untitled EARTH SCIENCES RESEARCH JOURNAL Earth Sci. Res. J. Vol. 13, No.1 (June 2009): 6-15 VERTEBRATES OF THE MARÍLIA FORMATION (LATE MAASTRICHTIAN) FROM THE PEIRÓPOLIS PALEONTOLOGICAL SITE: TOWARD A BETTER UNDERSTANDING Carlos Roberto A. Candeiro Laboratório de Geologia, Curso de Geografia, Campus do Pontal, Universidade Federal de Uberlândia, Avenida José João Dib, 2545, Bairro Progresso, Zip Code 38302-000, Ituiutaba, Minas Gerais State, Brazil Corresponding author. Tel.: ++ (55) 32692389. E-mail address: candeiro@yahoo.com ABSTRACT The Peirópolis paleontological site (Late Cretaceous, Maastrichtian) in Minas Gerais State, Brazil yielded an important assemblage of fossil vertebrates. The typical occurrence of South American widespread taxa in Peirópolis is important for correlation between the Brazilian Bauru Basin and Argentinean Late Cretaceous bas- ins. The fishes, turtles, anuran, crocodilians and dinosaurs known from Peirópolis (Marília Formation) resemble the Patagonian latest Late Cretaceous vertebrate faunas but lacks ornithischian dinosaurs. Key words: Vertebrates, Late Cretaceous, Marília Formation, Minas Gerais State, Brazil. RESUMEN En el Sítio Paleontológico de Peirópolis (cretácico superior, Maastrichtiano) ubicado en la província de Minas Gerais, Brasil, hay un contenido muy importante de vertebrados fósiles. La ocurrencia en Peirópolis de taxa típicos y de amplia distribución en la América del Sur es importante para la correlación entre la Cuenca Bauru y cuencas argentinas del cretácico superior. Los pesces, tortugas, anuros, cocodrilianos y dinosaurios de Peirópolis (Formación Marília), se equivalen a las faunas de vertebrados del Neocretácico de Patagonia, sin em- bargo, sin la presencia de dinosaurios ornitísquios. Palabras claves: Vertebrados, Neocretácico, Formación Marília, Província de Minas Gerais, Brasil. 6 Manuscript received: January 10th, 2009. Accepted for publication: April 30th, 2009. JULIO 6-GEOCIENCIAS-VOL 13-1 2009.prn D:\GEOCIENCIAS 13-1 JUN 2009\GEOCIENCIAS-VOL 13-1 2009.vp martes, 07 de julio de 2009 18:10:37 p p p Composite 133 lpi at 45 degrees Introduction The Upper Cretaceous beds at the Peirópolis paleontological site of (Uberaba municipality, Minas Gerais State; Fig. 1) consist on an entirely continental sedimentary succession. The Upper Cretaceous Bauru Group units at Peirópolis are divided into the Uberaba Formation (Coniacian-Santonian) and Marília Formation (late Maastrichtian) in ascending order. The Uberaba Formation occurs as small patches mainly in the Peirópolis village. Although extensive paleontological studies have been carried out in Peirópolis area (e.g., Price, 1955; Estes and Price, 1973; Báez and Peri, 1989; Gayet and Brito, 1989; Carvalho et al., 2004; França and Langer, 2005; Kellner and Campos, 2005; Kellner et al., 2005; Campos et al., 2005; Novas et al., 2005; Santucci and Bertini, 2006; Novas et al., 2008; Salgado and Carvalho, 2008; Candeiro et al., 2008), fossil vertebrates are unknown so far from Uberaba Formation, except for unstudied dinosaur eggs found near Peirópolis. On the other hand, abundant vertebrate remains have been reported from the late Maastrichtian Marília Formation (specimens and localities are listed in the Tab. 1 and 2). The first discovery of ver- tebrate fossils was vertebrate fragments from the “Ponto 1 do Price” from the Serra da Galga Member (Marília Formation) in the 1940´s (Price, 1951; Candeiro and Bergqvist, 2004). Since then, scattered dinosaur bones and other vertebrate fossils including fishes, turtles, and crocodilians have been found in quarries, roads cuts, and stream beds as a result of a Centro de Pesquisas Paleontológicas Llewelyn Ivor Price (Peirópolis) team survey. Abundant plants (palynomorphs) and mollusks have been located from the Peirópolis quarries. They clearly indicate that diversity of vertebrate communities existed dur- ing the