Quick viewing(Text Mode)

Theropod Teeth from the Marília Formation (Late Maastrichtian) at the Paleontological Site of Peirópolis in Minas Gerais State, Brazil

Theropod Teeth from the Marília Formation (Late Maastrichtian) at the Paleontological Site of Peirópolis in Minas Gerais State, Brazil

DOI: 10.5327/Z0375-75362012000200008 42(2): 323-330, junho de 2012

Theropod teeth from the Marília Formation (late ) at the paleontological site of Peirópolis in Minas Gerais State, Brazil

Carlos Roberto A. Candeiro1*, Philip J. Currie2, Lílian P. Bergqvist3

Abstract Theropod teeth can be separated at the level using the cross-section, and by the presence or absence of wrinkles in the enamel of the crowns. Theropods comprise the most common vertebrate remains in the assemblage from the Serra da Galga Member, Marília Formation (late Maastrichtian) in the Bauru Basin Peirópolis paleontological site (Uberaba Town, Minas Gerais State), in southeastern Brazil. In this study, cross-sectional outlines of the teeth and wrinkles on the surfaces of the crowns were used to group the teeth into nine morphological classes. The tooth cross-sections were drawn using a copper wire to contour the outlines of the most proximal portions of the tooth crowns. Other features were analyzed using a microscope. Within the nine morphological classes, morphotypes one and two can be assigned to the , and morphotypes three to seven belong to the . This is the most abundant record of Abelisauridae in Minas Gerais State, Brazil. Morphotypes eight and nine were not identified because they do not have any diagnostic features of known Gondwanan or Laurasian taxa. The presence of Carcharodontosauridae teeth in the Member of Serra da Galga (late Maastrichtian) is the youngest record for this family.

Keywords: ; Late ; theropod teeth; Minas Gerais State; Brazil.

Resumo Dentes de terópodes da Formação Marília (Neomaastrichtiano) do sítio paleontológico de Peirópolis do estado de Minas Gerais, Brasil. Dentes de dinossauros terópodes possuem várias características morfológicas a nível familiar. As principais características diagnósticas são: a forma do dente em seção transversal e a presença de enrugamentos na coroa do dente. Terópodes representam os restos de vertebrados mais comuns na assembleia fóssil do Membro Serra da Galga, Formação Marília (Neomaastrichtiano), Bacia Bauru, no sítio paleontológico de Peirópolis (cidade de Uberaba, estado de Minas Gerais), no Sudeste do Brasil. Neste estudo, seções transversais dos dentes e enrugamentos nas faces das coroas foram utilizadas para agrupar os dentes em nove classes morfológicas. As seções transversais dos dentes foram desenhadas utilizando-se um fio de cobre para contornar as porções mais proximais de suas respectivas coroas. Outras características foram analizadas, usando-se um microscópio. Dentro das dez classes morfológicas, os morfotipos um e dois podem ser atribuídos ao Abelisauridae, enquanto que os morfotipos três a sete pertencem a Carcharodontosauridae. Os dentes de Abelisauridae são os mais abudantes no registro fossilífero no Sítio de Peirópolis, em Minas Gerais, no Brasil. Os morfotipos oito e nove não foram identificados pois não apresentam nenhuma característica diagnóstica de terópodes conhecidos no ou na Laurásia. A presença de dentes Carcharodontosauridae, no Membro Serra da Galga (Neomaastrichtiano), é o registro geológico mais recente desta família.

Palavras-chave: dinossauros; Neocretáceo; dentes de terópodes; estado de Minas Gerais; Brasil.

INTRODUCTION Theropod teeth (carnivorous Another problem with isolated teeth is the lack ) are relatively abundant in the geological of studies that have been done on the taxonomic sig- record, but they are rarely studied and identified. The nificance of theropod dental morphology (Currie, 1987; main reason for this situation is the lack of sufficiently Farlow & Brinkman, 1987; Currie et al., 1990; Farlow diagnostic material, as usually only isolated teeth and et al., 1991). Many studies have been carried out on den- tooth fragments are recovered. Identification can be dif- tition replacement (Cooper et al., 1970; Osborn, 1971, ficult in the absence of association with more readily di- 1975; Kline & Cullum, 1984; Bolt & Demar, 1986), but agnostic cranial bones. Additionally, it is necessary to most of them focus on the relationships between diet have complete dentitions in to recognize variation and dentition (Hotton, 1955; Massare, 1987; Tanke & within the jaws (Chandler, 1990; Currie et al., 1990). Currie, 1998; Candeiro & Tanke, 2008).

