Fauna Flora Cretacea Uberaba

www.elsevier.com/locate/jsames

Journal of South American Earth Sciences 25 (2008) 203–216

The Late Cretaceous fauna and flora of the Uberaba area(Minas Gerais State, Brazil)

Carlos Roberto A. Candeiro a,b,*, Adriano R. Santos a, Lılian P. Bergqvist b,Luiz Carlos B. Ribeiro c,d, Sebastian Apesteguıa e

a Museu de Minerais e Rochas, Instituto de Geografia, Universidade Federal de Uberlandia, Av. Joao Naves de Avilla, 2160,

Sta. Monica, Uberlandia, Minas Gerais, Brazilb Laboratorio de Macrofosseis, Departamento de Geologia, Universidade Federal do Rio de Janeiro, Av. Brig. Trompski,

S/N, Cidade Universitaria, Rio de Janeiro, RJ, Brazilc Fundac�ao Municipal de Ensino Superior de Uberaba-FUMESU/Centro de Ensino Superior de Uberaba-CESUBE/

Centro de Pesquisas Paleontologicas Llewellyn Ivor Price, BR-262, KM 784, Peiropolis, Uberaba, Minas Gerais, Brazild Universidade de Uberaba-UNIUBE/Instituto de Formac�ao de Educadores, Dep. De Biologia, Av. Nene Sabino, 1801, Universitario,

38055-500, Uberaba, Minas Gerais, Brazile Museo Patagonico de Ciencias Naturales, Av. Roca 1250, General Roca, Rıo Negro, Argentina

Abstract

The Uberaba area, in Minas Gerais State, Brazil, yields a rich continental fauna and flora from the Late Cretaceous Uberaba andMarılia formations. This paper reviews the diversity of the biota recorded from these formations. The most significant taxa from Pei-ropolis are the frog Baurubatrachus pricei, the turtle Cambaremys langertoni, the lizard Pristiguana brasiliensis, the crocodyliforms Itasu-

chus jesuinoi, Peirosaurus tormini and Uberabasuchus terrificus, the titanosaurian Baurutitan britoi, Trigonosaurus pricei, Aeolosaurus sp.,indeterminate titanosaurians, and abelisaurid, carcharodontosaurid and maniraptoran theropods. Together with faunas of a similar agein Argentina and Madagascar, the assemblages contribute to a better understanding of Late Cretaceous Gondwanan faunas as a whole.� 2007 Elsevier Ltd. All rights reserved.

Keywords: Upper Cretaceous; Uberaba; Fauna and flora; Brazil; Gondwana

Resumo

A fauna e flora da regiao de Uberaba (Neocretaceo), Estado de Minas Gerais, Brasil provem das formac�oes Uberaba e Marılia sendorepresentada de taxa de origem continental. Este artigo revisa a diversidade da biota registrada nestas formac�oes. Os taxa mais signifi-cantes de Peiropolis sao conhecidos pelo sapo, Baurubatrachus pricei, a tartaruga Cambaremys langertoni, o lagarto, Pristiguana brasil-

iensis, e pelos crocodilianos Itasuchus jesuinoi, Peirosaurus tormini e Uberabasuchus terrificus. Os dinossauros sao representados porrestos de Baurutitan britoi, Trigonosaurus pricei, Aeolosaurus sp., titanossaurıdeos, abelisaurıdeos, carcarodontosaurıdeos e maniraptor.A assembleia fossilıfera proveniente de Uberaba e de origem gondwanica e apresentando afinidades com a fauna e flora do Neocretaceoda Argentina e Madagascar.� 2007 Elsevier Ltd. All rights reserved.

Palavras chave: Cretaceo Superior; Uberaba; Fauna e flora; Brasil; Gondwana

0895-9811/$ - see front matter � 2007 Elsevier Ltd. All rights reserved.

doi:10.1016/j.jsames.2007.06.005

* Corresponding author. Address: Museu de Minerais e Rochas, Instituto de Geografia, Universidade Federal de Uberlandia, Av. Joao Naves de Avilla,2160, Sta. Monica, Uberlandia, Minas Gerais, Brazil. Fax: +51 34 3214 2925.

E-mail addresses: [email protected], [email protected] (C.R.A. Candeiro).

mailto:[email protected]

mailto:[email protected]

204 C.R.A. Candeiro et al. / Journal of South American Earth Sciences 25 (2008) 203–216

1. Introduction

The Uberaba area (UA) in southwestern Minas GeraisState, Brazil, has yielded important Late Cretaceous faunaand flora (Fig. 1). The paleontological importance of thissite has been recognized since the beginning of the 1940s(Price, 1961), but a comprehensive, systematic review ofthe rich UA Bauru Group fossil biota has yet to be per-formed. In the past 15 years, a large number of beautifullypreserved fossil invertebrates and vertebrates (Appendix A)have been recovered from the Coniacian–Santonian Uber-aba Formation (Dias-Brito et al., 2001) and late Maas-trichtian Marılia Formation (Dias-Brito et al., 2001) atUberaba. Despite these advances, current understandingof the Upper Cretaceous biota of the site remains far fromcomplete. We review the up-to-date knowledge of thefauna of the site and compare it with faunas of other con-temporaneous Gondwanan localities, notably those com-ing from nearby Patagonia (Argentina) and Madagascar.

