Silva et al.
2009), the Bauru Group being one of the most
important Mesozoic vertebrate-bearing units of Brazil.
In this contribution, the first fossil amiid fishes
(Halecomorphi, Amiiformes) from the Marı́lia For-
mation (Bauru Group) are described from three sites
in Uberaba County, Triângulo Mineiro region (Minas
Gerais State, Brazil; Fig. 1). The specimens from
Uberaba constitute the first Late Cretaceous records
of this family in Brazil and one of the few in South
America (Bogan et al. 2010).
At present, the record of Amiiformes in Brazil is
limited to three species of vidalamiines (Amiidae,
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Vidalamiinae) from a Lower Cretaceous deposit in the
northeast of the country. Calamopleurus Agassiz, 1841
is represented by two species: C. cylindricus Agassiz,
1841 from the Aptian Santana Formation and C.
mawsoni (Woodward, 1902) from the Barremian Ilhas
Formation (Maizey 1991, Grande & Bemis 1998).
Cratoamia gondwanica Brito, Yabumoto & Grande,
2008 is a monospecific genus from the Aptian Crato
Formation (Brito et al. 2008). Finally, a possible
Vidalamiini was reported from the early Late Cretac-
eous of the Sergipe Basin (Gallo et al., 2007).
Material and methods
The specimens reported here were discovered during
fieldwork conducted by staff of the CPPLIP (Peirópolis,
Uberaba County, Minas Gerais State, Brazil) in
partnership with national and foreign universities and
museums, from the 20th century to the present. The
material was collected by surface sampling at various
sites, during excavations of crocodyliform and dinosaur
bones, or after screen and dry wash processings from
Fig. 1. Location maps of the three localities with Late Cretaceous amiid remains from Uberaba County, Minas Gerais State, Brazil. 1, Site
‘Ponto 1’. 2, Site ‘Ponto 2’. 3, Site BR-050 km 153 (Serra da Galga).
ALCHERINGA
levels with relatively abundant fossil remains. Each
specimen studied was found isolated; therefore, there is
no certainty that elements from the same locality and
stratigraphic level correspond to a single individual.
The studied material derives from the Serra da Galga
Member, Marı́lia Formation (Bauru Group; Barcelos
1984, Fernandes & Coimbra 1996). The Serra da Galga
Member is composed of fine- to coarse-grained
sandstones, associated with conglomerates in fining-
upwards cycles (Soares et al. 1980, Salgado & Carvalho
2008). According to Goldberg & Garcia (2000), they
were deposited by braided fluvial systems that produced
wide alluvial plains with small lakes. Amiid remains
were found both in beds of coarse sandstone and fine
sandstone (Fig. 2). The age of the Marı́lia Formation is
interpreted to be Maastrichtian based upon fossil
charophytes and ostracods (Dias-Brito et al. 2001).
Institutional Abreviations. CPPLIP, Centro de Pesqui-
sas Paleontológicas Llewellyn Ivor Price, Complexo
Cultural e Cientı́fico de Peirópolis, UFTM, Peiropo-
lis, Uberaba, MG, Brazil.
Systematic palaeontology
Subclass ACTINOPTERYGII Cope, 1887
Division HALECOSTOMI Regan, 1923
Order AMIIFORMES Hay, 1929
Superfamily AMIOIDEA Bonaparte, 1838
Family AMIIDAE Bonaparte, 1838
Subfamily VIDALAMIINAE Grande & Bemis, 1998
Vidalamiine gen. et sp. indet. (Figs 3–7)
Referred material. CPPLIP 167, right cleithrum (Fig.
5B); CPPLIP 266, tooth (Fig. 6A); CPPLIP 1303,
partial right maxilla (Fig. 3); CPPLIP 1304,
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Fig. 2. Stratigraphic profiles from the Marı́lia Formation (Bauru Group) exposed at the studied localities (modified from Novas et al. 2008,
Salgado & Carvalho 2008) indicating the levels at which the amiid remains were found.
precaudal vertebral centrum (Fig. 7); CPPLIP 1308,
anterior portion of right maxilla (Fig. 4A); CPPLIP
1310, posterior portion of left maxilla (Fig. 4C);
CPPLIP 1311, precaudal vertebtral centrum; CPPLIP
1312, posterior portion of right maxilla (Fig. 4B);
CPPLIP 1320, caudal vertebral centrum; CPPLIP
1321, caudal vertebral centrum; CPPLIP 1322, tooth
(Fig. 6B); CPPLIP 1323, left dermopterotic (Fig. 5A).
