Vertebrate fossils from the Adamantina Formation (Late Cretaceous), Prata paleontological district, Minas Gerais State, Brazil

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Geobios 39 (2006) 319–327

Original article

* CorrE-ma

0016-69doi:10.1

Vertebrate fossils from the Adamantina Formation (Late Cretaceous),

Vertébrés fossiles du Crétacé supérieur de la Formation Adamantina,district paléontologique de Prata, État de Minas Gerais, Brésil

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Vertebrados fósseis da Formação Adamantina (Neocretáceo)

Carlos Roberto A. Candeiro a,b,*, Adriano R. Santos a, Thomas H. Rich c,d, Thiago S. Marinho a,b,Emerson C. Oliveira a

aMuseu de minerais e rochas, Universidade federal de uberlândia (UFU), Avenida João Naves de Ávila 2160, 38408-100 Uberlândia, MG, Brazil

g authoss: can

e frontbios.20

b Laboratório de macrofósseis, departamento de geologia, Universidade federal do Rio de Janeiro (UFRJ),

cMuseum of Victoria, PO Box 666E, Melbourne, Victoria, Australiad School of geosciences, PO Box 28E, monash University, Victoria 3800, Australia

Received 11 May 2004; accepted 21 October 2004Available online 22 May 2006

Abstract

In this contribution is given a preliminary up-to-date annotated list of all fossil vertebrates from the Turonian–Santonian Adamantina Forma-tion, Bauru Group where it occurs in the Prata paleontological district which is located 45 km to the west of Prata in Minas Gerais State, Brazil.The Adamantina Formation is a reddish sandstone in the Triângulo Mineiro region. These fluviolacustrine sediments were deposited in a semiaridenvironment. Three fossil vertebrate sites occur in the Prata paleontological district. The diversity of vertebrates in the Adamantina Formation ismodest, and its components comprise a mixture of typical austral Gondwana taxa (such as abelisaurids, Aeolosaurus) and boreal Gondwana taxa(such as Carcharodontosauridae). The absence of Laurasian taxa in the upper part of the Bauru Group is noteworthy. Excepting some turtles,crocodylians, and one titanosaurid, most of the specimens discovered in this Group are based on fragmentary and isolated remains, which make itdifficult to correctly identify the fossils. For this reason, many of the taxa have only been identified in a preliminary manner. Despite thisdrawback, the dinosaurs are typical Gondwana forms with no evidence of Laurasian affinities. This is concordant with the idea that the Laurasiandinosaurs entered South America at the end of the Late Cretaceous; i.e. after the Turonian–Santonian, the age of the Adamantina Formation.© 2006 Elsevier SAS. All rights reserved.

Résumé

La liste des vertébrés fossiles est mise à jour dans la Formation Adamantina datée du Turonien-Santonien affleurant dans le district paléonto-logique de Prata à 45 km à l’est de Prata (Minas Gerais, Brésil). La Formation Adamantina est composée de grès rougeâtres dans la région duTriangle Minier. Les sédiments fluviolacustres se sont déposés dans un environnement semi-aride. Trois sites à vertébrés fossiles sont connusdans le district paléontologique de Prata. La diversité des vertébrés de la Formation Adamantina est réduite et comprend un mélange de taxonstypiques du Gondwana austral (comme des Abelisauridés, Aeolosaurus) et des taxons du Gondwana boréal (comme des Carcharodontosauridae).

[email protected] (C.R.A. Candeiro).

matter © 2006 Elsevier SAS. All rights reserved.05.10.003

C.R.A. Candeiro et al. / Geobios 39 (2006) 319–327320

L’absence de taxons laurasiatiques dans la partie supérieure du Groupe Bauru est remarquable. Excepté quelques tortues, des crocodiliens et untitanosauridé, la plupart des spécimens découverts dans ce Groupe Bauru sont représentés par des restes fragmentaires et isolés. Pour cette raison,seules des identifications préliminaires sont proposées. Toutefois, les dinosaures sont typiques du Gondwana sans évidence d’affinité Laura-sienne. Ceci est en accord avec l’hypothèse que les dinosaures laurasiens ne seraient arrivés en Amérique du Sud qu’à la fin de Crétacé supérieur,c’est-à-dire après le Turonien-Santonien, âge de la Formation Adamantina.© 2006 Elsevier SAS. All rights reserved.