2. Geological setting

The Uberaba area yields fossils from two of the threeformations of the Bauru Group: Uberaba and Marılia;the Adamantina Formation is absent (Fig. 2). The fossil-bearing units present in the Triangulo Mineiro region,which includes the Uberaba area, and crop out in thenortheastern region of the Bauru Basin (Fernandes andCoimbra, 1996). They were deposited above the basaltsof the Serra Geral Formation (Sao Bento Group, ParanaBasin), the sandstones of the Botucatu Formation (SaoBento Group), the metamorphic Proterozoic outcrops ofthe Araxa and Canastra groups (Sanfranciscana Basin),and the Mesozoic intrusive event of the Soerguimento doAlto Paranaıba (Suguio et al., 1979).

The older unit of the site is the Coniacian–Santonian(Dias-Brito et al., 2001), 140 m thick, Uberaba Formation(Fernandes and Coimbra, 1996), which is mainly exposedat Uberaba in Minas Gerais State. It is composed of lime-stones, sandstones, and a carbonate-cemented basal con-glomerate and includes an important volcanoclastic unit(Barcelos, 1984). Among the fossils collected are microfos-sils, turtles, titanosaurian sauropods, indeterminate dino-saur remains, and ichnofossils. The younger unit, the180 m thick Marılia Formation (Fernandes and Coimbra,1996), is exposed in Goias and Sao Paulo states and inthe Triangulo Mineiro region. It consists of fine- to med-ium-grained sandstones intercalated with conglomerates.The sandstones are well cemented and bear calcium car-bonate concretions (Fulfaro and Barcelos, 1991). Someauthors, such as Bertini et al. (1993), suggest the MarıliaFormation is Campanian–Maastrichitan, based on verte-brate fossils. Recently, Dias-Brito et al. (2001) have pro-posed an Upper Maastrichtian age for the MarıliaFormation, based on ostracods and charophytes.

The Marılia Formation has an irregular outcrop patternwithin the Triangulo Mineiro, as is the case in western Sao

Paulo state. According to Suguio (1973, 1980) and Barcelosand Suguio (1987), it was deposited as a series of coalescentalluvial fans, previously reworked and redeposited by abraided fluvial system, associated with calcretes andcalcareous lacustrine environments with intermittent dryperiods. Barcelos (1984) subdivides the Marılia Formationinto three members, Echapora, Ponte Alta, and Serra daGalga, the latter two restricted to the Uberaba area. Thebasal Ponte Alta Member (Fig. 3) was characterized sedi-mentologically by Garrido et al. (1992) as a ‘‘calcareouswhite member” composed of thin and medium limestoneswith pebbles and calcareous nodules deposited in playalakes, with alkaline water supersaturated with calciumbicarbonates. The upper Serra da Galga Member (Fig. 3)is characterized as a ‘‘member of white limestone andconglomerate” and composed of conglomerate and thin-to medium-grained limestones, with bleached colorationin an arkosic matrix. It shows an abundance of cross-strat-ification and invertebrate bioturbation (Suguio, 1973,1980).

The Marılia Formation includes several Cretaceouscontinental fossiliferous units (Fernandes and Coimbra,1996) (Fig. 3). In addition to ostracods and charophytes,there is a diverse tetrapod fauna that includes chelonians,crocodyliforms, and several dinosaurs, including abelisau-rid, carcharodontosaurid and maniraptoran theropods(Candeiro, 2002; Candeiro et al., 2004a,b; Novas et al.,2005). In Minas Gerais State, titanosaurian sauropod(sensu Wilson and Upchurch, 2003) remains are abundantin the regions of Monte Alegre de Minas and Uberabaalong the Uberaba–Uberlandia highway, as well as atCampina Verde in Minas Gerais State (von Huene,1931; Campos and Bertini, 1985; Kellner, 1996; Kellnerand Campos, 2000; Santucci and Bertini, 2002, 2006; Can-deiro et al., 2004b). The record includes Aeolosaurus sp.(from the UA) and indeterminate forms (from the UAand Monte Alto in Sao Paulo State).

3. Age of fossil-bearing sediments in the Uberaba area

The Upper Cretaceous age of the Bauru Group wasemphasized by Bertini et al. (1993) and Santucci andBertini (2002) on the basis of similarities between theUA titanosaurian and Aeolosaurus sp., among others.In addition, the microfossil assemblage reported byDias-Brito et al. (2001) favors a late Maastrichtian agefor the Marılia Formation and a Coniacian–Santonianage for the Uberaba Formation. The Upper Cretaceousostracod fauna reported from Marılia Formation, asexposed in the UA, includes Ilyocypris setembrinopetrii,Ilyocpris sp. 1, Neuquenocypris minor mineira, Virgatocy-

pris mezzalirai, Altanicypris australis, Lycopterocypris

angulata, Candonopsis sp., and gen. et sp. Indet. 1 and2. Of these, Neuquenocypris minor and Ilyocypris particu-larly link this fauna with that of the Neuquen Basin,Argentina (Musacchio and Simeoni, 1991; Dias-Britoet al., 2001).

