Journal of South American Earth Sciences 25 (2008) 203216 www.elsevier.

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The Late Cretaceous fauna and ora of the Uberaba area (Minas Gerais State, Brazil)
lian P. Bergqvist b, Carlos Roberto A. Candeiro a,b,*, Adriano R. Santos a, L c,d n Apestegu a e Luiz Carlos B. Ribeiro , Sebastia
villa, 2160, ndia, Av. Joa Museu de Minerais e Rochas, Instituto de Geograa, Universidade Federal de Uberla o Naves de A nica, Uberla ndia, Minas Gerais, Brazil Sta. Mo b rio de Macrofo sseis, Departamento de Geologia, Universidade Federal do Rio de Janeiro, Av. Brig. Trompski, Laborato ria, Rio de Janeiro, RJ, Brazil S/N, Cidade Universita c Fundac a o Municipal de Ensino Superior de Uberaba-FUMESU/Centro de Ensino Superior de Uberaba-CESUBE/ gicas Llewellyn Ivor Price, BR-262, KM 784, Peiro polis, Uberaba, Minas Gerais, Brazil Centro de Pesquisas Paleontolo d Sabino, 1801, Universita rio, Universidade de Uberaba-UNIUBE/Instituto de Formac a o de Educadores, Dep. De Biologia, Av. Nene 38055-500, Uberaba, Minas Gerais, Brazil e nico de Ciencias Naturales, Av. Roca 1250, General Roca, R o Negro, Argentina Museo Patago
a

Abstract The Uberaba area, in Minas Gerais State, Brazil, yields a rich continental fauna and ora from the Late Cretaceous Uberaba and lia formations. This paper reviews the diversity of the biota recorded from these formations. The most signicant taxa from PeiMar polis are the frog Baurubatrachus pricei, the turtle Cambaremys langertoni, the lizard Pristiguana brasiliensis, the crocodyliforms Itasuro chus jesuinoi, Peirosaurus tormini and Uberabasuchus terricus, the titanosaurian Baurutitan britoi, Trigonosaurus pricei, Aeolosaurus sp., indeterminate titanosaurians, and abelisaurid, carcharodontosaurid and maniraptoran theropods. Together with faunas of a similar age in 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 ora; Brazil; Gondwana

Resumo ceo), Estado de Minas Gerais, Brasil prove m das formac lia sendo A fauna e ora da regia o o de Uberaba (Neocreta es Uberaba e Mar representada de taxa de origem continental. Este artigo revisa a diversidade da biota registrada nestas formac o es. Os taxa mais signi polis sa cantes de Peiro o conhecidos pelo sapo, Baurubatrachus pricei, a tartaruga Cambaremys langertoni, o lagarto, Pristiguana brasiliensis, e pelos crocodilianos Itasuchus jesuinoi, Peirosaurus tormini e Uberabasuchus terricus. Os dinossauros sa o representados por deos, abelisaur deos, carcarodontosaur deos e maniraptor. restos de Baurutitan britoi, Trigonosaurus pricei, Aeolosaurus sp., titanossaur ia fossil fera proveniente de Uberaba e de origem gondwa ceo nica e apresentando anidades com a fauna e ora do Neocreta A assemble da Argentina e Madagascar. 2007 Elsevier Ltd. All rights reserved.
ceo Superior; Uberaba; Fauna e ora; Brasil; Gondwana Palavras chave: Creta

villa, ndia, Av. Joa Corresponding author. Address: Museu de Minerais e Rochas, Instituto de Geograa, Universidade Federal de Uberla o Naves de A nica, Uberla ndia, Minas Gerais, Brazil. Fax: +51 34 3214 2925. 2160, Sta. Mo E-mail addresses: candeiro@yahoo.com.br, candeiro@ras.ufu.br (C.R.A. Candeiro).
*

0895-9811/$ - see front matter 2007 Elsevier Ltd. All rights reserved. doi:10.1016/j.jsames.2007.06.005