Resumo

Na presente contribuição é apresentada uma listagem preliminar de todos os fósseis de vertebrados do Distrito Paleontológico do Prata (For-mação Adamantina, Turoniano-Santoniano, Grupo Bauru). As rochas da Formação Adamantina são compostas por arenitos avermelhados queafloram na região do Triângulo Mineiro. Esses sedimetnos de origem fluvio-lacustre foram depositados em um ambiente semi-árido. São conhe-cidos três sítios de vertebrados fósseis que ocorrem no Distrito Paleontológico do Prata. A diversidade de vertebrados na Formação Adamantina émodesta sendo representados por uma mistura típica de taxa austrais do Gondwana (abelisaurídeos e Aeolosaurus) e boreais (carcarodotontosau-rídeos). A ausência de taxa laurasianos na parte superior do Grupo Bauru é notável. Exceto pela presença de tartarugas, crocodilianos e umtitanossaurídeo a maioria dos espécimes descobertos nessa unidade geológica é baseada em restos fragmentários e isolados os quais são de difícilidentificação. Por essa razão, a maioria desses taxa têm sido somente identificados de forma preliminar. Considerando essa desvantagem, se podeafirmar que os dinossauros do Prata são tipicamente de formas gondwânicas com nenhuma evidência de formas laurasianas. Isso é concordantecom a idéia de que os dinossauros laurasianos entraram na América do Sul somente no final do Neocretáceo, ex. depois do Turoniano-Santo-niano, que é a idade da Formação Adamantina.© 2006 Elsevier SAS. All rights reserved.

Keywords: Late Cretaceous; Brazil; Crocodiles; Dinosaurs; Turtles

Mots clés : Crétacé supérieur ; Brésil ; Crocodiliens ; Dinosaures ; Tortues

Palabras-chaves: Neocretáceo; Brasil; Crocodilos; Dinossauros; Tartarugas

1. Introduction

Since the beginning of the last century, numerous specimensof fossil vertebrates have been collected from continental Cre-taceous rocks of the Adamantina Formation, Bauru Group, inthe Bauru Basin, Triângulo Mineiro region, Minas GeraisState. In the same area of Minas Gerais State, the Marília For-mation (Ponte Alta and Serra da Galga Members) also hasyielded fossil vertebrates. Both formations have produced fau-nas consisting primarily of fishes, turtles, crocodylians, and di-nosaurs. All of these fossils occur in sandstones deposited inbraided river channels.

A large quantity of fossil bones has been found in the con-tinental Cretaceous rocks exposed in the Prata paleontologicaldistrict of the Triângulo Mineiro region of Minas Gerais State,Brazil (Fig. 1; Goldberg et al., 1995a, 1995b; Silva et al., 2001;Marinho, 2003; Candeiro et al., 2003; Marinho et al., 2003).For this reason, the Prata paleontological district, located45 km west of the town of Prata, is known as one of the mostimportant fossil locations, as it contains abundant remains ofsauropod and theropod dinosaurs as well crocodylians and tur-tles.

Many authors (Soares et al., 1980; Suguio and Barcelos,1983; Barcelos, 1984; Candeiro, 2002) assign the sedimentsof Prata paleontological district to the Adamantina Formation(Bauru Group), which was dated as Turonian–Santonian in age(Dias-Brito et al., 2001). This unit is exposed at the Serra daBoa Vista Hill, in the vicinity of Prata.

Some of the richest Cretaceous fossil vertebrate localities inCentral Brazil occur in the Adamantina Formation. Many ofthe Adamantina Formation sites in the Prata paleontological

district have been known since the 1920s (Price, 1961). Owingto a lack of reliable lithological and paleontological data, thegeological ages of these sites are not well established.

The Adamantina Formation was formally proposed bySoares et al. (1980). The unit, 2–20 m thick, consists of roseto brown fine grained, cross-bedded sandstone. The type sec-tion of this formation was in the town of Adamantina, SãoPaulo State. In Triângulo Mineiro, the Adamantina Formationis the basal lithostratigrafic unit of the Bauru Group and iswidely exposed in the area.

In the Prata paleontological district, Triângulo Mineiro(Fig. 1), three fossil vertebrate sites occur in the AdamantinaFormation. These three fossil sites represent the best record ofLate Cretaceous vertebrates in the Minas Gerais State (Brazil).This makes those sites some of the most important when dis-cussing the fossil record of Late Cretaceous in Minas GeraisState.

The vertebrates reported here are from the AdamantinaFormation near Prata, and belong to the following groups: abe-lisaurs, carcharodontosaurs, Aeolosaurus, and titanosaur dino-saurs, turtles, and crocodylians. More is known about the dino-saurs than any other group from the Prata paleontologicaldistrict. The paleontologist Llewellyn Ivor Price first reportedto the Departamento Nacional de Produção Mineral fragmen-tary dinosaur bones in 1940. The fossils were transferred toRio de Janeiro (Price, 1961). Since then, there have been nu-merous fossil collecting trips to the Prata paleontological dis-trict that have yielded new dinosaurs and other vertebrate fos-sils (Silva et al., 2001). However, detailed stratigraphic studiesbased on dinosaur remains were not attempted.

Fig. 1. Location of the Prata paleontological district, and geographic distribution of the Cretaceous fossiliferous location.