1Laboratório de Geologia, Curso de Geografia,Campus Pontal, Universidade Federal de Uberlândia - UFU, Ituiutaba (MG), Brazil. E-mail: [email protected] 2Department of Biological Sciences, University of Alberta, CW405 Biological Sciences Building, Edmonton, Alberta, Canada. E-mail: [email protected] 3Laboratório de Macrofósseis, Departamento de Geologia, Universidade Federal do Rio de Janeiro - UFRJ, Rio de Janeiro (RJ), Brazil. E-mail: [email protected] *Autor correspondente

Arquivo digital disponível on-line no site www.sbgeo.org.br 323 Theropod teeth from Marília Formation at the paleontological site of Peirópolis

Theropod teeth are the most abundant cranial • CPP: Centro de Pesquisas Paleontológicas material found at the Paleontological Site of Peirópolis Llewellyn Ivor Price, Peirópolis; (Fig. 1) near Uberaba Town in Minas Gerais State, in • DG/UFRJ: Departamento de Geologia of Brazil. The exposures at the site are composed of Marília Universidade Federal do Rio de Janeiro, Rio de Formation (late Maastrichtian). Other verte- Janeiro; brates from the site include disarticulated bones from • MACN: Museo Argentino de Ciencias Naturales, fish, , lizards, crocodiliforms, titanosaurs, and Buenos Aires; theropods (Candeiro et al., 2006, 2008, Candeiro, 2007, • MCF-PVPH: Museo Municipal Carmen Funes, 2009 and references therein). However, the only crani- Paleontologia de Vertebrados, Plaza Huincul; al theropod elements found are teeth. These are labio- • UNPSJB-PV: Universidad Nacional de la lingualy compressed and have carinae with denticles. Patagonia, “San Juan Bosco”, Comodoro Previous to this study, the theropod tooth collections Rivadavia; from the Paleontological Site of Peirópolis housed in • PVL: Fundación-Instituto Miguel Lillo, Centro de Pesquisas Paleontológicas Llewellyn Ivor Tucumán; Price (Peirópolis) included 55 specimens. Kellner (1995, • MUCPV-CH: Museo de la Universidad Nacional 1996) and Candeiro et al. (2004) studied the morpho- de Comahue, El Chocon collection; logic aspects of these theropod teeth. Other studies by • RTMP, Royal Tyrrell Museum of Palaeontology, Franco & Bertini (1997) and Franco (2000) focused on Drumheller. theropod tooth . This study presents a detailed description of the teeth from Peirópolis, based on tooth MATERIAL AND METHODS Fifty-five isolated morphology. theropod teeth were micro-photographed using a scan- ning electron microscope (SEM) and a ZENIT 12 cam- Institutional abbreviations era. The following measurements were taken: tooth • AMNH: American Museum of Natural History, crown height (TCH); fore aft basal width – FABL New York; (Currie et al., 1990, Farlow et al., 1991; Sankey et al.,

0°

10°S 1N 70°W 60°W 20°S

Brazil 18° 40°W 50°W

Paranaíba River Minas Gerais Goiás State State

Grande River 20°

São Paulo State

0 100 km Marília Formation

Figure 1 – Locality and geological map of the Triângulo Mineiro region, Minas Gerais State, Brazil (modified from Fernandes & Coimbra 1996).