time of Marília Formation deposition in the Peirópolis area. Except for some the crocodilians, however, the vertebrate faunas remain inadequately published. Therefore, the purpose of this paper is to review the vertebrate fossils from the Marília Forma- tion at this locality and report their occurrence to the paleontological community. Comments and correla- tions with other vertebrate-bearing formations of Late Cretaceous age from Argentina are discussed. The potential significance of Marília Formation ver- tebrates from Peirópolis may be seen especially in the 7 VERTEBRATES OF THE MARÍLIA FORMATION (LATE MAASTRICHTIAN) FROM THE PEIRÓPOLIS PALEONTOLOGICAL SITE: TOWARD A BETTER UNDERSTANDING N 0° 70°W 10°S 20°S 40°W 50°W 60°W 51”00’ 18”00’ 47”30’ 18”00’ 20”30’ 51”00’ Brazil 20”30’ GOIÁS STATE SÃO PAULO STATE 47”30’ 0 10 20 km Legend Town Localization study area Road River Par ana iba Rive r Araguari River Tejuco River Peirópolis Grande River Uberaba Prata Ituiutaba Araguari Uberlãndia Figure 1. Geographic distribution of the Upper Cretaceous “Marília-Points” of Peirópolis paleontological site. Point 1 (5 quarries); Point 2 (1 quarry); Point 3 (1 quarry); Point 4 (2 quarries); Point 5 (1 quarry). JULIO 6-GEOCIENCIAS-VOL 13-1 2009.prn D:\GEOCIENCIAS 13-1 JUN 2009\GEOCIENCIAS-VOL 13-1 2009.vp martes, 07 de julio de 2009 18:10:39 p p p Composite 133 lpi at 45 degrees context of roughly contemporaneous areas from southern South America (Candeiro and Bergqvist, 2004). Methodology The data on vertebrates from Peirópolis pa- leontological site for the present work are mostly based in literature sources as well as in the direct ob- servation of the specimens deposited at Museu dos Dinossauros (Peirópolis District, Uberaba Town, Minas Gerais State, Brazil). For the stratigraphic units in the Bauru Group I follow the arrangement proposed by Dias Brito et al. (2001). Results Geological setting The Peirópolis paleontological site is northeast of Bauru Basin (Upper Cretaceous) (Fernandes and Coimbra, 1996). The Bauru Group in Triângulo Mineiro region is represented by sediments belong- ing to the Adamantina, Marília and Uberaba forma- tions (Fig. 2). These layers overlie basalts belonging to Serra Geral Formation (São Bento Group, Paraná Basin), sandstone belonging to Botucatu Formation (São Bento Group), metamorphic and Proterozoic outcrops belonging to Araxá and Canastra groups (Sanfransiscana Basin) and Mesozoic intrusion of the 8 CARLOS ROBERTO A. CANDEIRO Figure 2. Geological map of Bauru Group in Triângulo Mineiro region (modified Fernandes and Coimbra, 1996). JULIO 6-GEOCIENCIAS-VOL 13-1 2009.prn D:\GEOCIENCIAS 13-1 JUN 2009\GEOCIENCIAS-VOL 13-1 2009.vp martes, 07 de julio de 2009 18:10:39 p p p Composite 133 lpi at 45 degrees Soerguimento of Alto Paranaíba (Suguio et al., 1979). Outcrops of the Marília Formation are unevenly distributed throughout the Peirópolis paleontological site. According to Barcelos and Suguio (1987), that unit was deposited by coalescing alluvial fans, and later reworked by a braided system in association with calcretes and lacustrine calcareous sediments. Barcelos (1984) subdivided the Marília Formation into the Echaporã, Ponte Alta, and Serra da Galga members, although only the latter two members are exposed in the Peirópolis area. Ponte Alta and Serra da Galga members show the following characteris- tics (Garrido et al., 1992): Ponte Alta member– a basal “calcareous white member”, characterized by thin and medium calcareous beds with pebbles and calcareous nodules which were deposited in lakes; Serra da Galga member– “member of white sand- stone and conglomerate”, composed of bleached conglomerate and sandstone with feldspar matrix de- posited by alluvial fans, braided rivers and lakes. Those two members, according to Suguio (1973, 1980), were formed in a partially dry climate, which would have favored the fossil preservation found