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2. Geological setting

The Adamantina Formation was defined by Soares et al.(1980) and is known in the following states of Southeast Brazil:Goiás, Mato Grosso do Sul, Minas Gerais and São Paulo. Theseauthors regarded it as part of the Bauru Group, which occurs inthe Paraná Basin. Based on variation in the regional lithology,Barcelos (1984) divided the Adamantina Formation into twounits: the Araçatuba and São José do Rio Preto Members.

The Prata paleontological district is currently regarded asformed by rocks of the Bauru Group (Late Cretaceous)(Fig. 2). In the Triângulo Mineiro region, the Adamantina,

Marília and Uberaba formations sensu Fernandes and Coimbra(1996) form part of Bauru Group. These rocks overlay the ba-salt layers of the Serra Geral Formation.

Goldberg and Garcia (2000) described the Adamantina For-mation (Fig. 3) in the Prata paleontological district as very fineto fine grained, well-sorted, cross-bedded red sandstones.Furthermore, these authors recognized intraformational con-glomerates and subordinate red shales, which are massive orlaminar. The top of this unit is characterized by the interbed-ding of (1) conglomerates, (2) fine- to medium-grained cross-bedded sandstones and (3) abundant nodular and horizontalcarbonate concretions, which coarsen and thicken upward.

Fig. 2. Geological map of the Bauru Group (Upper Cretaceous) in the Triângulo Mineiro region, Minas Gerais State, Brazil (modified de Fernandes and Coimbra, 1996).

Fig. 3. Bauru Group lithostratigraphic chart in Prata area (modified fromSuguio and Barcelos, 1983).

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Von Huene (1927, 1939) regarded the Adamantina Fm asSenomanian based on a correlation made between the turtle Na-jadochelys and the sauropod “Titanosaurs australis” with similartaxa recorded in northern Patagonia, Argentina. Based on ostra-cods and caryophytes, Dias-Brito et al. (2001) considers the ageof this Formation to be Turonian–Santonian (92–83 my).

3. Vertebrate paleontology

The Late Cretaceous sequence of the Prata paleontologicaldistrict contains fossil remains of crocodylians, dinosaurs andturtles (Table 1; Silva et al., 2001; Marinho, 2003; Albuquer-que et al., 2003; Marinho et al., 2003; Moraes et al., 2003).

The fossil occurrences are in hilly areas located west of Pra-ta (Fig. 1). Because of the fragmentary nature of the fossilscollected thus far, they have not been identified to a refinedtaxonomic level except the genus Aeolosaurus.

Table 1Vertebrate fauna of the Adamantina Formation from Prata paleontological district. See footnotes for source references included. 1 Goldberg et al. (1995a, 1995b), 2

Silva et al. (2001), 3 Marinho (2003), 4 Albuquerque et al. (2003), 5 Candeiro et al. (2003), 6 Marinho et al. (2003), 7 Moraes et al. (2003), 8 Henriques et al. (2002), 9

Almeida et al., 2003 (in press)

Taxa Location 1 Location 2 Location 2Boa vista hill GPS. 19°27′26.0″Sand 49°14′42.6″W.

Outskirt GPS. 19°26′21.0″Sand 49°15′41.3″W.

“Idelmo Bramh farm” GPS. 19°26′31.8″Sand 49°14′27.8″W.

VertebrataVertebrata indet 1–7 x x xReptiliaChelonia indet. 2 xCrocodylomorpha indet. 2 xDinosauriaSauropodaTitanosauria 2–8 xAeolosaurus 9 xTheropodaTheropoda indet. 3-7 xAbelisauridae 3, 7 xCarcharodontosauridae 3 x

Fig. 4. Dinosaur remains from Prata paleontological district. A, labial view, B, lingual view Abelisauridae tooth (MMR/UFU-PV 0006); C, labial view, D, lingualview, Carcharodontosauridae tooth (MMR/UFU-PV 005); E, labial view, F, lingual view, Theropoda tooth (MMR/UFU-PV 007). Bar scale represent 30 mm. G,lateral view, Titanosauria caudal vertebrae (MMR/UFU-PV 0001); H, I, lateral view, Titanosauria ribs. Bar scale A, B, C, D, E, F bar scale = 30 mm; G = 10 mm; H,I = 50 mm.

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3.1. Remarks about the dinosaurs of the Prata district

Dinosaurs are the best-known and most diverse taxa fromPrata Site. This material is housed mainly at MMR andUFRJ-DG and briefly commented here.

Abbreviations: MACN, Museo Argentino de Ciencias Nat-urales, Buenos Aires, Argentina; MUCPV-CH, Museo de laUniversidad Nacional de Comahue, El Chocon collection;MMR/UFU-PV, Museu de Minerais e Rochas, UniversidadeFederal de Uberlândia, Uberlândia, Minas Gerais, Brazil;

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RTMP, Royal Tyrrell Museum of Palaeontology, Drumheller,Canada; UFRJ-DG-R, Departamento de Geologia, Universi-dade Federal do Rio de Janeiro, Rio de Janeiro, Brazil, fossilreptilian collection.