Locality, formation and age. CPPLIP 1303, CPPLIP
1310 and CPPLIP 1311 derive from the locality
known as ‘Ponto 1 do Price’ (198430 2400 S,
478440 4500 W), located approximately 2 km north of
the rural town of Peirópolis, Uberaba County,
Minas Gerais State, Brazil. CPPLIP 167, CPPLIP
266, CPPLIP 1323 and CPPLIP 1312 come from the
locality known as ‘Ponto 2 do Price’ (198430 1300 S,
478450 0600 W), located approximately 2.3 km north of
the rural town of Peirópolis, Uberaba County,
Minas Gerais State, Brazil. CPPLIP 1304, CPPLIP
1308, CPPLIP 1320, CPPLIP 1321 and CPPLIP 1322
LATE CRETACEOUS AMIIDS FROM BRAZIL 3
were collected from the road cut on highway BR–050
km 153 (198350 3300 S, 488010 4200 W), Serra da Galga,
located 25 km north of Uberaba city, where the
dinosaur Uberabatitan ribeiroi Salgado & Carvalho,
2008 was unearthed (Salgado & Carvalho 2008). All
specimens derive from the Serra da Galga Member,
Marı́lia Formation (Maastrichtian), Bauru Group.
Description. Available jaw material includes only four
isolated remains (Figs 3–4). Specimen CPPLIP 1303
preserves nine subcircular tooth bases. The first
preserved tooth base is smaller than the others, which
are subequal in size and morphology (Fig. 3). The
external surface of the entire maxilla is smooth, as are
the surfaces of the remaining bones. CPPLIP 1303 is
dorsoventrally low; its dorsal margin is concave and
its ventral margin gently convex. The anterior ramus
is robust, rod-like and dorsally oriented, being
subcircular in cross-section. Unfortunately, the ante-
rior process for contact with the premaxilla is only
partially preserved. The posterior portion of the
 4 AGUSTÍN G. MARTINELLI et al.
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Fig. 3. Amiidae gen. et sp. indet. CPPLIP 1303, right maxilla in
lateral (A), medial (B), ventral (C) and dorsal (D) views. Grey areas
indicate broken surfaces. Scale bar ¼ 5 mm.
maxilla is transversely flattened and dorsoventrally
expanded. This portion has a wide and deep medial
concavity at its posterior portion. There is no
supramaxillary notch in the preserved portion.
Based on CPPLIP 1308, the anterior ramus of the
maxilla is robust, acute and dorsomedially protruding
(Fig. 4A). This process has a deep and wide pit located
on its ventral surface for articulation with the
premaxilla. This feature extends posteriorly towards
the level of the first tooth base (Fig. 4A). This
specimen preserves only the bases of the first three
teeth.
The available posterior portions of maxillae are
fragmentary. In CPPLIP 1312, the maxilla is dorso-
ventrally high and transversely narrow, with small
vascular foramina on its external surface. This portion
of bone preserves only the bases of four teeth, which
slightly decrease in size posteriorly. The tooth bases
are closer to the external edge in this specimen (Fig.
ALCHERINGA
4B). CPPLIP 1310 is a slightly smaller specimen with
a slender dorsal process, which is broken at the top
(Fig. 4C). On the internal surface, it bears a shallow
concavity. As in CPPLIP 1312, there is evidence of
small vascular foramina on the external surface. Only
four poorly preserved tooth bases are present.
A single and incomplete left dermopterotic
(CPPLIP 1193; Fig. 5A) can be referred tentatively
to Amiidae. It is sub-rectangular in dorsal outline,
with a slightly protruding posterolateral process. The
shape of this element is similar to Pachyamia Chalifa
& Tchernov, 1982 with a slightly concave lateral
margin (Chalifa & Tchernov 1982, Grande & Bemis
1998), but with a rather more developed poster-
olateral corner. The sub-rectangular shape with the
anterior portion broad is diagnostic of vidalamiins
and different to the anteriorly tapering dermopterotic
of other amiids (Grande & Bemis 1998). The
ornamentation consists of simple and elongated ridges
and grooves radiating from the posterolateral corner
(Fig. 5A). The lateral edge is blunt and more elevated
than the dorsal plate. Laterally, the descending
process (i.e., ventral process) is laminar, smooth and
concave.