324 Revista Brasileira de Geociências, volume 42(2) 2012 Carlos Roberto A. Candeiro et al.

2002; Smith & Dodson, 2003); and numbers of denti- Horizon and age All the teeth are from the cles per millimeter on the proximal and distal portions Serra da Galga Member of Marília Formation – late of the anterior and posterior carina. Cross-sections of Maastrichtian (Dias-Brito et al., 2001). each specimen were traced from a copper thread, which had been wrapped around the base of the tooth crown. Description Teeth of morphotype one (Figs. 2A to The geological subdivision of the Bauru Group (Bauru D) are characterized by slightly convex labial and lingual Basin) hereafter follows Fernandes & Coimbra (1996). surfaces, which are compressed along the margins where they have contact at the carina. The denticles at the prox- SYSTEMATIC imal end of the anterior carina are smaller than those in (Marsh, 1881); the posterior carina, but the middle denticles of the ante- (Marsh, 1884); rior carina are larger than at either end. All the denticles at NEOCERATOSAURIA (Novas, 1991); the distal ends of both carinae point towards the apex of ABELISAURIA (Novas, 1992); the crown, but this is less pronounced in the proximal and (Bonaparte, 1991); middle denticles. All the denticles of both carinae are re- ABELISAURIDAE (Bonaparte & Novas, 1985). stricted to the enameled part of the crown. The denticles of the proximal and distal regions of the posterior carina Morphotype 1 are the same size, while the middle ones are larger. Referred material CPP 002, 020, 021, 121, 134, 136, 144, 150, 158, 205, 206, 207, 242, 452, 463, 477. Morphotype 2 Referred material CPP 123, 129b, 129c, 131, Locality Peirópolis site, near Uberaba Town, 132, 135, 154, 161/1, 198, 211, 372, 375/2, 446, 451/1, Minas Gerais State, Brazil. 452/1, 476, 478.

500 µm 5 mm 200 µm

500 µm

C 200 µm H A E F G B 2 mm 2 mm F I D

200 µm 200 µm 500 µm

500 µm 200 µm 200 µm P T L R J N

O 2 mm S 2 mm K 2 mm

U M Q

Figure 2 – Abelisauridae teeth. (A and C) SEM microphotograph of anterior and posterior denticles, CPP 20 (Labial); (D) cross-section of base; (B) labial view. (E and G) SEM microphotograph of posterior and anterior denticles, CPP 123 (lingual); (F) labial view; (H) cross-section of base. (I and K) SEM edge microphotograph edge with absence of denticles, CPP 129b (lingual); (F) labial view; (L) cross-section of base. (M and O) SEM of posterior and anterior denticles, CPP 131 (lingual); (N) lingual view; (P) cross-section of base. (Q) lingual view; (R) cross-section. (S and U) SEM of posterior and anterior denticles CPP 452/22 (lingual); (T) lingual view; (V) cross-section.

Revista Brasileira de Geociências, volume 42(2) 2012 325 Theropod teeth from Marília Formation at the paleontological site of Peirópolis