3.1.1. Systematic paleontologySuborder THEROPODA Marsh, 1881ABELISAURIA Novas, 1992Family ABELISAURIDAE Bonaparte and Novas, 1985Material: (MMR/UFU-PV 0006) (Fig. 4A, B).Locality: 2.Remarks: the Abelisauridae are represented by a single

tooth (MMR/UFU-PV 0006) from locality 2 on the westernside of the Prata paleontological district. This tooth was de-scribed recently by Marinho (2003). It lacked a root but thepreservation of the crown is good. It is small (32 mm), withserrated anterior and posterior carinae, the denticles on theproximal part of the anterior carina are smaller than those onthe distal part, and denticles on the middle are largest. All thedenticles on the distal part are pointed toward the crown apex.The denticles on the proximal and middle part of the anteriorcarina are perpendicular to the crown apex. The denticles onboth carinae are higher than wide. The denticles of the proxi-mal and distal parts of the posterior carinae are the same size;those in the middle are larger. The cross-section of the toothhas the anterior face compressed and posterior face convex(MMR/UFU-PV 0006) has the labial and lingual faces moreconvex in cross-section nears the anterior edge and flatter pos-teriorly; the lingual face is also slightly concave proximodis-tally.

The characters exhibited by the low crown tooth fromMMR/UFU-PV 0006 clearly indicate that it represents an abe-lisaurid. Bonaparte (1996), Lamanna et al. (2002), Wilson andUpchurch (2003), and Candeiro et al. (2004) noted the pre-sence of a low crown in the Abelisauridae (e.g. Aucasaurusgarridoi, Carnotaurus sastrei, Rajasarus narmadensis) teethand regarded it as a diagnostic character of the family. Can-deiro et al. (2004) pointed that abelisaurid teeth are character-ized by labial and lingual faces more convex in cross-sectionnear the anterior edge and flatter posteriorly; the lingual face isalso slightly concave proximodistally. The cross-section of thespecimen (MMR/UFU-PV 0006) is exactly as in the Abelisaur-idae.

Family CARCHARODONTOSAURIDAE Stromer, 1931Material: (MMR/UFU-PV 005) (Fig. 4C, D). Tooth.Locality: 2.Remarks: Marinho (2003) recently described MMR/UFU-

PV 005, which has a labial side of the tooth that is convex onits more anterior portion and posteriorly flattened. As on thelabial side, the anterior part of the lingual side is slightly con-vex, becoming posteriorly flattened. The posterior region of thelingual side bears acute wrinkles on the enamel, which are verywell developed, on the proximal and medial region, but thesevanish toward the crown tip. Each wrinkle is obliquely orientedwith respect to the tooth margin, but bends along its length

away from tooth margin until its orientation is transverse withrespect to the tooth’s long axis. The wrinkle then continuesacross the tooth surface as a horizontal band. The posteriordenticles on the proximal and distal ends are equal, while themiddle denticles are larger. The anterior denticles are smallerthan the posterior ones. All the denticles of both carinae areperpendicular to the crown apex, and are rather higher thanwide. The cross-section is chestnut-shaped. The outline of thetooth in lateral view is in the form of a rough isosceles triangle.

The presence of wrinkles on a theropod teeth tooth crownhas been considered as a diagnostic feature for Carcharodonto-sauridae (Carcharodontosaurus saharicus and Giganotosauruscarolinii) by many authors (Larsson, 1996; Sereno et al., 1996;Vickers Rich et al., 1999; Candeiro et al., 2004). In Giganoto-saurus carolinii different forms of wrinkles occur (Candeiropers. obsers.) which can be strong or subdued. These wrinklesmay or may not be close to the crown border (e.g. Carcharo-dontosaurus and Giganotosaurus).

The wrinkles on the crown of MMR/UFU-PV 005 arestrong and similar to the wrinkles found in carcharodontosaur-ids. This specimen is mediolaterally compressed and thus quitesimilar to Carcharodontosaurus saharicus (Stromer, 1931) andthe Giganotosaurus (MUCPV-CH-1 and casts of MACN S/Nand RTMP 94.43.1, 94.43.2, 94.43.3, 94.43.4).

The presence of carcharodontosaurids in the PPD is espe-cially noteworthy as it represents the youngest record of thisclade found to date (Turonian–Santonian). Previously, theyoungest known records of carcharodontosaurids are fromCenomanian beds in Africa, Argentina and Maranhão State inBrazil. Carcharodontosaurid teeth from the MMR/UFU-PV005 of Late Cretaceous age contrast with the osteological re-cord of Carcharodontosauridae, which is mainly restricted toolder parts of the Cretaceous (e.g. Albian–Turonian) of Africaand South America.