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1. Introduction The Uberaba area (UA) in southwestern Minas Gerais State, Brazil, has yielded important Late Cretaceous fauna and ora (Fig. 1). The paleontological importance of this site has been recognized since the beginning of the 1940s (Price, 1961), but a comprehensive, systematic review of the rich UA Bauru Group fossil biota has yet to be performed. In the past 15 years, a large number of beautifully preserved fossil invertebrates and vertebrates (Appendix A) have been recovered from the ConiacianSantonian Uberaba Formation (Dias-Brito et al., 2001) and late Maaslia Formation (Dias-Brito et al., 2001) at trichtian Mar Uberaba. Despite these advances, current understanding of the Upper Cretaceous biota of the site remains far from complete. We review the up-to-date knowledge of the fauna of the site and compare it with faunas of other contemporaneous Gondwanan localities, notably those coming from nearby Patagonia (Argentina) and Madagascar. 2. Geological setting The Uberaba area yields fossils from two of the three lia; formations of the Bauru Group: Uberaba and Mar the Adamantina Formation is absent (Fig. 2). The fossil ngulo Mineiro region, bearing units present in the Tria which includes the Uberaba area, and crop out in the northeastern region of the Bauru Basin (Fernandes and Coimbra, 1996). They were deposited above the basalts of the Serra Geral Formation (Sa o Bento Group, Parana Basin), the sandstones of the Botucatu Formation (Sa o Bento Group), the metamorphic Proterozoic outcrops of and Canastra groups (Sanfranciscana Basin), the Araxa and the Mesozoic intrusive event of the Soerguimento do ba (Suguio et al., 1979). Alto Parana The older unit of the site is the ConiacianSantonian (Dias-Brito et al., 2001), 140 m thick, Uberaba Formation (Fernandes and Coimbra, 1996), which is mainly exposed at Uberaba in Minas Gerais State. It is composed of limestones, sandstones, and a carbonate-cemented basal conglomerate and includes an important volcanoclastic unit (Barcelos, 1984). Among the fossils collected are microfossils, turtles, titanosaurian sauropods, indeterminate dinosaur remains, and ichnofossils. The younger unit, the lia Formation (Fernandes and Coimbra, 180 m thick Mar s and Sa 1996), is exposed in Goia o Paulo states and in ngulo Mineiro region. It consists of ne- to medthe Tria ium-grained sandstones intercalated with conglomerates. The sandstones are well cemented and bear calcium car lfaro and Barcelos, 1991). Some bonate concretions (Fu lia authors, such as Bertini et al. (1993), suggest the Mar Formation is CampanianMaastrichitan, based on vertebrate fossils. Recently, Dias-Brito et al. (2001) have prolia posed an Upper Maastrichtian age for the Mar Formation, based on ostracods and charophytes. lia Formation has an irregular outcrop pattern The Mar ngulo Mineiro, as is the case in western Sa within the Tria o

Paulo state. According to Suguio (1973, 1980) and Barcelos and Suguio (1987), it was deposited as a series of coalescent alluvial fans, previously reworked and redeposited by a braided uvial system, associated with calcretes and calcareous lacustrine environments with intermittent dry lia Formation periods. Barcelos (1984) subdivides the Mar into three members, Echapora , Ponte Alta, and Serra da Galga, the latter two restricted to the Uberaba area. The basal Ponte Alta Member (Fig. 3) was characterized sedimentologically by Garrido et al. (1992) as a calcareous white member composed of thin and medium limestones with pebbles and calcareous nodules deposited in playa lakes, with alkaline water supersaturated with calcium bicarbonates. The upper Serra da Galga Member (Fig. 3) is characterized as a member of white limestone and conglomerate and composed of conglomerate and thinto medium-grained limestones, with bleached coloration in an arkosic matrix. It shows an abundance of cross-stratication and invertebrate bioturbation (Suguio, 1973, 1980). lia Formation includes several Cretaceous The Mar continental 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 abelisaurid, 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 abundant in the regions of Monte Alegre de Minas and Uberaba ndia highway, as well as at along the UberabaUberla Campina Verde in Minas Gerais State (von Huene, 1931; Campos and Bertini, 1985; Kellner, 1996; Kellner and Campos, 2000; Santucci and Bertini, 2002, 2006; Candeiro et al., 2004b). The record includes Aeolosaurus sp. (from the UA) and indeterminate forms (from the UA and Monte Alto in Sa o Paulo State). 3. Age of fossil-bearing sediments in the Uberaba area The Upper Cretaceous age of the Bauru Group was emphasized by Bertini et al. (1993) and Santucci and Bertini (2002) on the basis of similarities between the UA titanosaurian and Aeolosaurus sp., among others. In addition, the microfossil assemblage reported by Dias-Brito et al. (2001) favors a late Maastrichtian age lia Formation and a ConiacianSantonian for the Mar age for the Uberaba Formation. The Upper Cretaceous lia Formation, as ostracod fauna reported from Mar exposed in the UA, includes Ilyocypris setembrinopetrii, Ilyocpris sp. 1, Neuquenocypris minor mineira, Virgatocypris mezzalirai, Altanicypris australis, Lycopterocypris angulata, Candonopsis sp., and gen. et sp. Indet. 1 and 2. Of these, Neuquenocypris minor and Ilyocypris particu n Basin, larly link this fauna with that of the Neuque Argentina (Musacchio and Simeoni, 1991; Dias-Brito et al., 2001).