Description Teeth classified as morphotype two Morphotype 3 (Figs. 2E to V) are characterized by slightly convex lin- Referred material CPP 124, 129a, 208, 375/1. gual and labial surfaces, which are compressed where they meet at the posterior carina. The denticles on the Description Morphotype three (Figs. 3A to D) has proximal portion of the anterior carina are smaller than a convex lingual face in cross-section, whereas the la- those of the distal part, but denticles in the middle are bial face is almost flat. The posterior part of the crown the largest. All the denticles near the tooth tip point to- is labiolingually compressed, especially at mid-height. ward the apex of the crown. The denticles on the proxi- Enamel wrinkles are present on both faces of the crown. mal and middle parts of the anterior carina point toward The denticles on the distal portions of the carinae are to the apex of the crown. Each denticle of either carina smaller than those on the middle and proximal parts. is proximodistally longer than its basal width in labial The denticles are larger at mid-height of the crown. All or lingual views. The denticles of the proximal and dis- of the distal denticles of both carinae point toward the tal parts of the posterior carinae are the same size, and apex of the crown, while the proximal and middle den- those in the middle are larger. The cross-section (Figs. ticles are directed perpendicular to the longitudinal axis 2F, J, N, R, V) shows the anterior face compressed and of the tooth. All the anterior denticles have the same posterior face slightly convexes. heights and widths. The posterior denticles are propor- Cross-sectional shape has already been used tionally the same size in the proximal and distal regions to study theropod teeth (Bakker et al., 1988; Currie of the carina, whereas the middle ones are larger. All the et al., 1990; Farlow et al., 1991; Sankey et al., 2002), denticles of this carina are proximodistally higher than and it is useful for identifying teeth from Brazil. The their corresponding basal widths. In cross-section, the cross-sections of teeth in the jaws of gar- base of morphotype three is oval. ridoi (MCF-PVPH-236), sastrei (MACN 894), Noasaurus leali (PVL-4061), rapto- Morphotype 4 rius (AMNH 1960), and an unidentified Carnotaurinae Referred material CPP 447. (UNPSJB-PV-247) are similar to the Peirópolis teeth. Morphotypes one and two have convex lingual and la- Description Morphotype four (Figs. 3F to H) has bial faces of the crown, but are compressed along the a smoothly convex lingual face, and a strongly convex carina margins. This type of cross-section is different labial one. The anterior region islabiolingually com- from that of a carcharodontosaurid (Stromer, 1931; pressed, while the posterior part of the tooth is broadly DG/UFRJ S/N; RTMP 94.43.1, 94.43.2, 94.43.3, rounded (Fig. 3D, H). Enamel wrinkles occur on both 94.43.4, MCPV-CH-1). Abelisaurid faces of the crown. The wrinkles originate proximally (Aucasaurus garridoi, Carnotaurus sastrei and from the center of the vertical convex ridge on each side, Indosuchus raptorius) teeth are more compressed la- andxtend toward the carina. The denticles on the proxi- biolingually than those of Carcharodontosaurids mal and distal ends of the posterior carina are equiva- (Fig. 3). Dentary teeth of Carnotaurus sastrei and lent in size, while the middle ones are larger. The ante- Indosuchus raptorius (AMNH 1960) are labiolingual- rior denticles are smaller than the posterior ones. All the ly compressed and slightly recurved on the posterior denticles of both carinae point perpendicular to the lon- carina whereas in Noasaurus leali and gitudinal axis of the tooth, and they are proximodistally knopfleri (Carrano et al., 2002) they are curved more higher than basally wide. The cross-section of the tooth strongly along the posterior carina. Morphotypes one (Fig. 3H) is shallowly convex on the lingual surface and and two have similar characteristics to those in the strongly convex on the labial face. It is compressed to- jaws of Carnotaurus sastrei and Indosuchus raptori- wards the front and thick posteriorly so that the cross- us – both have slightly convex lingual and labial faces section looks roughly like an isosceles triangle. and compressed edges. THEROPODA (Marsh, 1881) Morphotype 5 (Gauthier, 1986) Referred material CPP 156. NEOTETANURAE (Sereno, 1994) (Currie and Zhao, 1993) Description Morphotype five (Figs. 3I to L) is pe- cf. CARCHARODONTOSAURIDAE (Stromer, culiar in that the lingual face is smoothly convex where- 1931) as the labial face is strongly convex. The posterior part

