Geobios3 9( 2006)3 19–327

http://france.elsevier.com/direct/GEOBIO/

Original article

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

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

Vertebrados fósseis da Formação Adamantina (Neocretáceo)do Distrito Paleontológico do Prata, Estado de Minas Gerais, Brasil

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, Universidadef ederal de uberlândia (UFU), Avenida J oão N aves de Á vila 2160, 38408-100 Uberlândia, MG, B razilbLaboratório de macrofósseis, departamento de g eologia, Universidadef ederal do Rio de Janeiro (UFRJ),

avenida B rigadeiro Trompowski, s /n, I lha do Fundão, 21949-900 Rio de Janeiro, RJ, B razilcMuseum of Victoria, P O Box 666E, M elbourne, Victoria, A ustralia

dSchool of geosciences, P O Box 28E, monash University, Victoria 3800, A ustralia

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

Abstract

In this contribution is given a p reliminary u p-to-date annotated list of all fossil vertebrates from the Turonian–Santonian AdamantinaForma- tion, Bauru Group where it occurs in the Prata p aleontological district which is located 45 km to the west of Prata in Minas GeraisState, Brazil. The A damantina Formation is a reddish sandstone in the Triângulo Mineiro region. These fluviolacustrine sediments weredeposited in a semiarid environment. Three fossil vertebrate sites occur in the Prata p aleontological district. The diversity of vertebrates in theAdamantina Formation is modest, and its components comprise a mixture of typical austral Gondwana t axa (such as abelisaurids,A eolosaurus) and b oreal Gondwana taxa (such as Carcharodontosauridae). The absence of Laurasian taxa in the u pper p art of the BauruGroup is noteworthy. Excepting some turtles, crocodylians, and one titanosaurid, most of the specimens discovered in this Group are b ased onfragmentary and isolated remains, which make it difficult to correctly identify the fossils. For this reason, many of the t axa have onlyb een identified in a p reliminary manner. Despite this drawback, the dinosaurs are typical Gondwana forms with no evidence of Laurasianaffinities. This is concordant with the idea that the Laurasian dinosaurs entered South America at the end of the Late Cretaceous; i.e. after theTuronian–Santonian, the age of the A damantina Formation.©2006 Elsevier SAS. All r ights reserved.

Résumé

La liste des v ertébrés fossiles est mise àj our dans la Formation Adamantina datée du Turonien-Santonien affleurant dans le districtp aléonto- logique de Prata à45 km à l’est de Prata (Minas Gerais, Brésil). La Formation A damantina est composée de grès rougeâtres dansla région du Triangle Minier. Les sédiments fluviolacustres se sont déposés dans u n environnement semi-aride. Trois sites à vertébrésfossiles sont connus dans le district p aléontologique de Prata. La diversité des vertébrés de la Formation A damantina est réduite et comprendu n mélange de taxons typiques du Gondwana austral (comme des Abelisauridés, A eolosaurus) et des taxons du Gondwana b oréal (comme desCarcharodontosauridae).

* Corresponding author.E-mail address: candeiro@yahoo.com.br (C.R.A. Candeiro).

0016-6995/$ - see front matter ©2006 Elsevier SAS. All rights r eserved.doi:10.1016/j.geobios.2005.10.003320 C.R.A. Candeiro et al. / Geobios 39 (2006) 319–327

L’absence de t axons laurasiatiques dans la p artie supérieure du Groupe Bauru est r emarquable. Excepté quelques tortues, des crocodiliens etu n titanosauridé, la plupart des spécimens découverts dans ce Groupe Bauru sont r eprésentés par des restes fragmentaires et isolés. Pour cetteraison, seules des identifications p réliminaires sont p roposées. Toutefois, les dinosaures sont typiques du Gondwana sans évidenced’affinité Laura- sienne. Ceci est en accord avec l’hypothèse que les dinosaures laurasiens ne seraient arrivés en Amérique du Sud qu’à la finde Crétacé supérieur, c’est-à-dire après le Turonien-Santonien, âge de la Formation A damantina.©2006 Elsevier SAS. All r ights reserved.