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

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

4. Remarks on the biota of the Uberaba area Few invertebrates are known from UA. Pulmonate gaslia Formation, and as tropods occur mainly in the Mar mentioned previously, ostracods are well represented and very diverse in the UA fauna (Appendix A). The charophyte 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 a broad distribution in the Bauru Basin (Dias-Brito et al., 2001). There have been a signicant number of new fossil discoveries in the UA in the past 15 years (Appendix A; Figs. 4ae, 5ae). Among the tetrapods, a single leptodactylid species, Baurubatrachus pricei Baez and Peri, 1989 (Fig. 5a), represents the only anuran from Uberaba, whereas crocodyliforms are represented by large mesoeucrocodylian

taxa, including Peirosaurus tormini and Itasuchus jesuinoi Price, 1955, and Uberabasuchus terricus Carvalho et al., 2004 (Figs. 5ce). Chelonians are represented by Cambaremys langertoni Franc a and Langer, 2005 and turtle remains (Fig. 4d) are well known from the Uberaba region, but when diversity and abundance are taken into account, dinosaurs form 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 titanosaurians and abelisaurid and carcharodontosaurid theropods occur frequently in the UA (Fig. 4a, b, and e). Reported UA titanosaurs include Baurutitan britoi Kellner et al., 2005, Trigonosaurus pricei Campos et al., 2005, and Aeolosaurus sp. and indeterminate titanosaurian species, as well as eggshell fragments referred to Megaloolithidae (Fig. 4c) (Magalha esRibeiro, 2002a). Among the theropods, the presence of abelisaurid and carcharodontosaurid teeth is noteworthy (e.g., Candeiro, 2002; Marinho et al., 2003; Candeiro

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

et al., 2004a), as well a rst maniraptoran claw described recently by Novas et al. (2005). Some teeth were previously mistaken as those of dromaeosaurids and troodontids (Franco and Bertini, 1997; Franco, 1999), but most actually belong to carcharodontosaurids. The former family, however, is present in the Cretaceous of South America (Candeiro, 2002). The Carcharodontosauridae are characterized by their enamel wrinkles (Candeiro, 2002; Candeiro et al., 2004a) and apparently survived well into the Late Cretaa et al., ceous (Martinelli and Forasepi, 2004; Apestegu a, 2002). With regard to sauropods, 2004a; cf. Apestegu Santucci and Bertini (2002) and Candeiro et al. (2006a,b) demonstrate the presence of the titanosaurian Aeolosaurus lia Formation), approximately coeval sp. at Uberaba (Mar with Aeolosaurus species from Argentina. In addition, Santucci (2002) reports a new titanosaurian species from both lia formations. the Uberaba and Mar 4.1. Fossiliferous localities within the UA Fourteen Upper Cretaceous fossiliferous localities within the Uberaba region are considered in this study (Fig. 1, Appendix A). The localities in chronological order are as follows:

polis district, Locality 1: Ponto 1 do Price, Peiro lia Formation, Serra da Uberaba town. Horizon: Mar Galga Member. Age: late Maastrichtian. Fossils: Plantae: Pyteridophyta: Marsilea sp.?; invertebrates: Mollusca; Bivalvia; Musculium sp., Gastropoda; Physidae; Pysa aridi, Vivirapidae; Anodontites (Anodontites) pricei, Florencia peiropolensis, Viviparus souzai; Vertebrata: Pisces; Characiformes indet., Perciformes indet., Siluriformes indet.; Chelonia: gen. et sp. Indet., Podocnemidae indet.; Lepidosauria: Pristiguana brasiliensis; Mesoeucrodylia: Itasuchus jesuinoi, Peirosaurus tormini, Uberabasuchus terricus; Dinosauria: Sauropoda: Megaloolithidae indet., Baurutitan britoi, Trigonosaurus pricei Titanosauria (sensu Wilson and Upchurch, 2003); Theropoda: Abelisauridae indet., Carcharodontosauridae indet, Maniraptora indet. Latitude/longitude coordinates: 19430 13.200 S/ 0 00 4744 17.9 W. References: Price (1951, 1955, 1961), Estes and Price (1973), Mezzalira (1974), Bonaparte (1978, 1996, 1999), Buetaut (1982, 1985), Gayet and Brito (1989), Gasparini et al. (1991), Bertini et al. (1993), Kellner (1995), Albino (1996), Gasparini (1996), Salgado et al. (1997), Candeiro (1998, 2002), Fernandes

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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; ce = 20 mm; f = 150 mm.