326 Revista Brasileira de Geociências, volume 42(2) 2012 Carlos Roberto A. Candeiro et al.

200 µm

200 µm 200 µm G 200 µm

B C F E 2 mm A 2 mm H D

200 µm 200 µm

200 µm 7 mm 200 µm O N K J

I 2 mm M L P

200 µm 7 mm 200 µm

5 mm S 2 mm 200 µm R

200 µm W Y

X T 10 mm Q

V U

Z

Figure 3 – Carcharodontosauridae teeth. (A and C) SEM microphotograph of anterior and posterior denticles, CPP 124; (B) labial view; (D) cross-section of base. (E and G) SEM microphotograph of anterior and posterior denticles, CPP 157; (F) lingual view; (H) cross-section of base. (J and K) SEM of anterior and posterior denticles, CPP 200; (I) lingual view; (L) cross-section of base. (M) lingual view, (N and O) SEM of anterior and posterior denticles, CPP 216; (P) cross-section of base. (Q) labial view, (R and S) SEM of anterior and posterior denticles, CPP 448; (T) wrinkles on posterior edge; (U) cross-section of base. (V) lingual view, (W and X) SEM of anterior and posterior denticles CPP 474, (Y) wrinkles on distal region; (Z) cross-section of base. of the cross-sectional outline of the tooth is somewhat middle portions of the carina point perpendicular to the compressed. The tooth has a double posterior carina, longitudinal axis of the tooth. The anterior denticles are with both parts converging toward the tip of the tooth. smaller at the proximal end of the carina, and the middle Smooth enamel wrinkles occur on both faces of the denticles are the largest. The distal denticles of both cari- crown. They originate on the surface of the vertical con- nae point toward the apex of the crown. The denticles on vex ridge on each of the lingual and labial surfaces, and the proximal and middle portions of the anterior carina they extend towards the anterior and posterior carinae. point away from the longitudinal axis of the crown. The The paired posterior carina is formed by a large, al- denticles are basally wider than proximodistally high most straight component, and a smaller secondary part on the anterior carina, while they are higher than wide (Fig. 3J). The anterior denticles on the proximal and posteriorly. The posterior denticles are larger at the mid-

Revista Brasileira de Geociências, volume 42(2) 2012 327 Theropod teeth from Marília Formation at the paleontological site of Peirópolis height of the crown. The cross-section (Fig. 3L) is ante- roseae) by many authors (Larsson, 1996; Sereno et al., riorly compressed, and is posteriorly thicker. 1996; Vickers-Rich et al., 1999). In Giganotosaurus carolinii, different forms of wrinkles occur (Calvo Morphotype 6 pers. com), which can be strong or shallow in re- Referred material CPP 152, 199, 216, and 376. lief. These features may also extend close to the cari- nae (e.g., and Giganotosaurus), Description Morphotype six (Figs. 3M to P) is which are also characteristic of morphotypes three to characterized in cross-section by having a shallow- seven. Morphotypes three, four, six, and seven have ly convex labial and a strongly convex lingual surface compressed cross-sections, giving each tooth a knife- (Fig. 3N). The anterior part of the tooth is labiolingualy like appearance as in Carcharodontosaurus saharicus compressed, whereas the posterior edge is thicker in the (Stromer, 1931) and Giganotosaurus (MUCPV-CH-1). cross-section. Enamel wrinkles are similar to those of THEROPODA (Marsh, 1881) morphotypes three, four, and five. Denticles on the prox- imal and distal portions of each carina are of equal size, Morphotype 8 while the middle ones are larger. All anterior denticles Referred material CPP 128, 243, 271, and 371. are slightly larger than the posterior ones. On both cari- nae, the denticles point toward the apex of the crown. Description A morphotype eight (Figs. 4A to B) is The denticles of the proximal and mid-height portions characterized by a shallowly convex lingual face and a of the anterior carina are perpendicularly to the longi- more strongly convex labial surface in cross-section. tudinal axis of the tooth. All of the denticles are prox- The anterior and posterior margins of the tooth are rela- imodistally higher than basally wide on both carinae. tively thick in cross-section. On the anterior carina, the Slender blood grooves originate between adjacent den- proximal denticles are smaller than the distal ones, and ticles. The cross-section is relatively convex on the lin- the mid-height denticles are larger than both proximal gual side, but it is broadly expanded between the poste- and distal denticles. The distal denticles on both cari- rior carinae. nae pointed toward the apex of the crown. The denticles on the proximal and mid-height portions of the anterior Morphotype 7 carina are perpendicular to the longitudinal axis of the Referred material CPP127, 197, 200, 241, 448, tooth. All denticles are as high as they are wide on both 449, 474, 475. carinae. Slender blood grooves are present. The poste- rior carina forms an angle of almost 90º with the crown Description Morphotype seven (Figs. 3Q to Z) teeth base. The denticles of the proximal and distal re- have shallowly convex lingual and labial surfaces, and gions of the posterior carina are equal; however, they are labiolingually narrow at the anterior and posterior are smaller than the mid-height denticles. The denticles carinae. Enamel wrinkles (Figs. 3T and Y) are present on the proximal and mid-height regions of the posterior on both labial and lingual faces of the crown, and mor- carina are perpendicular to the longitudinal axis of the phologically are similar to those of morphotypes three tooth. The cross-section (Fig. 4B) has all the sides with to six. Denticles from the proximal portions of the cari- concave or oval shapes. nae are smaller than the distal ones, and both are small- er than the middle ones. Denticles of both carinae point Morphotype 9 toward the apex of the crown. The denticles from the Referred material CPP 374. proximal and mid-height areas of the anterior carina are perpendicular to the longitudinal axis of the tooth. All Description Morphotype nine (Figs. 4C to F) has a denticles are proximodistally higher than they are basal- relatively flat lingual surface and a more strongly con- ly wide. The lingual and labial surfaces of the tooth are vex labial face. The anterior part of the tooth base is la- shallowly convex in cross-section, and the tooth is nar- biolingually compressed and the posterior part is thicker row and bladelike. and has a more strongly convex profile in section. The presence of wrinkles on a theropod tooth The proximal and distal denticles are subequal crown has been considered a diagnostic feature for in size, although all anterior denticles are slightly Carcharodontosauridae (e.g., Carcharodontosaurus larger than the equivalent posterior ones. Both ante- saharicus, Giganotosaurus carolinii and rior and posterior denticles near the tip of the tooth