Resumo

Na presente contribuição é apresentada uma listagem p reliminar 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 arenitosavermelhados que afloram na região do Triângulo Mineiro. Esses sedimetnos de origem fluvio-lacustre foram depositados em u m ambientesemi-árido. São conhe- cidos três sítios de vertebrados fósseis que ocorrem no Distrito Paleontológico do Prata. A diversidade de v ertebradosna Formação Adamantina é modesta sendo representados por uma mistura típica de taxa austrais do Gondwana (abelisaurídeos e A eolosaurus)e boreais (carcarodotontosau- rídeos). A ausência de taxa laurasianos na p arte superior do Grupo Bauru é notável. Exceto p ela presença detartarugas, crocodilianos e u m titanossaurídeo a maioria dos espécimes descobertos nessa unidade geológica é baseada em restosfragmentários e isolados os quais são de difícil identificação. Por essa razão, a maioria desses taxa têm sido somente identificados de formap reliminar. Considerando essa desvantagem, se p ode afirmar que os dinossauros do Prata são tipicamente de formas gondwânicas comnenhuma evidência de formas laurasianas. Isso é concordante com a idéia de que os dinossauros laurasianos entraram n a América do Sulsomente no final do N eocretáceo, ex. depois do Turoniano-Santo- niano, que é a idade da Formação Adamantina.©2006 Elsevier SAS. All r ights reserved.

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

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

Palabras-chaves: N eocretáceo; Brasil; Crocodilos; Dinossauros; Tartarugas

1. Introduction

Since the b eginning of the last century, numerous specimens

of fossil vertebrates have b een collected from continental Cre-

taceous r ocks of the Adamantina Formation, Bauru Group, in

the Bauru Basin, Triângulo Mineiro region, Minas Gerais

State. In the same area of Minas Gerais State, the Marília For-

mation (Ponte Alta and Serra da Galga Members) also has

yielded fossil vertebrates. Both formations have produced fau-

nas consisting p rimarily of fishes, turtles, crocodylians, and di-

nosaurs. All of these fossils occur in sandstones deposited in

braided river channels.

A large quantity of fossil bones has b een found in the con-

tinental Cretaceous rocks exposed in the Prata p aleontological

district 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 p aleontological district, located

45 km w est of the town of Prata, is k nown as one of the most

important fossil locations, as it contains abundant remains of

sauropod and theropod dinosaurs as w ell crocodylians and tur-

tles.

Many authors (Soares et al., 1980; Suguio and Barcelos,

1983; Barcelos, 1984; Candeiro, 2002) assign the sediments

of Prata p aleontological district to the Adamantina Formation

(Bauru Group), which was dated as Turonian–Santonian in age

(Dias-Brito et al., 2001). This u nit is exposed at the Serra da

Boa 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 p aleontologicaldistrict have b een k nown since the 1920s (Price, 1961). Owingto a lack of reliable lithological and p aleontological data, thegeological ages of these sites are not w ell established.

The Adamantina Formation was formally proposed b ySoares et al. (1980). The u nit, 2–20 m t hick, consists of r oseto b rown fine grained, cross-bedded sandstone. The type sec-tion of this formation was in the town of Adamantina, SãoPaulo State. In Triângulo Mineiro, the A damantina Formationis the b asal lithostratigrafic u nit of the Bauru Group and iswidely exposed in the area.

In the Prata p aleontological district, Triângulo Mineiro(Fig. 1), three fossil vertebrate sites occur in the AdamantinaFormation. These three fossil sites represent the b est record ofLate Cretaceous vertebrates in the Minas Gerais State (Brazil).This makes those sites some of the most important w hen dis-cussing the fossil r ecord of Late Cretaceous in Minas GeraisState.