in these layers. Dias-Brito et al. (2001) assigned a late Maastrichtian age to the Marília Formation. This lithostratigraphic unit yielded invertebrates and plants, and abundant vertebrates (Fig. 3). Verte- brates from the Marília Formation are represented by anurans (Baurubatrachus pricei), lizards (Pristiguana brasiliensis), crocodilians (Itasuchus jesuinoi, Peirosaurus tormini, Uberabasuchus terrificus), chelonians (Cambaremys langertoni), dinosaurs (Abelisauridae, Carcharodontosauridae, Maniraptora, and eutitanosaurian Baurutitan britoi, Trigonosaurus pricei, Uberabatitan ribeiroi, Aeolosaurus). Vertebrate paleontology The vertebrate fossils of the Triângulo Mineiro re- gion have been known since 1920, however only with the pioneer studies of paleontologist Llewellyn Ivor Price, which started in 1947, the fossils found in the Peirópolis paleontological site became not only more known, but at the same time better classified. Just a small part of collected material in this paleontological site was described to the level of spe- cies due to fragmentary nature of most the specimens which made a more refined description impossible (e.g., Fig. 4). Abundant fish fossils have recently been found in the Peirópolis. To date four localities have yielded isolated scales and teeth. They produced Characiformes, Perciformes and Siluriformes. The species which were described formally are repre- sented by amphibian Baurubatrachus pricei Báez and Peri, 1989; lepidosaurian Pristiguana brasiliensis Estes and Price, 1973 and crocodilians Itasuchus jesuinoi, Peirosaurus tormini Price, 1955 9 VERTEBRATES OF THE MARÍLIA FORMATION (LATE MAASTRICHTIAN) FROM THE PEIRÓPOLIS PALEONTOLOGICAL SITE: TOWARD A BETTER UNDERSTANDING BAURU GROUP PEIRÓPOLIS SITE Serra da Galga Mb. (Marília Fm.) Ponte Alta Mb. (Marília Fm.) Uberaba Fm. Legend Plant Vertebrate Fish Frog Lizard Turtle Crocodilian Sauropod Theropod Figure 3. Upper Cretaceous Bauru Group generalized geo- logical section in Peirópolis showing occurrence of the biota. JULIO 6-GEOCIENCIAS-VOL 13-1 2009.prn D:\GEOCIENCIAS 13-1 JUN 2009\GEOCIENCIAS-VOL 13-1 2009.vp martes, 07 de julio de 2009 18:10:39 p p p Composite 133 lpi at 45 degrees and Uberabasuchus terrificus Carvalho, Ribeiro and Avilla, 2004. The dinosaurs represent the most abundant remains and they are represented by theropods which were attributed by Candeiro (2002) and Candeiro et al. (2004) to Abelisauridae and Carcharodontosauridae, first Abelisauridade bone remains by (Novas et al., 2008) and Maniraptoran claw recently described by Novas et al. (2005). The sauropods are represented by the titanosaurians Baurutitan britoi Campos and Kellner, 2005 and Trigonosaurus pricei Campos, Kellner, Bertini and Santucci, 2005; Uberabatitan ribeiroi Salgado and Carvalho, 2008 and others dif- ferent elements e.g., teeth, vertebrae, ribs, hemal arches, phalanges and osteoderms. These speci- mens were studied by Campos and Kellner (1999), 10 CARLOS ROBERTO A. CANDEIRO Figure 4. Reptilian fossils from the Peirópolis paleontological site. A, lower jaw Itasuchus jesuinoi (modi- fied from Price, 1955); B, pelvis of the “Titanosauridae” (modified from Campos and Kellner, 1999); C, Abelisauridae teeth; D, Carcharodontosauridae tooth; E, Theropoda indet. tooth. Scale bar in mm. Table 1. Vertebrate fauna of the Marília Formation from Peirópolis paleontological site. Area 1 Area 2 Area 3 Area 4 Area 5 Ponte Alta Member x x x Serra da Galga Member x x VERTEBRATA Vertebrata indet x x x x x PISCES Characiform x x Perciform x x Siluriform x x ANURA Baurubatrachus pricei x REPTILIA Squamata Pristiguana brasiliensis Chelonia Chelonia indet. Podcnemidae Crocodylomorpha Itasuchus jesuinoi Peirosaurus tormini Uberabasuchus terrificus Dinosauria Sauropoda Titanosauria x x x x x x JULIO 6-GEOCIENCIAS-VOL 13-1 2009.prn D:\GEOCIENCIAS 13-1 JUN 2009\GEOCIENCIAS-VOL 13-1 2009.vp martes, 07 de julio