Order THEROPODA indet.Material: (MMR/UFU-PV 0007) (Fig. 4E, F). Tooth.Locality: 2.Remarks: no family assignment could be made for MMR/

UFU-PV 0007, but there is no doubt that this specimen be-longs to a theropod dinosaur as it is strongly labio-linguallycompressed and shows denticles with great variation in sizeand shape throughout the carina. Their cross-sections are dif-ferent from any of the known Gondwanan Theropoda, but it isimportant to note that the teeth of some Gondwanan dinosaursare still unknown (Novas, 1997).

TITANOSAURIA Bonaparte and Coria, 1993Genus Aeolosaurus Powell, 1986Aeolosaurus indet.Material: (UFRJ-DG 270-R). Caudal vertebra.Locality: 2.Remarks: the sauropods are the most common and abun-

dant fossil vertebrate remains found in the region (Fig. 4G–I).Numerous postcranial bones represent them. Almeida et al.(2003, 2004) (UFRJ-DG 270-R caudal vertebra) (Fig. 5A–C)

Fig. 5. Aeolosaurus caudal vertebra UFRJ-DG 270-R. Lateral view (A),posterior (B) and anterior (C). Abbreviations: CN, neural canal; PRZ,postzigapophysis; PT, transverse process; PZ, postzygapophysy; SN, neuralspine. Scale bar 50 mm.

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and Henriques et al. (2002) reported the presence of titanosaur-id sauropods. Almeida et al. (2003, 2004) recognized the tita-nosaurid genus Aeolosaurus, which was first described fromthe Late Cretaceous of Patagonia (Argentina) (Powell, 1986,2003; Salgado and Coria, 1993; Salgado, 2000). Although thisvertebra is almost complete, it did suffer some damage. Thisdamage made it possible to observe the internal spongy com-position (Fig. 5B) of the bone. The vertebral centrum is deepand relatively low, procoelous, and bears a heart-shaped ante-rior articular facet (Fig. 5C). The transverse process is welldeveloped and positioned at the centro-neural suture. The baseof the neural arch is positioned anteriorly in relation to thevertebral centrum. The neural canal is oval—the lateral dia-meter is the longer one. The neural spine is large and poster-iorly oriented, showing a well-developed prespinal laminae(PRSL) (Fig. 5A). The spinoprezygapophyseal laminae are alsowell developed and both are joined to the base of the PRSL.The postzygapophysis is well marked. The haemal arch isstraight and laterally compressed. Although its proximal regionis fragmented, its Y-shaped nature is readily seen. The openhaemal canal indicates that this caudal vertebra belongs to theCamarasauromorpha. Within that group, the procoelous condi-

tion of this caudal vertebrate is a synapomorphic feature thatallies it with the “Titanosauridae” (Bonaparte and Coria, 1993;Wilson and Upchurch, 2003). The heart-shaped posterior ar-ticular facet and wide prezygapophyses allow us to relate thisvertebra to the taxa Gondwanatitan/Aeolosaurus. Recently, atitanosaurid (MN 5013-V) was recognized at the same locality(the Boa Vista Hill) (Table 1). However, Almeida et al. (2004)pointed out that the specimen lacked features of the Gondwa-natitan/Aeolosaurus complex. Henriques et al. (2002) sug-gested that this fossil was a new taxon closely related to thesegenera.

TITANOSAURIA indetMaterial: (MMR-UFU-PV 001). Caudal vertebrae.Locality: 2.Remarks: there are several fragments of disarticulated

bones found in Adamantina Formation sediments in the Pratapaleontological district that have not previously been attributedto the Titanosauridae (sensu Powell, 1986, 2003; McIntosh,1990). These specimens are caudal vertebrae (MMR-UFU-PV001; Fig. 4G) that are strongly procoelous, having “ball sock-et” posterior articular faces. According Powell (1986, 2003)and McIntosh (1990) among Titanosauria, titanosaurids devel-oped anterior caudals with a deeply excavated anterior facemuch like a “socket” and a posterior articulation like a promi-nent “ball”.

3.1.2. Adamantina Formation fossil localities in the Pratapaleontogical district

Price studied the first fossil site in the Prata paleontologicaldistrict in the 1950’s. However, it was only in 2002 that fossillocalities in the Adamantina Formation were discovered in thatarea by researchers from the Universidade Federal de Uberlân-dia (Museu de Minerais e Rochas). Three fossil vertebrate sitesin the Adamantina Formation are mentioned in the literature(e.g. Albuquerque et al., 2003). The best-known vertebrate fos-sils of Adamantina Formation occur in the Prata paleontologi-cal district and are listed in Table 1.

4. Concluding remarks

The fossils from the Prata paleontological district are allvertebrates mainly collected at the Boa Vista Hill fossil site,Locality 2. The best-known taxa from the Adamantina Forma-tion are referred to the Abelisauridae, Carcharodontosauridaeand Titanosauria (Aeolosaurus and Titanosauria indet.).