The vertebrates reported here are from the A damantinaFormation near Prata, and b elong to the following groups: abe-lisaurs, carcharodontosaurs, A eolosaurus, and titanosaur dino-saurs, turtles, and crocodylians. More is known about the dino-saurs than any other group from the Prata p aleontologicaldistrict. The p aleontologist Llewellyn Ivor Price first reportedto the Departamento Nacional de Produção Mineral fragmen-tary dinosaur b ones in 1940. The fossils were transferred toRio de Janeiro (Price, 1961). Since then, there have b een nu-merous fossil collecting trips to the Prata p aleontological dis-trict that have yielded new dinosaurs and other v ertebrate fos-sils (Silva et al., 2001). However, detailed stratigraphic studiesbased on dinosaur remains were not attempted.

C.R.A. Candeiro et al. / Geobios 39 (2006) 319–327 321

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

2. Geological setting

The Adamantina Formation was defined b y 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 p art 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 A raçatuba and São José do Rio Preto Members.

The Prata p aleontological district is currently regarded asformed by rocks of the Bauru Group (Late Cretaceous)(Fig. 2). In the Triângulo Mineiro region, the A damantina,Marília and Uberaba formations sensu Fernandes and Coimbra(1996) form p art of Bauru Group. These rocks overlay the ba-

salt layers of the Serra Geral Formation.Goldberg and Garcia (2000) described the A damantina For-

mation (Fig. 3) in the Prata p aleontological district as very fineto fine grained, w ell-sorted, cross-bedded red sandstones.Furthermore, t hese authors r ecognized intraformational con-glomerates and subordinate red shales, which are massive orlaminar. The top of this u nit is characterized b y the interbed-ding of (1) conglomerates, (2) fine- to m edium-grained cross-bedded sandstones and (3) abundant nodular and h orizontalcarbonate concretions, which coarsen and thicken upward.

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

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).

Von Huene (1927, 1939) regarded the Adamantina Fm asSenomanian b ased 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 t his Formation to b e Turonian–Santonian (92–83 my).

3. Vertebrate paleontology

The Late Cretaceous sequence of the Prata p aleontologicaldistrict 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 h illy areas located west of Pra-ta (Fig. 1). Because of the fragmentary nature of the fossilscollected thus far, they have not b een identified to a refinedtaxonomic level except the genus A eolosaurus.

C.R.A. Candeiro et al. / Geobios 39 (2006) 319–327 323

Table 1Vertebrate fauna of the Adamantina Formation from Prata p aleontological district. See footnotes for source r eferences 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 etal. (2002), 9 Almeida et al., 2003 (in p ress)TaxaLocation1 Location2 Location2

Boa vista hill GPS. 19°27′26.0″S Outskirt GPS. 19°26′21.0″S “Idelmo Bramh farm” GPS. 19°26′31.8″Sand 49°14′42.6″W. and 49°15′41.3″W. and 49°14′27.8″W.

Vertebrata

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

x

Fig. 4. Dinosaur remains from Prata p aleontological district. A, labial view, B, lingual view Abelisauridae tooth (MMR/UFU-PV 0006); C, labial view, D,lingual view, Carcharodontosauridae tooth (MMR/UFU-PV 005); E, labial v iew, F, lingual view, Theropoda tooth (MMR/UFU-PV 007). Bar scale represent30 mm. G, lateral view, Titanosauria caudal vertebrae (MMR/UFU-PV 0001); H, I, lateral view, Titanosauria r ibs. Bar scale A , B, C, D, E, F bar scale = 30 mm;G = 10 mm; H, I= 50 mm.

3.1. Remarks about the dinosaurs of the P rata district Abbreviations: MACN, Museo Argentino de Ciencias

N at-

urales, Buenos Aires, Argentina; MUCPV-CH, Museo de la

Dinosaurs are the best-known and most diverse taxa from Universidad Nacional de Comahue, El Chocon

collection; Prata Site. This material is housed mainly at MMR and MMR/UFU-PV, Museu de Minerais e Rochas,

Universidade UFRJ-DG and briefly commented here. Federal de Uberlândia, Uberlândia, Minas

Gerais, Brazil;