(1998), Magalha es-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 and Bertini (2002), Evans et al. (2002), Powell (2003), Ribeiro (2003), Franc a and Langer (2003), Marinho and 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), Campos et al. (2005), Kellner et al. (2005), Novas et al. (2005). polis district, UberLocality 2: Ponto 2 do Price, Peiro lia Formation, Serra da Galga aba town. Horizon: Mar Member. Age: late Maastrichtian. Fossils: Chelonia: Cambaremys langertoni; Dinosauria: Saurpoda: Titanosauria indet.; Theropoda: Abelisauridae indet., Carcharodontosauridae indet. Latitude/longitude coordinates: 19430 12.000 S/ 0 00 4745 04.4 W.

References: Price (1961), Bertini et al. (1993), Fernandes z (1998), Franco (1999), Candeiro (2002), (1998), Sanch Santucci and Bertini (2002), Marinho and Candeiro (2003b), Venczel and Csiki (2003), Candeiro and Bergqvist (2004), Candeiro et al. (2004a,b), Franc a and Langer (2005). polis Quarry, Peiro polis district, UberLocality 3: Peiro lia Formation, Ponte Alta aba town. Horizon: Mar Member. Age: late Maastrichtian. Fossils: Pisces indet. Latitude/longitude coordinates: 19o420 52.400 S/ o 0 00 47 45 16.1 W. References: Candeiro (1998), Fernandes (1998), Candeiro and Bergqvist (2004). polis district, Locality 4: Ponto 1 Langerton, Peiro Uberaba town. Horizon: Uberaba Formation. Age: Coniacian-Santonian. Fossils: Dinosauria indet. (nests). Latitude/longitude: 19440 42.500 S/47440 32.200 W. References: Magalha es-Ribeiro (2002a,b).

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

Locality 5: Mumbuco Freeway BR-262/KM-780, lia Formation, Serra da Uberaba town. Horizon: Mar Galga Member. Age: late Maastrichtian. Fossils: Dinosauria: Titanosauria indet. Latitude/longitude: 19430 29.300 S/47420 55.200 W. References: Campos and Kellner (1999), Kellner and Campos (2000), Santucci and Bertini (2002), Trotta et al. (2002), Candeiro et al. (2004b).

Fossils: Dinosauria: gen. et sp. indet. (eggs). Latitude/longitude: 19420 38.300 S/47400 27.900 W. References: Magalha es-Ribeiro (2002a), Santucci (2002), Candeiro et al. (2004b). Locality 7: Manoel Mendes Neighborhood/Avenue Leopoldina de Oliveira, Uberaba town. Horizon: Uberaba Formation. Age: Coniacian-Santonian. Fossils: Anura: Baurubatrachus pricei; Dinosauria: gen. et sp. indet.; Sauropoda indet. Latitude/longitude: 19450 29.0000 S/475300 53.500 W. References: Baez (1985), Baez and Peri (1989), Santucci (2002).