Although the vertebrate fossil record of Prata District is cur-rently poorly known, with few well-preserved specimens,many fragmentary remains have been found. Fossil localitiesare centered in the western parts of Prata municipality, andresemble the fauna from southern South America (Patagonia).The dinosaurs of the Prata District comprise two families ofGondwanan theropods (Abelisauridae and Carchadorodonto-sauridae), one genus of titanosaurs (Aeolosaurus). Single cro-codilian and turtle records are reported in the literature fromthe Prata District. Their presence supports the earlier conclu-

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Bo

Bo

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Ca

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Vo

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sions of Von Huene (1927, 1939) that correlated the fauna ofAdamantina Formation with those found in the Late Cretac-eous of Argentina or even those of Africa, India and Madagas-car.

Although only a few of the named Prata taxa have beenfully reported, a picture is emerging which suggests a Gondwa-nan origin for the assemblage (Bonaparte, 1986). Carcharodon-tosaurs are common in the early Late Cretaceous in SouthAmerica (Coria and Salgado, 1995; Leanza et al., 2004), butin the Prata District of Minas Gerais State a record of thisgroup indicates that it possibly survived into the Late Cretac-eous. Titanosaurs and abelisaurs dominate the Late Cretaceousin the Gondwana landmasses. They are found also in Late Cre-taceous of the Prata District. Aeolosaurus is found not only inthe Prata paleontological district but also in the Late Cretac-eous of Argentina.

Typical Laurasian taxa such as ornithischians have beenwidely documented in several Late Cretaceous units of South-ern South America (e.g. Bonaparte et al., 1984; Salgado andCoria, 1996; De Valais et al., 2003; Novas et al., 2004). Be-cause they are absent from earlier Turonian–Santonian depositsincluding the Adamantina Formation, this suggests that thephysical connection between North and South America wasnot established until the end of the Cretaceous.

Acknowledgments

We thank Agustin G. Martinelli (Museo Argentino de Cien-cias Naturales, Buenos Aires) for thorough and helpful com-ments that greatly improved this manuscript. For references,we thank Jeffrey A. Wilson (University of Michigan), P. Up-church (University of Cambridge) and F. Novas (Museo Ar-gentino de Ciencias Naturales). The authors also thank AlbertoGarrido (Museo Carmen Funes, Cipoletti) and an anonymousreviewer for critically reading the manuscript and providingvaluable suggestions. We specially thank Lílian PaglarelliBergqvist, Leonardo S. Avilla and Erika Abranches (Labora-tório de Macrofósseis, Departamento de Geologia, Universi-dade Federal do Rio de Janeiro) for allowing us free accessto laboratories and specimens in their charge. Adelino Carval-ho, Cláudia Regina A. Candeiro, Edeilson P. Silva, EdivaneCardoso, Flávio Oliveira, Jorge Augusto Albuquerque, PatriciaTavares (Uberlândia), and Sérgio Moraes (Prata) deserve manythanks for assistance during 2002–2003 field seasons. CláudiaRegina A. Candeiro (Uberlândia) and Éric Buffetaut (Researchat Centre National de la Recherche Scientifique, Paris) maderésumé observations and correction of the grammar.

References

Albuquerque, J.A.C., Oliveira, E.C., Marinho, T.S., Candeiro, C.R.A., 2003.Jazigos fossilíferos cretácicos do Triângulo Mineiro, indentificação e ma-peamento. Boletim de Resumos do 18e Congresso Brasileiro de Paleonto-logia. Brasília-DF, Brazil (2003, pp. 322).

Almeida, E.B., Avilla, L.S., Candeiro, C.R.A., 2003. Restos caudais associa-dos a Titanosauridae do Cretáceo Superior da Formação Adamantina, Ba-cia Bauru, Município do Prata – MG. Boletim de Resumos do 18e Con-gresso Brasileiro de Paleontologia. Brasília-DF, Brazil (2003, pp. 36).