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

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

3.1.1. S ystematic p aleontologySuborder THEROPODA Marsh, 1881ABELISAURIA N ovas, 1992Family ABELISAURIDAE Bonaparte and N ovas, 1985Material: (MMR/UFU-PV 0006) (Fig. 4A, B).Locality: 2.Remarks: the Abelisauridae are represented b y a single

tooth (MMR/UFU-PV 0006) from locality 2 on the westernside of the Prata p aleontological district. This tooth was de-scribed recently b y Marinho (2003). It lacked a root but thepreservation of the crown is good. It is small (32 mm), w ithserrated anterior and p osterior carinae, the denticles on theproximal p art of the anterior carina are smaller than t hose onthe distal p art, and denticles on the m iddle are largest. All thedenticles on the distal p art are pointed toward the crown apex.The denticles on the proximal and middle p art of the anteriorcarina are p erpendicular to the crown apex. The denticles onboth carinae are higher than wide. The denticles of the p roxi-mal and distal p arts of the p osterior carinae are the same size;those in the middle are larger. The cross-section of the t oothhas the anterior face compressed and p osterior face convex(MMR/UFU-PV 0006) has the labial and lingual faces moreconvex in cross-section nears the anterior edge and flatter p os-teriorly; the lingual face is also slightly concave p roximodis-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 p re-sence of a low crown in the Abelisauridae (e.g. A ucasaurusgarridoi, Carnotaurus sastrei, R ajasarus narmadensis) teeth

and regarded it as a diagnostic character of the family. Can-deiro et al. (2004) pointed that abelisaurid teeth are character-ized b y labial and lingual faces more convex in cross-sectionnear the anterior edge and flatter p osteriorly; 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, w hich has a labial side of the tooth that is convex onits more anterior portion and p osteriorly flattened. A s on thelabial side, the anterior p art of the lingual side is slightly con-vex, becoming p osteriorly flattened. The p osterior r egion of thelingual side b ears acute wrinkles on the enamel, which are verywell developed, on the proximal and medial region, but t hesevanish toward the crown tip. Each wrinkle is obliquely orientedwith respect to the tooth margin, but b ends along its lengthaway from tooth margin until its orientation is transverse withrespect t o the tooth’s long axis. The wrinkle then continuesacross the tooth surface as a horizontal b and. The p osteriordenticles on the proximal and distal ends are equal, while themiddle denticles are larger. The anterior denticles are smallerthan the p osterior ones. All the denticles of b oth carinae areperpendicular to the crown apex, and are r ather 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 b een considered as a diagnostic feature for Carcharodonto-sauridae (Carcharodontosaurus s aharicus and Giganotosauruscarolinii) b y 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.) w hich can b e strong or subdued. These wrinklesmay or may not b e close to the crown b order (e.g. Carcharo-

dontosaurus and Giganotosaurus).The wrinkles on the crown of MMR/UFU-PV 005 are

strong and similar to the wrinkles found in carcharodontosaur-ids. This specimen is mediolaterally compressed and thus quitesimilar to Carcharodontosaurus s aharicus (Stromer, 193 1) 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 r epresents 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 p arts 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 b e 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 w ith great variation in sizeand shape throughout the carina. Their cross-sections are dif-ferent from any of the k nown Gondwanan Theropoda, but it isimportant to note that the teeth of some Gondwanan dinosaursare still unknown (Novas, 1997).

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

dant fossil v ertebrate remains found in the region (Fig. 4G–I).

Numerous p ostcranial b ones r epresent them. Almeida et al.

(2003, 2004) (UFRJ-DG 270-R caudal vertebra) (Fig. 5A–C)

C.R.A. Candeiro et al. / Geobios 39 (2006) 319–327 325

Fig. 5. A eolosaurus caudal vertebra UFRJ-DG 270-R. Lateral v iew (A),posterior (B) and anterior (C). Abbreviations: CN, neural canal; PRZ,postzigapophysis; PT, transverse process; PZ, postzygapophysy; SN, neural

spine. Scale bar 50 mm.