Fig. 1. Approximate geographic distribution of the fossiliferous localities of the Uberaba area, Minas Gerais State, Brazil.

C.R.A. Candeiro et al. / Journal of South American Earth Sciences 25 (2008) 203–216 205

Fig. 2. Geologic map of the Triangulo Mineiro region, Minas Gerais State, Brazil (modified from Fernandes and Coimbra, 1996).


4. Remarks on the biota of the Uberaba area

Few invertebrates are known from UA. Pulmonate gas-tropods occur mainly in the Marılia Formation, and asmentioned previously, ostracods are well represented andvery diverse in the UA fauna (Appendix A). The charo-phyte assemblage is represented by Feistiella cf. globosa,Feistiella cf. costata, Sphaerochara latifasciata, Nitellopsis?sp., Gobichara (Pseudoharrisichara) groeberi, Gobichara

(Pseudoharrisichara) sp., and Chara tomentosa and has abroad distribution in the Bauru Basin (Dias-Brito et al.,2001).

There have been a significant number of new fossil dis-coveries in the UA in the past 15 years (Appendix A; Figs.4a–e, 5a–e). Among the tetrapods, a single ‘leptodactylid’species, Baurubatrachus pricei Baez and Peri, 1989(Fig. 5a), represents the only anuran from Uberaba, whereascrocodyliforms are represented by large mesoeucrocodylian

taxa, including Peirosaurus tormini and Itasuchus jesuinoi

Price, 1955, and Uberabasuchus terrificus Carvalho et al.,2004 (Figs. 5c–e). Chelonians are represented by Cambare-

mys langertoni Franc�a and Langer, 2005 and turtle remains(Fig. 4d) are well known from the Uberaba region, but whendiversity and abundance are taken into account, dinosaursform the largest part of the record (see Kellner and Campos,2000; Marinho et al., 2003; Candeiro and Bergqvist, 2004;Candeiro et al., 2004a, 2006a). Remains of titanosauriansand abelisaurid and carcharodontosaurid theropods occurfrequently in the UA (Fig. 4a, b, and e). Reported UA tita-nosaurs include Baurutitan britoi Kellner et al., 2005, Trigo-

nosaurus pricei Campos et al., 2005, and Aeolosaurus sp. andindeterminate titanosaurian species, as well as eggshell frag-ments referred to ‘‘Megaloolithidae” (Fig. 4c) (Magalhaes-Ribeiro, 2002a). Among the theropods, the presence ofabelisaurid and carcharodontosaurid teeth is noteworthy(e.g., Candeiro, 2002; Marinho et al., 2003; Candeiro

Fig. 3. Stratigraphy of the Uberaba area (modified from Garrido et al., 1992).


et al., 2004a), as well a first maniraptoran claw describedrecently by Novas et al. (2005). Some teeth were previouslymistaken as those of dromaeosaurids and troodontids(Franco and Bertini, 1997; Franco, 1999), but most actuallybelong to carcharodontosaurids. The former family, how-ever, is present in the Cretaceous of South America (Cande-iro, 2002). The Carcharodontosauridae are characterized bytheir enamel wrinkles (Candeiro, 2002; Candeiro et al.,2004a) and apparently survived well into the Late Creta-ceous (Martinelli and Forasepi, 2004; Apesteguıa et al.,2004a; cf. Apesteguıa, 2002). With regard to sauropods,Santucci and Bertini (2002) and Candeiro et al. (2006a,b)demonstrate the presence of the titanosaurian Aeolosaurus

sp. at Uberaba (Marılia Formation), approximately coevalwith Aeolosaurus species from Argentina. In addition, Sant-ucci (2002) reports a new titanosaurian species from boththe Uberaba and Marılia formations.

4.1. Fossiliferous localities within the UA

Fourteen Upper Cretaceous fossiliferous localitieswithin the Uberaba region are considered in this study(Fig. 1, Appendix A). The localities in chronological orderare as follows:

Locality 1: ‘‘Ponto 1 do Price,” Peiropolis district,Uberaba town. Horizon: Marılia Formation, Serra daGalga Member. Age: late Maastrichtian.Fossils: Plantae: Pyteridophyta: Marsilea sp.?; inverte-brates: Mollusca; Bivalvia; Musculium sp., Gastropoda;Physidae; Pysa aridi, Vivirapidae; Anodontites (Anodon-

tites) pricei, Florencia peiropolensis, Viviparus souzai;Vertebrata: Pisces; Characiformes indet., Perciformesindet., Siluriformes indet.; Chelonia: gen. et sp. Indet.,Podocnemidae indet.; Lepidosauria: Pristiguana brasili-

ensis; Mesoeucrodylia: Itasuchus jesuinoi, Peirosaurus

tormini, Uberabasuchus terrificus; Dinosauria: Sauro-poda: ‘‘Megaloolithidae” indet., Baurutitan britoi, Trig-onosaurus pricei Titanosauria (sensu Wilson andUpchurch, 2003); Theropoda: Abelisauridae indet., Car-charodontosauridae indet, Maniraptora indet.Latitude/longitude coordinates: 19�43013.200S/47�44017.900W.References: Price (1951, 1955, 1961), Estes and Price(1973), Mezzalira (1974), Bonaparte (1978, 1996,1999), Buffetaut (1982, 1985), Gayet and Brito(1989), Gasparini et al. (1991), Bertini et al. (1993),Kellner (1995), Albino (1996), Gasparini (1996), Sal-gado et al. (1997), Candeiro (1998, 2002), Fernandes

Fig. 4. (a) Abelisaurid tooth in labia view; (b) carcharodontosaurid tooth in labial view; (c) ‘‘megaloolithid” eggshell; (d) turtle carapace in dorsal view; (e)crocodyliform cervical vertebrae in posterior view; (f) titanosaurian femur. Scale bar in a and b = 10 mm; c–e = 20 mm; f = 150 mm.