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

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nior Outcrop Locality 8: Avenue Randolfo Borges Ju near the Uberaba River, Uberaba town. Horizon: Uberaba Formation. Age: Coniacian-Santonian. Fossils: Dinosauria indet. Latitude/longitude: 19430 16.500 S/47570 3000 W. References: Santucci (2002), Kellner and Campos (2000). Locality 9: Mangabeira Station, Uberaba town. Horilia Formation, Serra da Galga Member. zon: Mar Age: late Maastrichtian. Fossils: Dinosaur eggs. Latitude/longitude: 19o360 2200 S/47o580 27.300 W. References: Price (1951), Campos and Bertini (1985), Bertini et al. (1993), Kellner and Campos (2000), Magalha es-Ribeiro (2001a), Magalha es-Ribeiro (2001b), Magalha es-Ribeiro (2002a). Locality 10: Freeway BR-050/Km-24, Uberaba town. lia Formation, Serra da Galga MemHorizon: Mar ber. Age: late Maastrichtian. Fossils: Pisces indet. (teeth, scales); Dinosauria: Theropoda indet. Latitude/longitude: 19350 1600 S/48010 4700 W. References: Bertini et al. (1993), Kellner and Campos (2000). Locality 11: Posto Cinquenta o BR-050, Uberaba lia Formation, Ponte Alta town. Horizon: Mar and Serra da Galga members. Age: late Maastrichtian. Fossils: Vertebrata indet.; Sauropoda indet.; Theropoda indet. Latitude/longitude coordinates: 19370 2300 S/47580 2600 W. References: Bertini et al. (1993), Kellner and Campos (2000). Locality 12: Quarry Partezan BR-050, Uberaba town. lia Formation, Ponte Alta and Serra da Horizon: Mar Galga members. Age: late Maastrichtian. Fossils: Plantae: Charophyceae: Amblyochara sp., Chara? sp., Gobichara groeberi; Ostracoda: Candonopsis sp., Ilyocypris setembrinopetrii, Ilyocypris sp. 1, Neuquenocypris minor mineira, Virgatocypris mezzalirai, gen. et sp. indet. 1; Vertebrata indet., Sauropoda indet., Theropoda indet. Latitude/longitude coordinates: 19370 6000 S/47270 0100 W. References: Bertini et al. (1993), Kellner and Campos (2000), Dias-Brito et al. (2001). ngulo, Uberaba town. Horizon: Locality 13: Quarry Tria lia Formation, Ponte Alta and Serra da Galga Mar members. Age: late Maastrichtian. Fossils: Plantae: Charophyceae: Amblyochara sp., Chara? 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., Sauropoda indet., Theropoda indet. Latitude/longitude: 19370 4200 S/47130 4100 W. References: Bertini et al. (1993), Kellner and Campos (2000), Dias-Brito et al. (2001). Locality 14: Freeway BR-050, Uberaba town. Horizon: lia Formation, Ponte Alta and Serra da Galga Mar members. Age: late Maastrichtian. Fossils: Vertebrata indet. Latitude/longitude: 19370 1900 S/47010 1300 W. References: Albuquerque and Candeiro (2003).

4.2. Descriptions of the vertebrate species from the polis area Peiro The institutional abbreviations used herein are as fol gicas Llewellyn lows: CPP, Centro de Pesquisas Paleontolo polis, Uberaba town, Minas Gerais State, Ivor Price, Peiro Brazil; and DGM, Departamento de Geologia e Mineralogia, Rio de Janeiro, Rio de Janeiro State, Brazil. 4.2.1. Systematic paleontology Anura Ranesque, 1815. Leptodactylidae Werner, 1896. Baurubatrachus pricei Baez and Peri, 1989 Holotype: DGM 1412-R, A and B (Fig. 5a). Incomplete skeleton, partially articulated, embedded in two blocks of sandstone. Locality: 2. lia ForHorizon and age: Serra da Galga Member, Mar mation, late Maastrichtian. Comments: Baurubatrachus pricei is the only known anuran from the Late Cretaceous of the UA. B. pricei was ` k and Lamaud (1995) to be representaconsidered by Roce tive of the Leptodactylidae and by Baez and Peri (1989) z (1998), as a possible Ceratophryinae. Leptoand Sanch dactylids are a large and varied group of frogs with extant species occurring in South and Central America, as well as the West Indies. Extant leptodactylids are extremely rare in the United States and absent from Africa and Eurasia. The fossil record of this family is sparse, including only Wawelia gerholdi Casamiquela, 1963 and Neoprocoela edentata Schaeer, 1949 recorded, respectively, in the Upper 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 skeleton of a single individual. The preserved elements include