de 2009 18:10:40 p p p Composite 133 lpi at 45 degrees 11 VERTEBRATES OF THE MARÍLIA FORMATION (LATE MAASTRICHTIAN) FROM THE PEIRÓPOLIS PALEONTOLOGICAL SITE: TOWARD A BETTER UNDERSTANDING Area 1 Area 2 Area 3 Area 4 Area 5 “Titanosauridae” A “Titanosauridae” B “Titanosauridae” C “Titanosaurinae” “Titanosaurus” Aeolosaurus sp. Baurutitan britoi Trigonosaurus pricei Theropoda Theropoda indet. Abelisauridae indet. Carcharadontosauridae indet. Maniraptora x x x x x x x x x x x x x Table 2. Fossiliferous areas (quarries), in the Peirópolis paleontological site, Minas Gerais State. Areas Generalities Reference selected Area 1 (quarry 1) This outcrop has been known as “Caieira” Campos and Kellner (1999) Area 1 (quarry 2) This outcrop is inside of “Caieira” Kellner (1996) Area 1 (quarry 3) This quarry has been known as “Point 2” Price (1955) Area 1 (quarry 4) This outcrop has been known as “Point 3” Campos and Kellner (1999), Trotta et al. (2002) Área 1 (quarry 5) This outcrop is known as “Mumbuca” Powell (1987, 2003) Area 2 The outcrop is situated on the BR-262 throughway about 3,5 km east of Peirópolis Campos and Kellner (1999) Area 3 The outcrop is situated on the Ponte Alta district throughway about 18 km east from Peirópolis Magalhães-Ribeiro (2002) Area 4 (quarry 1) Quarry 1 is known as the “ Rio Tejuco 1” Albuquerque and Candeiro (2003) Area 4 (quarry 2) Quarry 2 is known as the “Rio Tejuco 2” Albuquerque and Candeiro (2003) Area 5 This area has been known as “Partezan Quarry” Bertini et al. (1993) JULIO 6-GEOCIENCIAS-VOL 13-1 2009.prn D:\GEOCIENCIAS 13-1 JUN 2009\GEOCIENCIAS-VOL 13-1 2009.vp martes, 07 de julio de 2009 18:10:40 p p p Composite 133 lpi at 45 degrees Santucci (2002), Powell (2003) and Santucci and Bertino (2006) who attributed part of these remains to Titanosauria (see detailed assignments in Tab. 1). Dinosaur eggs found in sediments of the Serra da Galga Member are related to “Megaloolithidae” oofamily, according to Magalhães-Ribeiro (2002). Furthermore there are numerous cranial and post cranial remains belonging to crocodilians and tur- tles. Numerous fragments of disarticulated bones and coprolites are found in Marília Formation sedi- ments in the Peirópolis paleontological site, how- ever they have not been attributed to taxonomic level below Vertebrata; they are housed in the Centro de Pesquisas Paleontológicas Llewellyn Ivor Price and Museu de Ciências da Terra (Rio de Ja- neiro). Fossil sites from Marília Formation in the Peirópolis paleontological site of: The first fossil site was studied by Price in the 1940´s. Since then, new fossil localities of the Marília Formation have been found in the Uberaba municipality with five men- tioned in literature so far (Fig. 1; Tab. 2). The best known fossil vertebrates of Marília For- mation found in the Peirópolis paleontological site of are listed below. Discussion The present study and those of the previous works in the Marília Formation Peirópolis paleontological site), it is attempted here to point a comprehensive point of view of this area. The Peirópolis paleontological site contains a moderately diverse Late Cretaceous (Maastrichtian) vertebrate assemblage, from which some of the taxa are reported also in Argentina (e.g., Chubut, Neuquén and Malargüe groups; Candeiro and Martinelli, 2003; Lamanna et al., 2003; Martinelli and Forasiepi, 2004; Leanza et al., 2004). Some of the vertebrate taxa reported at Peirópolis occur with other vertebrates known from other areas of Argentina, and are important for corre- lation with Argentinean units. Supporting this corre- lation are the mesoeucrocodylian Peirosaurus tormini and the titanosaur Aeolosaurus, which are found in Bajo de la Carpa, Allen, Angostura Colorada, Los Alamitos, and Bajo Barreal forma- tions in Patagonia. Abelisaurid and carcharodon- tosaurid remains are known to occur in the Bauru Group. In addition to the titanosaurian