(1998), Magalhaes-Ribeiro (1998, 1999a,b, 2000a,b,2001a,b 2002a,b), Pol (1999, 2003), Salgado (2000),Ortega et al. (2000), Avilla (2002), Gobbo-Rodrigues(2002), Santucci (2002), Senra (2002), Santucci andBertini (2002), Evans et al. (2002), Powell (2003),Ribeiro (2003), Franc�a and Langer (2003), Marinhoand Candeiro (2003a,b, 2005), Krause et al. (2003),Wilson and Upchurch (2003), Candeiro and Bergqvist(2004), Candeiro et al. (2004a,b), Carvalho et al.(2004), Candeiro and Martinelli (2006), Camposet al. (2005), Kellner et al. (2005), Novas et al.(2005).

Locality 2: ‘‘Ponto 2 do Price,” Peiropolis district, Uber-aba town. Horizon: Marılia Formation, Serra da GalgaMember. Age: late Maastrichtian.Fossils: Chelonia: Cambaremys langertoni; Dinosauria:Saurpoda: Titanosauria indet.; Theropoda: Abelisauri-dae indet., Carcharodontosauridae indet.Latitude/longitude coordinates: 19�43012.000S/47�45004.400W.

References: Price (1961), Bertini et al. (1993), Fernandes(1998), Sanchız (1998), Franco (1999), Candeiro (2002),Santucci and Bertini (2002), Marinho and Candeiro(2003b), Venczel and Csiki (2003), Candeiro and Bergq-vist (2004), Candeiro et al. (2004a,b), Franc�a and Lan-ger (2005).

Locality 3: Peiropolis Quarry, Peiropolis district, Uber-aba town. Horizon: Marılia Formation, Ponte AltaMember. Age: late Maastrichtian.Fossils: Pisces indet.Latitude/longitude coordinates: 19o42052.400S/47o45016.100W.References: Candeiro (1998), Fernandes (1998), Cande-iro and Bergqvist (2004).

Locality 4: ‘‘Ponto 1 Langerton,” Peiropolis district,Uberaba town. Horizon: Uberaba Formation. Age:Coniacian-Santonian.Fossils: Dinosauria indet. (nests).Latitude/longitude: 19�44042.500S/47�44032.200W.References: Magalhaes-Ribeiro (2002a,b).

Fig. 5. (a) Partial skeleton of Baurubatrachus pricei in dorsal view (modified from Baez and Peri, 1989); (b) left humerus of Pristiguana brasiliensis incaudal view (modified from Estes and Price, 1973); (c) left premaxilla of Peirosaurus tormini in lateral view (modified from Price, 1955); (d) skull ofUberabasuchus terrificus in left view; (e) skull of Itasuchus jesuinoi in left view (modified from Rasmusson, 2002). Scale bar in a–c = 10 mm andd–e = 20 mm.


Locality 5: ‘‘Mumbuco” Freeway BR-262/KM-780,Uberaba town. Horizon: Marılia Formation, Serra daGalga Member. Age: late Maastrichtian.Fossils: Dinosauria: Titanosauria indet.Latitude/longitude: 19�43029.300S/47�42055.200W.References: Campos and Kellner (1999), Kellner andCampos (2000), Santucci and Bertini (2002), Trottaet al. (2002), Candeiro et al. (2004b).

Locality 6. Ponte Alta Quarry, Ponte Alta district,Uberaba town. Horizon: Marılia Formation, Ponte Altaand Serra da Galga Member. Age: late Maastrichtian.

Fossils: Dinosauria: gen. et sp. indet. (eggs).Latitude/longitude: 19�42038.300S/47�40027.900W.References: Magalhaes-Ribeiro (2002a), Santucci(2002), Candeiro et al. (2004b).

Locality 7: ‘‘Manoel Mendes Neighborhood/AvenueLeopoldina de Oliveira,” Uberaba town. Horizon:Uberaba Formation. Age: Coniacian-Santonian.Fossils: Anura: Baurubatrachus pricei; Dinosauria: gen.et sp. indet.; Sauropoda indet.Latitude/longitude: 19�45029.0000S/47�530053.500W.References: Baez (1985), Baez and Peri (1989), Santucci(2002).


Locality 8: ‘‘Avenue Randolfo Borges Junior Outcrop”

near the Uberaba River, Uberaba town. Horizon: Uber-aba Formation. Age: Coniacian-Santonian.Fossils: Dinosauria indet.Latitude/longitude: 19�43016.500S/47�5703000W.References: Santucci (2002), Kellner and Campos(2000).