328 Revista Brasileira de Geociências, volume 42(2) 2012 Carlos Roberto A. Candeiro et al. point toward the apex of the crown. More proximal 10 mm denticles are more or less perpendicular to the longi- 500 µm tudinal axis of the tooth. 500 µm E CONCLUSIONS Morphotypes eight and nine C do not have any of the characteristics (cross-section, A D presence of wrinkles, and tooth shape) typical of known 2 mm Gondwanan theropods, nor do they resemble the teeth B F of Laurasian theropods. Gondwanan theropods are rep- resented mainly by Abelisauroidea, Alvarezsauridae, Figure 4 – Theropoda indet. teeth. (A) lingual Carcharodontosauridae, , and view, CPP 271; (B) cross-section of base. (C and . Amongst South-American taxa, teeth E) SEM microphotograph of posterior and anterior are not known for exemple Alvarezsaurus calvoi, denticles, CPP 374 (lingual); (D) lingual view; (F) Neuquenornis volans, Yungavolucris brevipedalis, cross-section of base. Lectavis brelicola, Patagopteryx deferasi, Patagonykus puertai, namuhaiquii, challengeri, work was supported by a CAPES Fellowship. A assmetrica, and placidus, nor Foundation Grant made possible part of this study at the are they known for lapparenti of . Royal Tyrrell Museum of Palaeontology in Drumheller, Morphotypes nine and ten might represent any of these Alberta, Canada. The authors were given access by Luíz Gondwanan taxa, or might represent new taxa. Carlos Borges Ribeiro to the theropod teeth at the Centro Thus, in the present work theropod teeth re- de Pesquisas Paleontológicas Llewellyn Ivor Price. The cords were reported. They are from the Peirópolis Site, authors would like to thank Drs. Mark Norell and Chris in Minas Gerais State, a place that has yielded a great Collins (American Museum of Natural History); José number of bones of titanosaurid sauropods (Baurutitan F. Bonaparte (Museo Argentino de Ciencias Naturales); britoi and Uberabatitan riberoi) and crocodylo- Alexander A. Kellner (Museu Nacional); and Darren morphs ( jesuoinoi, tormini and Tanken, Eva B. Koppelhus and D. Brinkman (Royal terrificus). In contrast, to the theropo- Tyrrell Museum of Palaeontology) for the assistance in da record is only represented by teeth (this work), an studying collections in their institutions. Abelisauridae (Novas et al., 2008), a tetanuran The SEM work was done by Rogério Martins scapula (Machado et al., 2008), and a maniraptoran (Centro de Pesquisas da Petrobrás). We also thank claw (Novas et al., 2005). The present Abelisauridae Drs Farlow (University of Indiana), Peter Dodson and Carcharodontosauridae reports from Peirópolis in- (University of Pennsylvania), Kenneth Carpenter dicates that this group has a larger geographical distri- (Denver Museum of Natural History Denver, Colorado), bution in the Gondwana landmasses, which is the most and Ismar Carvalho (Universidade Federal do Rio de extensive theropod tooth assemblage from Brazil. Janeiro) for supplying references to the senior author during the development of this thesis. R. Candeiro and ACKNOWLEDGMENTS This work is a revision L. Berqgvist were supported by Conselho Nacional de of a Master’s thesis presented to the Departamento de Desenvolvimento Científico e Tecnológico (CNPq)/ Geologia, Universidade Federal do Rio de Janeiro. The Bolsista de Produtividade em Pesquisa fellow.