A single and incomplete right cleithrum (CPPLIP
167; Fig. 5B) is available. Its external surface shows
well-developed ornamentation in the junction between
the anteroventral and dorsal processes. In the
Brazilian specimen, as is the case in Melvius Bryant,
1987 (Bryant 1987, Grande & Bemis 1998), there are
well-developed grooves and ridges along the posterior
edge of the dorsal process of the bone. The angle
between the anteroventral and dorsal processes is ca
908 in this bone, a condition present in most
vidalamiines and several amiines (Grande & Bemis
1998). The posteroventral margin is gently curved.
The anteroventral process is very thick and is
subvertically oriented (Fig. 5B).
Several teeth were recovered from the three
localities that can be referred to Amiidae (see
discussion). For descriptive purposes we figure only
CPPLIP 266 and CPPLIP 1322 (Fig. 6). All teeth
found in Uberaba preserve only the rigid acrodine cup
and a small portion of the crown. Unfortunately,
most of the crown is always broken away due to its
fragile nature. The teeth are sharp, conical and are
gently curved towards the lingual margin. They have a
subcircular base in cross-section; the base is separated
from the crown by a well-developed constriction. The
crown is composed of an acrodine cup, that is
labiolingually compressed and shows very well-devel-
oped mesial and distal keels (Fig. 6).
There are four caudal and precaudal vertebral
centra preserved with amphicoelous articular surfaces.
The vertebral articular surfaces are subcircular to
rhombic in contour. The notochordal canal is closed
in CPPLIP 1304 (Fig. 7) and CPPLIP 1311. In
CPPLIP 1321 and CPPLIP 1320, the centrum is
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Fig. 4. Amiidae gen. et sp. indet. A, CPPLIP 1308, anterior portion of right maxilla in medial and ventral views. B, CPPLIP 1312, posterior
portion of right maxilla in lateral and ventral views. C, CPPLIP 1310, posterior portion of left maxilla in lateral and ventral views. The arrow
indicates the deep concavity for reception of the premaxillary bone. Grey areas indicate broken surfaces. Scale bars ¼ 5 mm.
perforated by means of a small notochordal canal. In
CPPLIP 1304 and CPPLIP 1320, there are four
depressions on the dorsal margin for the articulation
with the autogenous neural arch (Fig. 7). There are
well-developed depressions for the articulation with
the parapophyses, which were not fused with the
vertebral centrum. The ventral margin of both speci-
mens exhibits two longitudinal ridges for articulation
with the autogenous haemal arch (Fig. 7).
Discussion
Taxonomy
The described specimens have a combination of
characters that, collectively, are diagnostic of the
Amiidae clade Vidalamiinae (sensu Grande & Bemis
1998). Available maxillae are very similar in general
contour and gross morphology to amiids of the
Vidalamiinae. One of the most salient synapomor-
phies for this subfamily is the presence of a deep
posterior notch (i.e., postmaxillary notch; clearly
observed in Vidalamia White & Moy-Thomas, 1941,
Pachyamia and Cratoamia; Grande & Bemis 1998,
Brito et al. 2008) in the maxilla. Regrettably, this
feature is not preserved in the available specimens
from the Marı́lia Formation. Nevertheless, in the
maxilla CPPLIP 1303, there is a triangular posterior
edge, which could correspond to the anterior edge of
the postmaxillary notch (Fig. 3), resembling the
condition of vidalamiines. Unfortunately, the ventral
and dorsal borders are mostly broken off preventing
unambiguous interpretations.
Another diagnostic trait of Vidalamiinae is the
absence of a supramaxillary notch (Grande & Bemis
LATE CRETACEOUS AMIIDS FROM BRAZIL 5
1998, Brito et al. 2008). The absence of this notch is
also evident in specimen CPPLIP 1303 (Fig. 3).
Previously, Bryant (1987) considered this feature to
be diagnostic of the vidalamiine genus Melvius,
although later authors indicated its presence in other
vidalamiines (see discussion by Grande & Bemis
1998). Another feature considered diagnostic of
Vidalamiinae is the presence of a wide and deep pit
for the articulation of the premaxilla (Grande &
Bemis 1998). Specimen CPPLIP 1308 from the
Marı́lia Formation has a wide and well-defined
concavity at the anteroventral margin of the maxilla
(Fig. 4A) that nearly reaches the anterior margin of
the first tooth base. This condition was formerly
considered by Bryant (1987) to be synapomorphic for
Melvius. However, the presence of this feature in other
vidalamiines such as Calamopleurus and Pachyamia
indicates that it is a unique apomorphy of Vidalamii-
nae (Grande & Bemis 1998, p. 397; see also Patterson
& Longbottom 1989). In addition, a third diagnostic
feature allowing inclusion of the studied specimens
within Vidalamiinae (Grande & Bemis 1998) is the
sub-circular cross-section of the anterior portion of
the maxilla of the specimens CPPLIP 1303 (Fig. 3)
and CPPLIP 1308 (Fig. 4A).