Locality 9: Mangabeira Station, Uberaba town. Hori-zon: Marılia Formation, Serra da Galga Member.Age: late Maastrichtian.Fossils: Dinosaur eggs.Latitude/longitude: 19o3602200S/47o58027.300W.References: Price (1951), Campos and Bertini (1985),Bertini et al. (1993), Kellner and Campos (2000), Magal-haes-Ribeiro (2001a), Magalhaes-Ribeiro (2001b),Magalhaes-Ribeiro (2002a).

Locality 10: Freeway BR-050/Km-24, Uberaba town.Horizon: Marılia Formation, Serra da Galga Mem-ber. Age: late Maastrichtian.Fossils: Pisces indet. (teeth, scales); Dinosauria: Thero-poda indet.Latitude/longitude: 19�3501600S/48�0104700W.References: Bertini et al. (1993), Kellner and Campos(2000).

Locality 11: ‘‘Posto Cinquentao” BR-050, Uberabatown. Horizon: Marılia Formation, Ponte Altaand Serra da Galga members. Age: lateMaastrichtian.Fossils: Vertebrata indet.; Sauropoda indet.; Theropodaindet.Latitude/longitude coordinates: 19�3702300S/47�5802600W.References: Bertini et al. (1993), Kellner and Campos(2000).

Locality 12: Quarry Partezan BR-050, Uberaba town.Horizon: Marılia Formation, Ponte Alta and Serra daGalga members. Age: late Maastrichtian.Fossils: Plantae: Charophyceae: Amblyochara sp., Cha-

ra? sp., Gobichara groeberi; Ostracoda: Candonopsis

sp., Ilyocypris setembrinopetrii, Ilyocypris sp. 1, Neu-

quenocypris minor mineira, Virgatocypris mezzalirai,gen. et sp. indet. 1; Vertebrata indet., Sauropoda indet.,Theropoda indet.Latitude/longitude coordinates: 19�3706000S/47�2700100W.References: Bertini et al. (1993), Kellner and Campos(2000), Dias-Brito et al. (2001).

Locality 13: Quarry Triangulo, Uberaba town. Horizon:Marılia Formation, Ponte Alta and Serra da Galgamembers. Age: late Maastrichtian.Fossils: Plantae: Charophyceae: Amblyochara sp., Cha-

ra? sp., Feistiella cf. globosa, Feistiella cf. costata, Gobic-

hara groberi, Nitellopsis? sp., Sphaerochara latifasciata;Ostracoda: Altanicypris australis, ‘‘Lycopterocypris

angualta”, Neuquenocypris minor mineira, Virgatocypris

mezzalirai, gen. et sp. indet. 2; Vertebrata indet., Sauro-poda indet., Theropoda indet.Latitude/longitude: 19�3704200S/47�1304100W.References: Bertini et al. (1993), Kellner and Campos(2000), Dias-Brito et al. (2001).

Locality 14: Freeway BR-050, Uberaba town. Horizon:Marılia Formation, Ponte Alta and Serra da Galgamembers. Age: late Maastrichtian.Fossils: Vertebrata indet.Latitude/longitude: 19�3701900S/47�0101300W.References: Albuquerque and Candeiro (2003).

4.2. Descriptions of the vertebrate species from thePeiropolis area

The institutional abbreviations used herein are as fol-lows: CPP, Centro de Pesquisas Paleontologicas LlewellynIvor Price, Peiropolis, Uberaba town, Minas Gerais State,Brazil; and DGM, Departamento de Geologia e Mineralo-gia, Rio de Janeiro, Rio de Janeiro State, Brazil.

4.2.1. Systematic paleontology

Anura Rafinesque, 1815.‘‘Leptodactylidae” Werner, 1896.Baurubatrachus pricei Baez and Peri, 1989Holotype: DGM 1412-R, A and B (Fig. 5a). Incomplete

skeleton, partially articulated, embedded in two blocks ofsandstone.

Locality: 2.Horizon and age: Serra da Galga Member, Marılia For-

mation, late Maastrichtian.Comments: Baurubatrachus pricei is the only known

anuran from the Late Cretaceous of the UA. B. pricei wasconsidered by Rocek and Lamaud (1995) to be representa-tive of the ‘‘Leptodactylidae” and by Baez and Peri (1989)and Sanchız (1998), as a possible ‘‘Ceratophryinae.” Lepto-dactylids are a large and varied group of frogs with extantspecies occurring in South and Central America, as well asthe West Indies. Extant leptodactylids are extremely rare inthe United States and absent from Africa and Eurasia.

The fossil record of this family is sparse, including onlyWawelia gerholdi Casamiquela, 1963 and Neoprocoela

edentata Schaeffer, 1949 recorded, respectively, in theUpper Miocene and Lower Oligocene of Argentina.