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frontals, a left pterygoid with teeth, a right surangular, a left pterygoid, splenial, a left humerus (Fig. 5b), and many other skeletal elements. Locality: 1. lia ForHorizon and age: Serra da Galga Member, Mar mation, late Maastrichtian. Comments: The holotype specimen (DGM 552) of P. brasiliensis was originally placed in Iguanidae (Estes and Price, 1973). In Brazil, Mesozoic terrestrial fossil Eolacertilia records are limited to the latest Cretaceous (P. brasiliensis) and Early Cretaceous (Olindalacerta brasiliensis and Tijubina pontei) from the Crato Formation of the Araripe Group, Araripe Basin, northeastern Brazil nior and Marques, 1997; Evans and Yabum(Bonm-Ju oto, 1998). In addition to being considered an indeterminate squamate by later researchers (e.g., Gao and Hou, 1995), P. brasiliensis has not received any detailed systematic revision since its original description. The recent nding of possible iguanids in the Cenomanian of Argena et al., 2004b) reopens the interest in tina (Apestegu Pristiguana 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 ve teeth, a left palpebral, maxillary and dentary teeth, cervical, dorsal, and one caudal vertebrae, ribs, chevrons, a radius and ulna; a pubis and an incomplete left ischium; and several dermal plates. Locality: 1. lia ForHorizon and age: Serra da Galga Member, Mar mation, late Maastrichtian. Comments: With the Upper Cretaceous Lomasuchus ndez from palpebrosus Gasparini, Chiappe and Ferna n Province, Argentina (Gasparini et al., 1991), Neuque Peirosaurus tormini are included in the Peirosauridae (Gasparini, 1982), a group diagnosed on the basis of the following synapomorphies: distance from anterior orbital edge to anterior contour of rostrum equal to or longer than distance from the anterior orbital edge to posterior pariental contour; dentary extends caudally up to end of tooth row; splenials behind symphysis is broad and robust, and anterior projection of surangular is unique and acute in lateral view; no surangular and quadratojugal taking part in craniomandibular joint; ve premaxilary teeth; and occipital condyle caudally directed (Carvalho et al., 2004). Buckley and Brochu (1999) consider Mahajangasuchus closely related to Peirosauridae. Furthermore, sensu Carvalho et al. (2004), Mahajangasuchus is deeply included within the group, forming the Mahajangasuchini together with Uberabasuchus. According to Carvalho et al. (2004), these taxa share the following synapomorphies: premaxillamaxilla suture zigzag-shaped in lateral view; absence of a foramen at premaxilla-maxilla suture in lateral view;

premaxillamaxilla notch present; no supraoccipital exposure on cranial roof, parientals contact on occipital avoiding dorsal exposure of supraoccipital; basisphenoid almost hidden from ventral view and covered by pyterygoid and basioccipital in lateral view, ostoderms without anterior projection; posterior branch of quadrate at least as long as broad; and nasal and lacrimal do not contact or with a short contact. Just outside the tribe Mahajangasuchini is the Argentinian taxon Lomasuchus, a basal Lomasuchinae (sensu Carvalho et al., 2004). Uberabasuchus terricus Carvalho et al., 2004. Holotype: CPP 630. Skull (Fig. 5d), mandible and part of the axial and appendicular skeleton. Locality: 1. lia ForHorizon and age: Serra da Galga Member, Mar mation, late Maastrichtian. Comments: Uberabasuchus terricus is a most complete south Gondwanan mesoeucrododylian, as recently described by Carvalho et al. (2004). It bears moderate, narrow-snouted, serrate teeth; large orbits protected by supraorbital bones of triangular shape; and an antorbital fenestrae bounded posteriorly by a deep groove. U. terricus forms with Mahajangasuchus the tribe Mahajangasuchini. A recent proposal by Carvalho et al. (2004) coins Lomasuchinae to include Lomasuchus palpebrosus (from the Bajo de la Carpa Forman tion, Neuque Group, Patagonia) and Mahajangasuchini. They share the following synapomorphies: lateral contour of dentary sigmoidal in dorsal view, no separated nares, mandibular symphysis short, and largest mandibular teeth double, generally the 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, both articulars, four cervical, four dorsal and four caudal vertebrae, one chevron, complete right and incomplete left coracoid, left humerus, both ulnae, left pelvis; both femora, tibiae and bulae, four metapodials, and about 20 dermal plates. Locality: 1. lia ForHorizon and age: Serra da Galga Member, Mar mation, late Maastrichtian. Comments: Itasuchus (Fig. 5e) occurs in the same Maastrichtian deposits as Peirosaurus tormini. Although considered a Trematochampsidae by some authors (e.g., Buetaut, 1985; Rasmusson, 2002), Itasuchus jesuinoi remains a basal peirosauroid (the most recent common ancestor of Itasuchus and Peirosauridae and all its descendents), together with Malawisuchus mwakayungutiensis, from Karonga District, Northern Malawi, Africa,

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which together form the Itasuchidae (sensu Carvalho et al., 2004). 5. Conclusions The Uberaba area (UA) has yielded a wide diversity of taxa, especially charophytes, ostracods, and tetrapods (e.g., crocodyliforms, dinosaurs). This fauna represents a mixture of terrestrial and aquatic animals, associated with uviolacustrine environments. It includes at least three dinosaur taxa: abelisaurid, carcharodontosaurid and maniraptoran theropods, titanosaurian sauropods, including Baurutitan britoi, Trigonosaurus pricei, Aeolosaurus sp., and indeterminate forms; the crocodyliforms Peirosaurus tormini, Uberabasuchus terricus, and Itasuchus jesuinoi, the anuran Baurubatrachus pricei, the turle Cambaremys langertoni, and the lizard Pristiguana brasiliensis. The UA fauna is remarkably similar to other South American Late Cretaceous faunas such as those from Sa o Paulo, Brazil, and Argentina, and also share taxa with Late Cretaceous continental faunas from more distant Gondwanan localities (e.g., Madagascar). Crocodyliforms show the closer connection with Madagascar, based on the close relationships of the Mahajangasuchini peirosaurids. On the other side, more basal taxa, such as basal peirosauriformes, show close relationships with continental Africa, in support of known continental Gondwanan connections and in contrast to the conclusions of Franco and Bertini (1997) and Franco (1999), who suggest a Laurasian origin for the UA fauna. Carcharodontosaurid remains from the UA and Patagonia demonstrate the survival of these theropods late into the Cretaceous. The absence of those dinosaurs outside South America also suggests the existence of barriers.