Teeth from the Marı́lia Formation have an acrodine
cup that is labiolingually compressed and strongly
keeled on its mesial and distal margins (Fig. 6). This
morphology is synapomorphic for the Vidalamiinae
clade (Grande & Bemis 1998; see also below).
The vertebral centra from the Marı́lia Formation
(Fig. 7) are also reminiscent of amiid fishes. The
specimens lack fused parapophyses, a plesiomorphic
trait that distinguishes the studied specimens from the
Amiinae subfamily (Grande & Bemis 1998). The
general morphology of the available vertebrae is
 6 AGUSTÍN G. MARTINELLI et al.
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Fig. 5. Amiidae gen. et sp. indet. A, CPPLIP 1323, left
dermopterotic in dorsal view. B, CPPLIP 167, right cleithrum in
lateral and medial views. Scale bars ¼ 10 mm.
rather similar to other vidalamiines, being very similar
in proportions to that of Melvius (Bryant 1987).
However, the specimens reported here differ from the
latter genus in lacking the ventrolateral concavities of
the centrum (Grande & Bemis 1998).
The Vidalamiinae includes two subclades: Vidala-
miini and Calamopleurini (sensu Grande & Bemis
1998). The studied maxillae differ from the calamo-
pleurinine Calamopleurus (Grande & Bemis 1998) in
lacking external ornamentation and in having a
concave dorsal margin and a convex ventral one.
The absence of external ornamentation on the
maxillae distinguishes the Brazilian specimens from
remaining vidalamiines, with the exceptions of Cra-
toamia and Maliamia (Patterson & Longbottom 1989,
Brito et al. 2008). Cratoamia differs from the Uberaba
material by its weakly developed notch for the
supramaxilla in the dorsal edge of the maxilla (Brito
et al. 2008)—a feature that is absent in CPPLIP 1303.
Due to the isolated and fragmentary nature of the
specimens described here, a generic or specific
determination of the individuals is not possible, and
they are consequently assigned to Vidalamiinae indet.
ALCHERINGA
Fig. 6. Amiidae gen. et sp. indet. A, CPPLIP 266 and B, CPPLIP
1322, isolated teeth. Scale bar ¼ 1 mm in A and 0.5 mm in B.
Isolated teeth from the Bauru Group
Most fish fossils from the Bauru Group are isolated
specimens recovered from several localities of south-
eastern Brazil (Bertini et al. 1993, Brito et al. 2006,
Azevedo et al. 2007). Isolated teeth are the most
frequently discovered material. Hundreds of teeth
with the morphology described above were recovered
from the Marı́lia Formation at the three studied
localities in Uberaba County. These specimens have
the well-developed acrodin cup, constricted basally,
with strongly keeled mesial and distal margins (Fig. 6)
that is typical of vidalamiine amiids (Grande & Bemis
1998). Similar shaped isolated teeth have been
reported previously from exposures of the Bauru
Group at several fossiliferous localities, including the
sites reported here, and were referred to erythrinid-
like Characiformes fishes (Gayet & Brito 1989, Brito
et al. 2006, figs 4D, 4F). We consider this identification
to be not well founded and the teeth reported by those
authors are indistinguishable from those known in
Cretaceous vidalamiine fishes. Additionally, Azevedo
et al. (2007, fig. 2L, M) reported similar teeth from the
Flórida Paulista locality (São Paulo State) from the
Adamantina Formation, which were assigned to
Characiformes indet. Based on the features previously
noted, the material may belong to amiid fishes, thus
indicating the presence of this group also in the
Adamantina Formation. Based upon this resemblance
and the fact that other unambiguous material (e.g.,
maxillae, vertebrae) of vidalamiine amiids are now
known from three Bauru Group localities, these
isolated teeth from the Bauru Group are considered
here indeterminate Vidalamiinae (Amiidae).
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Fig. 7. Amiidae gen. et sp. indet. CPPLIP 1304, vertebral centrum in dorsal (left above), axial articular surfaces (left middle and left below),
ventral (right above) and lateral (right below) views. Scale bar ¼ 10 mm.