Lepidosauria Haeckel, 1866.Squamata Oppel, 1811.Iguania Cuvier, 1817.Pristiguana brasiliensis Estes and Price, 1973.Holotype: DGM 552. Portion of a disarticulated skele-

ton of a single individual. The preserved elements include


frontals, a left pterygoid with teeth, a right surangular, aleft pterygoid, splenial, a left humerus (Fig. 5b), and manyother skeletal elements.


mation, late Maastrichtian.Comments: The holotype specimen (DGM 552) of P.

brasiliensis was originally placed in Iguanidae (Estes andPrice, 1973). In Brazil, Mesozoic terrestrial fossil Eola-certilia records are limited to the latest Cretaceous (P.

brasiliensis) and Early Cretaceous (Olindalacerta brasili-

ensis and Tijubina pontei) from the Crato Formation ofthe Araripe Group, Araripe Basin, northeastern Brazil(Bonfim-Junior and Marques, 1997; Evans and Yabum-oto, 1998). In addition to being considered an indetermi-nate squamate by later researchers (e.g., Gao and Hou,1995), P. brasiliensis has not received any detailed sys-tematic revision since its original description. The recentfinding of possible iguanids in the Cenomanian of Argen-tina (Apesteguıa et al., 2004b) reopens the interest inPristiguana as a true iguanid.

Crocodyliformes Benton and Clark, 1988.Mesoeucrocodylia Whetstone and Whybrow, 1983.Peirosauridae Gasparini, 1982.Peirosaurus tormini Price, 1955.Holotype: DGM 433-R. The specimen is composed of a

left premaxilla (Fig. 5c) with five teeth, a left palpebral,maxillary and dentary teeth, cervical, dorsal, and one cau-dal vertebrae, ribs, chevrons, a radius and ulna; a pubis andan incomplete left ischium; and several dermal plates.


mation, late Maastrichtian.Comments: With the Upper Cretaceous Lomasuchus

palpebrosus Gasparini, Chiappe and Fernandez fromNeuquen Province, Argentina (Gasparini et al., 1991),Peirosaurus tormini are included in the Peirosauridae(Gasparini, 1982), a group diagnosed on the basis ofthe following synapomorphies: distance from anteriororbital edge to anterior contour of rostrum equal toor longer than distance from the anterior orbital edgeto posterior pariental contour; dentary extends caudallyup to end of tooth row; splenials behind symphysis isbroad and robust, and anterior projection of surangularis unique and acute in lateral view; no surangular andquadratojugal taking part in craniomandibular joint; fivepremaxilary teeth; and occipital condyle caudally direc-ted (Carvalho et al., 2004). Buckley and Brochu (1999)consider Mahajangasuchus closely related to Peirosauri-dae. Furthermore, sensu Carvalho et al. (2004), Maha-

jangasuchus is deeply included within the group,forming the ‘‘Mahajangasuchini” together with Uber-

abasuchus. According to Carvalho et al. (2004), thesetaxa share the following synapomorphies: premaxilla-maxilla suture zigzag-shaped in lateral view; absence ofa foramen at premaxilla-maxilla suture in lateral view;

premaxilla–maxilla notch present; no supraoccipitalexposure on cranial roof, parientals contact on occipitalavoiding dorsal exposure of supraoccipital; basisphenoidalmost hidden from ventral view and covered bypyterygoid and basioccipital in lateral view, ostodermswithout anterior projection; posterior branch of quad-rate at least as long as broad; and nasal and lacrimaldo not contact or with a short contact. Just outsidethe tribe ‘‘Mahajangasuchini” is the Argentinian taxonLomasuchus, a basal ‘‘Lomasuchinae” (sensu Carvalhoet al., 2004).

Uberabasuchus terrificus Carvalho et al., 2004.Holotype: CPP 630. Skull (Fig. 5d), mandible and part

of the axial and appendicular skeleton.Locality: 1.Horizon and age: Serra da Galga Member, Marılia For-

mation, late Maastrichtian.Comments: Uberabasuchus terrificus is a most com-

plete south Gondwanan mesoeucrododylian, as recentlydescribed by Carvalho et al. (2004). It bears moderate,narrow-snouted, serrate teeth; large orbits protected bysupraorbital bones of triangular shape; and an antor-bital fenestrae bounded posteriorly by a deep groove.U. terrificus forms with Mahajangasuchus the tribe‘‘Mahajangasuchini”. A recent proposal by Carvalhoet al. (2004) coins ‘‘Lomasuchinae” to include Lomasu-

chus palpebrosus (from the Bajo de la Carpa Forma-tion, Neuquen Group, Patagonia) and‘‘Mahajangasuchini”. They share the following synapo-morphies: lateral contour of dentary sigmoidal in dor-sal view, no separated nares, mandibular symphysisshort, and largest mandibular teeth double, generallythe third and the fourth.

Crocodyliformes Benton and Clark, 1988.Mesoeucrocodylia Whetstone and Whybrow, 1983.Itasuchus jesuinoi Price, 1955.Holotype: DGM 434-R. Incomplete left maxilla with

seven alveoli and the anterior part of the fourth alveolus,left jugal, quadratojugal and quadrate, a dentary, botharticulars, four cervical, four dorsal and four caudal verte-brae, one chevron, complete right and incomplete left cor-acoid, left humerus, both ulnae, left pelvis; both femora,tibiae and fibulae, four metapodials, and about 20 dermalplates.


mation, late Maastrichtian.Comments: Itasuchus (Fig. 5e) occurs in the same

Maastrichtian deposits as Peirosaurus tormini. Althoughconsidered a Trematochampsidae by some authors (e.g.,Buffetaut, 1985; Rasmusson, 2002), Itasuchus jesuinoi

remains a basal peirosauroid (the most recent commonancestor of Itasuchus and Peirosauridae and all itsdescendents), together with Malawisuchus mwakayunguti-

ensis, from Karonga District, Northern Malawi, Africa,


which together form the Itasuchidae (sensu Carvalhoet al., 2004).