Acknowledgments We thank reviewers Dra. Saswati Bandyopadhyay (Indian Statistical Institute, Calcutta), Dr. David Martill (School of Earth and Environmental Sciences, University ez (Departamento de Cienof Portsmouth), Dr. Ana M. Ba cias Geologicas, Universidad de Buenos Aires), Dr. Eric Buetaut (Research at Centre National de la Recherche Scientique, Paris), Dr. Diego Pol (American Museum of Natural History, New York), Dra. Cibele Schwanke (Instituto de Educac a o, Universidade Estadual do Rio de Janeiro), Dr. Paul Upchurch (University College London, London), and Dr. Paul Barret (The Natural History Museum Dept of Palaeontology, London) for criticisms and suggestions that greatly improved the manuscript. The manuscript was substantially improved after the revision work of Dr. Mattew Lamanna (Carnegie Museum of Natural History) and Darren Naish (University of Portsmouth). We thank Dr. Dimas Dias-Brito (Universidade Estadual Paulista, Rio Claro) who suggested relevant liter udia Regina A. ature. We are particularly grateful to Cla ndia, Minas Gerais State) and Felipe Candeiro (Uberla rio de Cartograa e Sensoriamento M. Costa (Laborato Remoto, Instituto de Geograa, Universidade Federal de ndia) for preparation of the some illustrations. VanUberla derlon S., (in memorium), Joa o I., and Carlos M. (Centro logicas Llewellyn Ivor Price) were de Pesquisas Paleonto among those who excavated specimens from the eld. cio Thanks are extended to the geologist Prof. Dr. Igna Machado Brito (in memorium), Universidade Federal de ndia, for advice during the undergraduate program Uberla of R. Candeiro. This work was part of a revised undergraduate project supported by grants by CNPq/166/98, 178/99 cio to R. Candeiro and FAPEMIG/UFU/801/98 to Ina Brito (for Visiting Professor research at the Departamento ndia). de Geograa, Universidade Federal de Uberla

Appendix A. Late Cretaceous fauna and ora from the Uberaba area Taxon PLANTAE Charophyceae Amblyochara sp. Chara tomentosa Chara sp. Feistiella cf. Costata Feistiella cf. globosa Gobichara (Pseudoharrisichara) groeberi Gobichara (Pseudoharrisichara) sp. Nitellopsis? sp. Sphaerochara latifasciata Pteridophyta Marsilea sp.? Locality Provenance Selected references

12, 13 13 12, 13 13 13 13 12 13 13 1

Ponte Ponte Ponte Ponte Ponte Ponte

Alta Alta Alta Alta Alta Alta

Mb., Mb., Mb., Mb., Mb., Mb.,

lia Mar lia Mar lia Mar lia Mar lia Mar lia Mar

Fm. Fm. Fm. Fm. Fm. Fm.

Dias-Brito Dias-Brito Dias-Brito Dias-Brito Dias-Brito Dias-Brito

et et et et et et

al. al. al. al. al. al.

(2001) (2001) (2001) (2001) (2001) (2001)

lia Fm. Ponte Alta Mb., Mar lia Fm. Ponte Alta Mb., Mar lia Fm. Ponte Alta Mb., Mar lia Fm. Serra da Galga Mb., Mar

Dias-Brito et al. (2001) Dias-Brito et al. (2001) Dias-Brito et al. (2001) Senra and Silva-Silva (1998)