and Henriques et al. (2002) reported the presence of titanosaur-id sauropods. Almeida et al. (2003, 2004) recognized the t ita-nosaurid genus A eolosaurus, w hich 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 b one. The v ertebral centrum is deepand r elatively low, p rocoelous, and b ears a h eart-shaped ante-rior articular facet (Fig. 5C). The transverse process is w elldeveloped and p ositioned at the centro-neural suture. The b aseof the neural arch is p ositioned anteriorly in relation t o 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 p respinal laminae(PRSL) (Fig. 5A). The spinoprezygapophyseal laminae are alsowell developed and b oth are j oined to the b ase of the PRSL.The postzygapophysis is w ell 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 p rocoelous condi-tion of this caudal vertebrate is a synapomorphic feature thatallies it w ith the “Titanosauridae” (Bonaparte and Coria, 1993;Wilson and U pchurch, 2003). The h eart-shaped p osterior ar-ticular facet and w ide p rezygapophyses allow u s to r elate thisvertebra to the taxa Gondwanatitan/Aeolosaurus. Recently, atitanosaurid (MN 5013-V) was recognized at the same locality(the Boa V ista 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 t o thesegenera.

TITANOSAURIA indetMaterial: (MMR-UFU-PV 001). Caudal v ertebrae.

Locality: 2.Remarks: there are several fragments of disarticulated

bones found in A damantina Formation sediments in the Pratapaleontological district that have not p reviously b een attributedto the Titanosauridae (sensu Powell, 1986, 2003; McIntosh,1990). These specimens are caudal v ertebrae (MMR-UFU-PV001; Fig. 4G) that are strongly p rocoelous, having “ball sock-et” p osterior articular faces. According Powell (1986, 2003)and McIntosh (1990) among Titanosauria, titanosaurids devel-oped anterior caudals w ith a deeply excavated anterior facemuch like a “socket” and a p osterior articulation like a p romi-nent “ball”.

3.1.2. A damantina F ormation f ossil localities in the P ratapaleontogical district

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

4. Concluding remarks

The fossils from the Prata p aleontological district are allvertebrates mainly collected at the Boa Vista Hill fossil site,Locality 2. The b est-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 p oorly known, w ith few w ell-preserved specimens,many fragmentary remains have b een found. Fossil localitiesare centered in the western p arts 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-

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

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 n amed Prata taxa h ave b eenfully reported, a picture is emerging w hich 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. A eolosaurus is found not only inthe Prata p aleontological district but also in the Late Cretac-eous of A rgentina.

Typical Laurasian taxa such as ornithischians have b eenwidely documented in several Late Cretaceous u nits of South-ern South America (e.g. Bonaparte et al., 1984; Salgado andCoria, 1996; De Valais et al., 2003; N ovas et al., 2004). Be-cause they are absent from earlier Turonian–Santonian depositsincluding the A damantina Formation, this suggests t hat thephysical connection between N orth and South America wasnot established u ntil the end of the Cretaceous.

Acknowledgments

We thank Agustin G. Martinelli (Museo A rgentino de Cien-cias N aturales, 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. N ovas (Museo Ar-

gentino de Ciencias N aturales). The authors also t hank Alberto

Garrido (Museo Carmen Funes, Cipoletti) and an anonymous

reviewer for critically reading the manuscript and p roviding

valuable suggestions. We specially thank Lílian Paglarelli

Bergqvist, Leonardo S. Avilla and Erika A branches (Labora-

tório de Macrofósseis, Departamento de Geologia, U niversi-

dade Federal do Rio de Janeiro) for allowing u s free access

to laboratories and specimens in their charge. Adelino Carval-

ho, Cláudia Regina A. Candeiro, Edeilson P. Silva, Edivane

Cardoso, Flávio Oliveira, Jorge Augusto Albuquerque, Patricia

Tavares (Uberlândia), and Sérgio Moraes (Prata) deserve many

thanks for assistance during 2002–2003 field seasons. Cláudia

Regina A. Candeiro (Uberlândia) and ÉricBuffetaut (Research

at Centre National de la Recherche Scientifique, Paris) made

résumé observations and correction of the grammar.

References

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