5. Conclusions

The Uberaba area (UA) has yielded a wide diversityof taxa, especially charophytes, ostracods, and tetrapods(e.g., crocodyliforms, dinosaurs). This fauna represents amixture of terrestrial and aquatic animals, associatedwith fluviolacustrine environments. It includes at leastthree dinosaur taxa: abelisaurid, carcharodontosauridand maniraptoran theropods, titanosaurian sauropods,including Baurutitan britoi, Trigonosaurus pricei, Aeolo-

saurus sp., and indeterminate forms; the crocodyliformsPeirosaurus tormini, Uberabasuchus terrificus, and Itasu-

chus jesuinoi, the anuran Baurubatrachus pricei, the turleCambaremys langertoni, and the lizard Pristiguana

brasiliensis.The UA fauna is remarkably similar to other South

American Late Cretaceous faunas such as those fromSao Paulo, Brazil, and Argentina, and also share taxawith Late Cretaceous continental faunas from more dis-tant Gondwanan localities (e.g., Madagascar). Croco-dyliforms show the closer connection with Madagascar,based on the close relationships of the ‘‘Mahajangasu-chini” peirosaurids. On the other side, more basal taxa,such as basal peirosauriformes, show close relationshipswith continental Africa, in support of known continen-tal Gondwanan connections and in contrast to the con-clusions of Franco and Bertini (1997) and Franco(1999), who suggest a Laurasian origin for the UAfauna. Carcharodontosaurid remains from the UA andPatagonia demonstrate the survival of these theropodslate into the Cretaceous. The absence of those dinosaursoutside South America also suggests the existence ofbarriers.

Acknowledgments

We thank reviewers Dra. Saswati Bandyopadhyay (In-dian Statistical Institute, Calcutta), Dr. David Martill(School of Earth and Environmental Sciences, Universityof Portsmouth), Dr. Ana M. Baez (Departamento de Cien-cias Geologicas, Universidad de Buenos Aires), Dr. EricBuffetaut (Research at Centre National de la RechercheScientifique, Paris), Dr. Diego Pol (American Museum ofNatural History, New York), Dra. Cibele Schwanke (Insti-tuto de Educac�ao, Universidade Estadual do Rio de Ja-neiro), Dr. Paul Upchurch (University College London,London), and Dr. Paul Barret (The Natural History Mu-seum Dept of Palaeontology, London) for criticisms andsuggestions that greatly improved the manuscript. Themanuscript was substantially improved after the revisionwork of Dr. Mattew Lamanna (Carnegie Museum of Nat-ural History) and Darren Naish (University of Ports-mouth). We thank Dr. Dimas Dias-Brito (UniversidadeEstadual Paulista, Rio Claro) who suggested relevant liter-ature. We are particularly grateful to Claudia Regina A.Candeiro (Uberlandia, Minas Gerais State) and FelipeM. Costa (Laboratorio de Cartografia e SensoriamentoRemoto, Instituto de Geografia, Universidade Federal deUberlandia) for preparation of the some illustrations. Van-derlon S., (in memorium), Joao I., and Carlos M. (Centrode Pesquisas Paleontologicas Llewellyn Ivor Price) wereamong those who excavated specimens from the field.Thanks are extended to the geologist Prof. Dr. IgnacioMachado Brito (in memorium), Universidade Federal deUberlandia, for advice during the undergraduate programof R. Candeiro. This work was part of a revised undergrad-uate project supported by grants by CNPq/166/98, 178/99to R. Candeiro and FAPEMIG/UFU/801/98 to InacioBrito (for Visiting Professor research at the Departamentode Geografia, Universidade Federal de Uberlandia).

Appendix A. Late Cretaceous fauna and flora from the Uberaba area

Taxon Locality Provenance Selected references

PLANTAECharophyceae

Amblyochara sp. 1
2, 13 Ponte Alta Mb., Marılia Fm. Dias-Brito et al. (2001) Chara tomentosa 1 3 Ponte Alta Mb., Marılia Fm. Dias-Brito et al. (2001) Chara sp. 1 2, 13 Ponte Alta Mb., Marılia Fm. Dias-Brito et al. (2001) Feistiella cf. Costata 1 3 Ponte Alta Mb., Marılia Fm. Dias-Brito et al. (2001) Feistiella cf. globosa 1 3 Ponte Alta Mb., Marılia Fm. Dias-Brito et al. (2001) Gobichara (Pseudoharrisichara)groeberi
1
3 Ponte Alta Mb., Marılia Fm. Dias-Brito et al. (2001)
Gobichara (Pseudoharrisichara) sp. 1
2 Ponte Alta Mb., Marılia Fm. Dias-Brito et al. (2001) Nitellopsis? sp. 1 3 Ponte Alta Mb., Marılia Fm. Dias-Brito et al. (2001) Sphaerochara latifasciata 1 3 Ponte Alta Mb., Marılia Fm. Dias-Brito et al. (2001)
Pteridophyta
Marsilea sp.? 1 Serra da Galga Mb., Marılia Fm. Senra and Silva-Silva (1998)