eida, E.B., Avilla, L.S., Candeiro, C.R.A. 2004, Restos associados a Tita-nosauridae do Cretáceo Superior da Formação Adamantina, Bacia Bauru,Município do Prata – MG. Revista Brasileira de Paleontologia 7 (2), 239–244.rcelos, J.H., 1984. Reconstrução paleogeográfica da sedimentação do GrupoBauru baseada na sua redefinição estratigráfica parcial em território paulis-ta e no estudo preliminar fora do estado de São Paulo. Unpublished Ph.D.thesis, Universidade Estadual Paulista, Rio Claro, Brasil.naparte, J.F., 1986. History of the terrestrial Cretaceous vertebrales ofGondwana. IVo Congreso Argentino de Paleontología y Bioestratigrafía(Mendonza), Actas 4, 63–95.naparte, J.F., 1996. Cretaceous tetrapods of Argentina. Münchner Geowis-senchaftlichen Abhandlungen 30, 73–130.naparte, J.F., Coria, R.A., 1993. Un Nuevo y gigantesco saurópodo titano-saurio de la Formación Río Limay (Albiano-Cenomaniano) de la Provinciadel Neuquén, Argentina. Ameghiniana 30, 271–282.naparte, J.F., Franchi, M.R., Powell, J.E., Sepúlveda, E.G., 1984. La For-mación Los Alamitos (Campaniano-Maastrichtiano) del sudeste de Río Ne-gro, con descripción de Kritosaurus australia n. sp. (Hadrosauridae). Sig-nificado paleogeográfico de los vertebrados. Revista de la AsociaciónGeológica Argentina 39, 284–299.ndeiro, C.R.A., 2002. Dentes de Theropoda da Formação Marília (Santo-niano–Maastrichtiano), Bacia Bauru, Região de Peirópolis, Uberaba, MinasGerais, Brazil. Unpublished Ms. thesis, Universidade Federal do Rio deJaneiro, Brasil.ndeiro, C.R.A., Abranches, C.T., Abrantes, E.A., Avilla, L.S., Martins, V.C., Moreira, A.L., Torres, S.R., Bergqvist, L.P., 2004. Dinosaur remainsfrom Western São Paulo State, Brazil (Bauru Basin, Adamantina Forma-tion, Upper Cretaceous). Journal of South American Earth Science 18, 1–10.ndeiro, C.R.A., Marinho, T.S., Novais, G.T., Franco, A.C., Albuquerque, J.A.C., Oliveira, E.C., 2003. Previously unreported result from the UpperCretaceous of the Paleontological Site of Prata, Minas Gerais State, Brazil.Boletim de Resumos do 3e Simpósio Brasileiro de Paleontologia de Ver-tebrados. Rio de Janeiro-RJ, Brazil (2003, pp. 20).ria, R.A., Salgado, L., 1995. A new giant carnivorous dinosaur from theCretaceous of Patagonia. Nature 377, 224–226.Valais, S., Apesteguía, S., Sauthier, D.U., 2003. Nuevas evidencias de di-nosaurios de la Formación Puerto Yeruá (Cretácico), Provincia de EntreRíos, Argentina. Ameghiniana 40, 631–635.s-Brito, D.E.A., Musacchio, J.C., Castro, M.S.A., Maranhão, J.M., Suárez,J.M., Rodrigues, R., 2001. Grupo Bauru: uma unidade continental do Cre-táceo do Brasil – concepções baseadas em dados micropaleontológicos,isotópicos e estratigráficos. Revue de Paléobiologie 2, 245–304.nandes, L.A., Coimbra, A.M., 1996. A Bacia Bauru (Cretáceo Superior,Brasil). Anais da Academia Brasileira de Ciências 68, 195–205.ldberg, K., Azevedo, S.A.K., Garcia, A.J., 1995a. A ocorrência de fósseisde dinossauros no Município de Prata, MG. Boletim de Resumos do 14e

Congresso Brasileiro de Paleontologia. Uberaba-MG, Brazil (1995, pp.53).ldberg, K., Garcia, A.J., Azevedo, S.A.K., 1995b. Geologia e paleoam-biente dos sedimentos associados aos depósitos fossíliferos do Municípiode Prata, MG. Boletim de Resumos do 14e Congresso Brasileiro de Pa-leontologia. Uberaba-MG, Brazil (1995, pp. 54).ldberg, K., Garcia, A.J.V., 2000. Palaeogeography of the Bauru Group, adinosaur-bearing Cretaceous unit, northeastern Paraná Basin, Brazil. Cre-taceous Research 21, 241–254.nriques, D.R., Kellner, A.W.A., Azevedo, S.A.K., Santos, D.D., Craik, M.M.T., 2002. On a sauropod dinosaur (Titanosauridae) from the AdamantinaFormation (Late Cretaceous). Journal of Vertebrate Paleontology, Abstracts22, 64A.n Huene, F., 1927. Contribución a la paleogeografía de Sud-América – II.Las relaciones paleogeograficas de Sud América durante el CretáceoSuperior. Academina Nacional de Ciencias de Córdoba 30, 256–294.n Huene, F., 1939. Carta de F. von Huene ao Dr. Euzébio de Oliveira.Mineração e Metalurgia 4, 1–190.