References

Bakker R.T., Williams M., Currie P.J. 1988. Nanotyrannus Candeiro C.R.A. 2007. Cretaceous biota of the Triângulo a new of pygmy tyrannosaur, from latest Mineiro region: A review of recent finds. Estudios Cretaceous of Montana. Hunteria, 1:1-30. Geológicos, 63:65-73. Bolt J.R. & Demar R.E. 1986. Computer simulation Candeiro C.R.A. & Tanke D. 2008. A pathological Late of tooth replacement with grow in lower tetrapods. Cretaceous carcharodontosaurid tooth from Minas Journal of Vertebrate Paleontology, 6:233-250. Gerais, Brazil. Bulletin of Geosciences, 83:351-354. Candeiro C.R.A. 2009. Vertebrates of the Marília Candeiro C.R.A., Bergqvist L.P., Novas F.E., Currie P.J. Formation (late Maastrichtian) from the Peirópolis 2004. Theropod teeth from the Marília Formation Paleontological Site: toward a better understanding. (Upper Maastrichtian), Minas Gerais State, Brazil. Earth Sciences Research Journal, 13:6-15. Journal of Vertebrate Paleontology, 24:43A-43A. Revista Brasileira de Geociências, volume 42(2) 2012 329 Theropod teeth from Marília Formation at the paleontological site of Peirópolis