Appendix A (continued)

Taxon
Locality P rovenance S elected references
‘‘INVERTEBRATES”

GastropodsOstracoda

Altanicupris australis
13 P onte Alta Mb., Marılia Fm. D ias-Brito et al. (2001) Candonopsis sp. 12 P onte Alta Mb., Marılia Fm. D ias-Brito et al. (2001) Ilyocypris setembrinopetrii 12 P onte Alta Mb., Marılia Fm. D ias-Brito et al. (2001) Ilyocpris sp. 1 12 P onte Alta Mb., Marılia Fm. D ias-Brito et al. (2001) Lycopterocypris angulata 13 P onte Alta Mb., Marılia Fm. D ias-Brito et al. (2001) Neuquenocypris minor mineira 12, 13 P onte Alta Mb., Marılia Fm. D ias-Brito et al. (2001) Virgatocypris mezzalirai 12, 13 P onte Alta Mb., Marılia Fm. D ias-Brito et al. (2001) gen. et sp. indet. 1 12 P onte Alta Mb., Marılia Fm. D ias-Brito et al. (2001) gen. et sp. indet. 2 13 P onte Alta Mb., Marılia Fm. D ias-Brito et al. (2001)
Gastropoda
Physidae Physa aridi 1 S erra da Galga Mb., Marılia Fm. S enra (2002) Viviparidadae Viviparus souzai 1 S erra da Galga Mb., Marılia Fm. S enra (2002)
Bivalvia
Florencia peiropolensis 1 S erra da Galga Mb., Marılia Fm. M ezzalira (1974) Anodontites (Anodontites) pricei 1 S erra da Galga Mb., Marılia Fm. M ezzalira (1974) Musculium sp. 1 S erra da Galga Mb., Marılia Fm. S enra (2002)
VERTEBRATA
Vertebrata indet. 11, 12,
13, 14PM

onte Alta and Serra da Galga mbs.,arılia Fm.

C
andeiro and Bergqvist (2004)
Pisces
Characiformes indet. 1 S erra da Galga Mb., Marılia Fm. G ayet and Brito (1989) Perciformes indet. 1 S erra da Galga Mb., Marılia Fm. G ayet and Brito (1989) Siluriformes 1 S erra da Galga Mb., Marılia Fm. G ayet and Brito (1989) Pisces indet. 3 P onte Alta Mb., Marılia Fm. F ernandes (1998)
Anura
Baurubatrachus pricei 7 S erra da Galga Mb., Marılia Fm. B aez and Peri (1989)
Chelonia
Cambaremys langertoni 2 S erra da Galga Mb., Marılia Fm. F ranc�a and Langer (2005) gen. et sp. indet. 1 S erra da Galga Mb., Marılia Fm. B ertini et al. (1993)
Lepidosauria
Pristiguana brasiliensis 1 S erra da Galga Mb., Marılia Fm. E stes and Price (1973)
Mesoeucrocodylia
Itasuchus jesuinoi 1 S erra da Galga Mb., Marılia Fm. P rice (1955) Peirosaurus tormini 1 S erra da Galga Mb., Marılia Fm. P rice (1955) Uberabasuchus terrificus 1 S erra da Galga Mb., Marılia Fm. C arvalho et al. (2004)
Dinosauria
Dinosauria indet. 4, 7, 8, 9 S erra da Galga Uberaba Fm., Mb.,
Marılia Fm.C
andeiro et al. (2004b)
Sauropoda
‘‘Megaloolithidae” indet. 1 U beraba Fm., Serra da Galga Mb.,
Marılia Fm.R
ibeiro (2002a, b)
(continued on next page)


Appendix A (continued)

Taxon L
ocality Provenance Selected references
gen. et sp. nov. 7
Uberaba Fm., Ponte Alta and Serrada Galga mbs, Marılia Fm.
Santucci (2002), Candeiro et al.(2004b)

Titanosauria
Baurutitan britoi 1 Serra da Galga Mb., Marılia Fm. Kellner et al. (2005) Trigonosaurus pricei 1 Serra da Galga Mb., Marılia Fm. Campos et al. (2000) Aeolosaurus sp. 1 Serra da Galga Mb., Marılia Fm. Bertini et al. (2000), Santucci
(2002)
gen. et sp. indet. 1 , 2, 5, 6,
7
Uberaba Fm., Serra da Galga Mb.,Marılia Fm.
Powell (1986, 1987, 2003),Santucci and Bertini (2006)

Theropoda
Abelisauridae indet. 1 ,2 Serra da Galga Mb., Marılia Fm. Candeiro (2002), Candeiro
et al. (2004a)
Carcharodontosauridae indet. 1 ,2 Serra da Galga Mb., Marılia Fm. Candeiro (2002), Candeiro
et al. (2004a)
Maniraptora indet. 1 Serra da Galga Mb., Marılia Fm. Novas et al. (2005) gen. et sp. indet. 1 ,2, 10,
11, 12, 13
Serra da Galga Mb., Marılia Fm. Candeiro (2002)
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Fauna Flora Cretacea Uberaba