Palaeobiogeographic importance of Amiidae in the
Bauru Group
During the Early Cretaceous, South America was
slightly separated from Africa by a shallow sea
(Scotese & Golonka 1992). At that time, all the South
American fossil amiid records derive from north-
eastern Brazil and are attributable to three taxa
referable to Vidalamiinae. Calamopleurus cylindricus
(Aptian Santana Formation; Maizey 1991) and C.
mawsoni (Barremian Ilhas Formation; Grande &
Bemis 1998) were, together with Cretaceous and
Cenozoic African taxa (see Grande & Bemis 1998),
included in the Tribe Calamopleurini. This genus was
also reported in Lower Cretaceous marine strata of
Equatorial Guinea (Africa; Taverne 1997). These taxa
were distributed along the coasts of the shallow
marine sea separating Africa and South America. The
other vidalamiine tribe, Vidalamiini, is represented in
South America by the single genus and species
Cratoamia gondwanica (Aptian Crato Formation;
Brito et al. 2008). The biogeography of the early
Vidalamiini was assessed by Grande & Bemis (1998)
and Brito et al. (2008) who indicated that these taxa
were distributed along shallow coastal waters of a
seaway extending from North America eastward to
LATE CRETACEOUS AMIIDS FROM BRAZIL 7
the Tethys region and Caribbean and northern South
American seas.
On the other hand, the Late Cretaceous record of
Amiidae in South America is rather patchy and
biased. Thus far, only three Late Cretaceous fossili-
ferous assemblages have yielded amiid remains from
the entire continent (Bogan et al. 2010; this contribu-
tion). Two fossiliferous localities (i.e., El Anfiteatro
and Cerro Tortuga localities) are located in the north
Patagonian Rı́o Negro province, Argentina (Bogan
et al. 2010). An isolated cleithrum from the Turonian
El Anfiteatro locality was originally described by
Salgado et al. (2009) as belonging to a probable
Osteoglossomorpha. However, this specimen was
later identified as an indeterminate Amiidae (Bogan
et al. 2010). From the uppermost Cretaceous (Maas-
trichtian Allen Formation) at the Cerro Tortugas
locality, Bogan et al. (2010) reported an isolated
dentary bone that belongs to the Amiidae. Both
isolated specimens from northern Patagonia have
indeterminate affinities below family level. To these
Late Cretaceous records, we add the material
described here from the Bauru Group.
Such findings in non-marine Upper Cretaceous
strata of Brazil and Argentina contrast with the
 8 AGUSTÍN G. MARTINELLI et al.
Lower Cretaceous record of Amiidae in South
America. The latter examples are preserved in
brackish-water deposits of marine coasts (Grande &
Bemis 1998, Brito et al. 2008). This palaeoenviron-
mental difference is interesting because living amiid
fishes currently inhabit freshwater environments
(Grande & Bemis 1998). Although the Late Cretac-
eous record from continental environments is still
sparse, this pattern may indicate a shift in the
behaviour of these fishes in the mid-Cretaceous.
However, this hypothesis is potentially biased by the
incomplete representation of freshwater sediments in
the Early Cretaceous of South America.
Downloaded by [American Museum of Natural History] at 06:08 25 October 2012
Conclusions
The described material represents some of the few
amiid records from the Late Cretaceous of South
America and the first records of this age in Brazil.
These fossils increase the known fish diversity of the
Bauru Group, which at present is represented by
Chondrichthyes (personal observation), Lepisostei-
formes, Osteoglossiformes, Characiformes, Perci-
formes, Siluriformes and Neoceratodontidae (e.g.,
Bertini et al. 1993, Brito et al. 2006).
The Uberaba specimens indicate the persistence of
Subfamily Vidalamiinae into the Late Cretaceous of
South America. The fragmentary and poorly infor-
mative nature of the Late Cretaceous specimens
precludes definitive referral to a genus. The Late
Cretaceous amiid remains from South America occur
in freshwater deposits, matching the habits of living
forms in the clade.
Acknowledgements
This work was aided by logistic and financial
support from the Fundação de Amparo à Pesquisa
do Estado de Minas Gerais (FAPEMIG), the
Conselho Nacional de Desenvolvimento Cientı́fico
e Tecnológico (CNPq), the Coordenação de Aper-
feiçoamento de Pessoal de Nı́vel Superior (CAPES),
the Fundação de Ensino e Pesquisa de Uberaba
(FUNEPU), the Fundação Peirópolis, the Funda-
ción de Historia Natural ‘Félix de Azara’, and the
Universidad Maimónides (UM). Technicians from
the CPPLIP provided invaluable support during
fieldwork. J. Kriwet, an anonymous reviewer and
the editor S. McLoughlin provided useful comments
and suggestions that greatly improved the manu-
script.
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