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Lar

Lea

Ma

Ma

Mc

Mo

No

No

Po

Po

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Sal

Sal

Sal

Ser

Sil

So

Str

Su

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Wi

C.R.A. Candeiro et al. / Geobios 39 (2006) 319–327 327

anna, M.C., Martínez, R.D., Smith, J., 2002. A definitive abelisaurid ther-opod dinosaur from the early Late Cretaceous of Patagonia. Journal of Ver-tebrate Paleontology 22, 58–69.sson, H.C.E., 1996. Cranial morphology of the African theropod Carchar-odontosaurus saharicus (Allosauroidea). Journal of Vertebrate Paleontol-ogy 16, 47a.nza, H.A., Apesteguía, S., Novas, F., de la Fuente, M., 2004. Cretaceousterrestrial beds from the Neuquén Basin (Argentina) and their tetrapod as-semblages. Cretaceous Research 25, 61–87.rinho, T.S., 2003. As ocorrências fossilíferas do Cretáceo Superior nos Mu-nicípios do Prata e Monte Alegre de Minas, no Triângulo Mineiro. Unpub-lished Undergraduate Monograph, Universidade Federal de Uberlândia,Uberlândia, Brazil.rinho, T.S., Oliveira, E.C., Albuquerque, J.A.C., Ribeiro, L.C.B., Candeiro,C.R.A., 2003. Dinosaurs from the Bauru Basin (Upper Cretaceous). Bole-tim de Resumos do 18e Congresso Brasileiro de Paleontologia. Brasília-DF, Brazil (2003, pp. 322).Intosh, J.S., 1990. Sauropoda. In: Weishampel, D.B., Dodson, P., Osmólka,H. (Eds.), The Dinosauria. University of California Press, Berkeley, pp.345–401.raes, S.F., Oliveira, E.C., Albuquerque, J.A.C., Novais, G.T., Marinho, T.S., Candeiro, C.R.A., 2003. New discoveries of vertebrates fossils and theoccurrence of a new fossil location in the Prata Site (Upper Cretaceous),Minas Gerais State, Brazil. Boletim de Resumos do 21e Semana Científicade estudos Biológicos, Universidade Federal de Uberlândia. Uberlândia–MG, Brazil (2003, pp. 36).vas, F.E., 1997. South American dinosaurs. In: Currie, P.J., Padian, K.(Eds.), Encyclopedia of Dinosaurs. Academic Press, San Diego, pp. 678–689.vas, F.E., Cambiaso, A.V., Ambrosio, A., 2004. A new basal iguanodontian(Dinosauria, Ornithischia) from the Upper Cretaceous of Patagônia. Ame-ghiniana 41, 75–82.well, J.E., 1986. Revisión de los titanosáuridos de América del Sur. Unpub-lished Ph.D. thesis, Universidad Nacional de Tucumán, Argentina.well, J.E., 2003. Revision of South American Titanosaurid dinosaurs: Paleo-biological, Paleobiogeographical and Phylogenetic aspects. Records of theQueen Victoria, Melbourne 111, 1–173.

ce, L.I., 1961. Sobre os dinossáurios do Brasil. Academia Brasileira deCiências, Anais 33, 3–4.gado, L., 2000. Evolución y Paleobiología de los Saurópodos Titanosauri-dae. Unpublished Ph.D. thesis, Universidad Nacional de La Plata, Argenti-na.gado, L., Coria, R.A., 1993. El gênero Aeolosaurus (Sauropoda, Titano-sauridae) em la Formacion Allen (Campaniano-Maastrichtiano) de la Pro-víncia de Rio Negro, Argentina. Ameghiniana 30, 119–128.gado, L., Coria, R., 1996. First evidence of an armoured ornithischian di-nosaur in the late Cretaceous of North Patagonia, Argentina. Ameghiniana33, 367–371.eno, P.C., Dutheil, D.B., Iarochene, M., Larsson, H.C.E., Lyon, G.H.,Magwenw, P., Sidor, C.A., Varrichio, D.J., Wilson, J.A., 1996. Predatorydinosaurs from the Sahara and Late Cretaceous faunal differentiation.Science 272, 986–991.va, E.P., Brito, I.M., Santos, A.R., Candeiro, C.R.A., Ribeiro, L.C., 2001.Sítio Paleontológico do Prata, Triângulo Mineiro. Boletim de Resumos do17e Congresso Brasileiro de Paleontologia Rio Branco–AC, Brazil, (2001,pp. 195).ares, P.C., Landim, P.M.B., Fulfaro, V.J., Neto, A.F.S., 1980. Ensaio decaracterização estratigráfica do Cretáceo no Estado de São Paulo: GrupoBauru. Revista Brasileira de Geociências 10, 177–185.omer, E., 1931. Wirbeltierreste der Baharíje-Stufe (untertes Cenoman). 10.Ein Skellett-Rest von Carcharodotnosaurus nov. gen. Abhandlungen derBayerischen Akademie der Wissenschaften Mathematich-naturwis-senschaftliche Abteilung Neue Folge 9, 1–23.guio, K., Barcelos, J.H., 1983. Paleoclimatic evidence from the BauruGroup, Cretaceous of Paraná Basin, Brazil. Revista Brasileira de Geociên-cias 13, 232–236.kers Rich, P., Rich, T.H., Lanus, D.R., Rich, L.S., Vacca, R., 1999. “Bigtooth” from the Early Cretaceous of Chubut province, Patagonia: a possi-ble Carcharodontosaurid. National Science Museum Monographs 15, 85–88.lson, J.A., Upchurch, P., 2003. A revision of Titanosaurus Ludekker (Dino-sauria-Sauropoda), the first dinosaur genus with a “Gondwanan” distribu-tion. Journal of Systematic Paleontology 1, 125–160.