material from Peirópolis area, remains of those sauropods are widely known in Upper Cretaceous units of Argen- tina. Additional support for correlation is given by the presence in Peirópolis of the Gondwanan “Megaloolithidae” eggshells. Other support of the correlation between Peirópolis area and Argentinean areas may be given by other vertebrate groups, but more detailed studies are needed before this can be done. For example, characiform, siluriform and perciform fishes and podocnemid turtles are known in most of the Neuquén and Malargüe groups, but they cannot be used as correlation tools since their remains are not well known from the Marília Formation. Ornithischian records are remarkable in Argen- tina (Late Cretaceous), however, not a single ornithischian remain has been identified so far in the Marília Formation or the Bauru Group. Considering that ornithischians occurred in Argentina, this bias may be from lack of extensive or least systematic prospecting in the Marília Formation rather than from depositional or preservational events. Conclusions So far, four fossil sites have been recorded at the Peirópolis paleontological site of, the Marília Forma- tion with a diverse vertebrate fauna of Upper Maastrichtian age. These fossils are represented mainly by fishes, frogs, turtles, crocodilians, and di- nosaurs. The diverse fossiliferous, but poorly known Marília Formation accumulated in the northwestern Bauru Basin during the latest Cretaceous. Deposition occurred on a fluvio-lacustrine environment. The Maastrichtian age of the Marília Formation sug- gested by previous authors has several important biogeographical and paleontological correlations. First, it indicates that the Marília Formation assem- blage is approximately contemporaneous in age with 12 CARLOS ROBERTO A. CANDEIRO JULIO 6-GEOCIENCIAS-VOL 13-1 2009.prn D:\GEOCIENCIAS 13-1 JUN 2009\GEOCIENCIAS-VOL 13-1 2009.vp martes, 07 de julio de 2009 18:10:40 p p p Composite 133 lpi at 45 degrees the vertebrate assemblage recovered from the Patagonia (e.g., Chubut, Malargüe, and Neuquén groups). This finding is certainly consistent with striking similarities between faunas from these two southern South America areas. Second, this Maastrichtian Marília Formation fauna, when con- sidered in the light of the Late Cretaceous discover- ies, was different by the lack of ornithischian records in Brazil, but which are found in Argentina. Acknowledgments This paper forms part of the undergraduate project of the R. Candeiro who gratefully acknowledges finan- cial assistance in the form of an Undergraduate Fel- lowship (1998-1999) received from the Council of Scientific from Brazil (CNPq/166/98) under Dr. Ignácio Brito advisor (financial supported by FAPEMIG/UFU/801/98 fellow). The manuscript was substantially improved after the revision work of Dr. Yaneth Muñoz-Saba (Universidad Nacional de Co- lombia). I would like to thank Prof. Luiz Carlos Borges Ribeiro and the Centro de Pesquisas Paleontológicas Llewellyn Ivor Price crew, Dr. Adriano Rodrigues dos Santos (Universidade Fed- eral de Uberlândia, UFU), Dr. Lílian Paglarelli Bergqvist (Universidade Federal do Rio de Janeiro, UFRJ) and Thiago da Silva Marinho (UFRJ) for re- viewing the earlier drafts of the manuscript. The manuscript was substantially improved by of Dr. Simone Magnuco (Museo Civico di Storia Naturale di Milano, Italy), Dr. Walter Hartwig (Touro Univer- sity, United States) and Dr. David Gillette (Museum of Northern Arizona, United States). The author also would like to thank the friends, Lisete, Ana and Fernando Brito, that in some ways helped us to start our Paleontological studies, since they are wife, daughter and son of Professor Ignácio Machado Brito, who opened the doors of his house and ignited my life into Paleontology. References Albuquerque, J.A.C. and Candeiro, C.R.A., (2003). 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