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213

Appendix A (continued) Taxon INVERTEBRATES Gastropods Ostracoda Altanicupris australis Candonopsis sp. Ilyocypris setembrinopetrii Ilyocpris sp. 1 Lycopterocypris angulata Neuquenocypris minor mineira Virgatocypris mezzalirai gen. et sp. indet. 1 gen. et sp. indet. 2 Gastropoda Physidae Physa aridi Viviparidadae Viviparus souzai Bivalvia Florencia peiropolensis Anodontites (Anodontites) pricei Musculium sp. VERTEBRATA Vertebrata indet. Pisces Characiformes indet. Perciformes indet. Siluriformes Pisces indet. Anura Baurubatrachus pricei Chelonia Cambaremys langertoni gen. et sp. indet. Lepidosauria Pristiguana brasiliensis Mesoeucrocodylia Itasuchus jesuinoi Peirosaurus tormini Uberabasuchus terricus Dinosauria Dinosauria indet. Sauropoda Megaloolithidae indet. Locality Provenance Selected references

13 12 12 12 13 12, 13 12, 13 12 13

Ponte Ponte Ponte Ponte Ponte Ponte Ponte Ponte Ponte

Alta Alta Alta Alta Alta Alta Alta Alta Alta

Mb., Mb., Mb., Mb., Mb., Mb., Mb., Mb., Mb.,

lia Mar lia Mar lia Mar lia Mar lia Mar lia Mar lia Mar lia Mar lia Mar

Fm. Fm. Fm. Fm. Fm. Fm. Fm. Fm. Fm.

Dias-Brito Dias-Brito Dias-Brito Dias-Brito Dias-Brito Dias-Brito Dias-Brito Dias-Brito Dias-Brito

et et et et et et et et et

al. al. al. al. al. al. al. al. al.

(2001) (2001) (2001) (2001) (2001) (2001) (2001) (2001) (2001)

1 1 1 1 1 11, 12, 13, 14 1 1 1 3 7 2 1 1 1 1 1

lia Fm. Serra da Galga Mb., Mar lia Fm. Serra da Galga Mb., Mar lia Fm. Serra da Galga Mb., Mar lia Fm. Serra da Galga Mb., Mar lia Fm. Serra da Galga Mb., Mar

Senra (2002) Senra (2002) Mezzalira (1974) Mezzalira (1974) Senra (2002)

Ponte Alta and Serra da Galga mbs., Candeiro and Bergqvist (2004) lia Fm. Mar lia Fm. Serra da Galga Mb., Mar lia Fm. Serra da Galga Mb., Mar lia Fm. Serra da Galga Mb., Mar lia Fm. Ponte Alta Mb., Mar lia Fm. Serra da Galga Mb., Mar lia Fm. Serra da Galga Mb., Mar lia Fm. Serra da Galga Mb., Mar lia Fm. Serra da Galga Mb., Mar lia Fm. Serra da Galga Mb., Mar lia Fm. Serra da Galga Mb., Mar lia Fm. Serra da Galga Mb., Mar Gayet and Brito (1989) Gayet and Brito (1989) Gayet and Brito (1989) Fernandes (1998) Baez and Peri (1989) Franc a and Langer (2005) Bertini et al. (1993) Estes and Price (1973) Price (1955) Price (1955) Carvalho et al. (2004) Candeiro et al. (2004b)

4, 7, 8, 9 Serra da Galga Uberaba Fm., Mb., lia Fm. Mar 1 Uberaba Fm., Serra da Galga Mb., lia Fm. Mar

Ribeiro (2002a, b) (continued on next page)

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Appendix A (continued) Taxon gen. et sp. nov. Titanosauria Baurutitan britoi Trigonosaurus pricei Aeolosaurus sp. gen. et sp. indet. Theropoda Abelisauridae indet. Carcharodontosauridae indet. Maniraptora indet. gen. et sp. indet. Locality 7 Provenance Selected references Uberaba Fm., Ponte Alta and Serra Santucci (2002), Candeiro et al. lia Fm. da Galga mbs, Mar (2004b) lia Fm. Serra da Galga Mb., Mar lia Fm. Serra da Galga Mb., Mar lia Fm. Serra da Galga Mb., Mar Kellner et al. (2005) Campos et al. (2000) Bertini et al. (2000), Santucci (2002) Powell (1986, 1987, 2003), Santucci and Bertini (2006) Candeiro (2002), Candeiro et al. (2004a) Candeiro (2002), Candeiro et al. (2004a) Novas et al. (2005) Candeiro (2002)

1 1 1

1, 2, 5, 6, Uberaba Fm., Serra da Galga Mb., lia Fm. 7 Mar 1,2 1,2 lia Fm. Serra da Galga Mb., Mar lia Fm. Serra da Galga Mb., Mar

lia Fm. 1 Serra da Galga Mb., Mar lia Fm. 1,2, 10, Serra da Galga Mb., Mar 11, 12, 13

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