Candeiro C.R.A., Martinelli A.G., Avilla L.S., Rich T. ______. 1995. Theropod teeth from the 2006. Tetrapods from the Upper Cretaceous ( Bauru Group near Peirópolis, Minas Gerais, Brazil. XIV Maastrichtian) Bauru Group of Brazil: a reappraisal. Congresso Brasileiro de Paleontologia, Atas 66-67. Cretaceous Research, 27:923-946. Kline L.W., Cullum A. 1984. A long term study of the tooth Candeiro C.R.A, Santos A.R., Bergqvist L.P., Ribeiro replacement phenomenon in the young green iguana, L.C. 2008. The Late Cretaceous fauna and flora of the Iguana iguana. Journal of Herpetology, 18:176-185. Uberaba area (Minas Gerais State, Brazil). Journal of Larsson H.C.E. 1996. Cranial morphology of the South American Earth Sciences, 25:203-216. African theropod, Carcharorontosaurus saharicus Carrano, M.T., Sampson, S.D., Forster, C.A. 2002. (Allosauroidea). Journal of Vertebrate Paleontology, The osteology of Masiakasaurus knopfleri, a small 16:3 (Abstracts). abelisauroid (Dinosauria: Theropoda) from the Late Machado E., Campos D.A., Kellner A.W.A. 2008. On a Cretaceous of . Journal of Vertebrate theropod scapula (Upper Cretaceous) from the Marília Paleontology, 22(3):510-534. Formation, Bauru Group, Brazil. Palaeontologische Chandler C.L. 1990. Taxonomic and functional significance Zeitschrift, 82:308-313. of serrated tooth morphology. In: Theropod Dinosaurs. Massare J.A. 1987. Tooth morphology and prey reference Unpublished Master Thesis, Yale University, New of marine . Journal of Vertebrate Haven, 163 p. Paleontology, 7(2):121-137. Cooper J.S., Poole D.F.G., Lawson R. 1970. The dentition Novas F.E., Ribeiro L.C.B., Carvalho I.S. 2005. of iguanid lizards with special reference to tooth Maniraptoran theropod ungual from the Marília replacement. Journal of , 162:85-98. Formation (Upper Cretaceous), Brazil. Revista del Currie P.J. 1987. -like characteristics of the jaws and Museo Argentino de Ciencias Naturales, 7:31-36. teeth of troodontid theropods (Dinosauria, ). Novas F.E., Carvalho I.S., Ribeiro L.C.B, Mendez A.H. Journal of Vertebrate Paleontology, 7:72-81. First abelisaurid bone remains from the Maastrichtian Currie P.J., Rigby J., Sloan R.E. 1990. Theropod teeth Marília Formation, Bauru Basin, Brazil. Cretaceous from the Judith River Formation of southern Alberta, Research, 29:625-635. Canada. In: Carpenter K. & Currie P. J. (eds.). Dinosaur Osborn J.W. 1975. Tooth replacement: efficiency, patterns, Systematics: Approaches and Perspectives, Cambridge and . Evolution, 29:180-186. University Press, New York, p. 107-125. Osborn J.W. 1971. The ontogeny of tooth succession in Dias-Brito D., Musacchio E.A., Castro J.C., Maranhão M.S.A., Lacerta vivipara Jacuin 1787. Proceedings Royal Suarez J.M., Rodrigues R. 2001. Grupo Bauru: uma Society of London, Serie B, 179:261-289. unidade continental do Cretáceo do Brasil – concepções Sankey J.T., Brinkman D.B., Guenther F., Currie P.J. baseadas em dados micropaleontológicos, isotópicos e 2002. Small theropod and bird teeth from the Late estratigráficos.Revue Paléobiologie, 20:245-304. Cretaceous (Late ) Judith Rivert Group, Farlow J.O. & Brinkman D.L. 1987. Serration and patterns Alberta. Journal of Paleontology, 76(4):751-763. of wear dinosaur teeth. 21st Annual Meeting, South- Sereno P.C., Dutheil D.B., Iarochene M., Larsson H.C.E., Central Section, Geological Society of America. Waco. Lyon G.H., Magwenw P. et al. 1996. Predatory Texas, Abstracts with Program, 151. dinosaurs from the Sahara and Late Cretaceous faunal Farlow J.O., Brinkman O.L., Abler W.L., Currie P.J. 1991. differentiation. Science, 272:986-991. Size, shape and serration density of theropod dinosaur Smith J.B. & Dodson P. 2003. A proposal for a standard lateral teeth. Modern Geology, 16:161-198. terminology of anatomical notation and orientation Fernandes L.A. & Coimbra A.M., 1996. A Bacia Bauru in fossil vertebrate dentitions. Journal of Vertebrate (Cretáceo Superior, Brasil). Anais da Academia Paleontology, 23(1):1-12. Brasileira de Ciências, 68(2):95-105. Stromer E. 1931. Ein skellett-Rest von Carcharodontosaurus. Franco A.C. 2000. Dentes teropodomorfos do Cretáceo Abhandlungen der Bayerischen Akademie der Superior da Bacia do Paraná. Análise em microscopia Wissenschaften, 9:1-23. eletrônica de varredura. Unpublished Master Thesis, Tanke D.H. & Currie P.J. 1998. Head-biting behavior in Universidade Estadual Paulista/UNESP-Rio Claro, 109 p. theropod dinosaurs: Paleopathological evidence. Gaia, Franco A.A. & Bertini R.J. 1997. Structure and ultra- 15:167-184. structure of the dental morphology in theropodomorphs Vickers-Rich P., Rich T.H., Lanus D.R., Rich L.S.V., Vacca dinosaurs (Sarischia: Reptilia). Ulility to taxonomy. XVI R. 1999. “Big tooth” from the of Chubut Meeting of Brazilian Society for Electron microscopy. Province, Patagonia: A possible carcharodontosaurid. p. Acta Microcopica, 6:688. 85-88 Proceedings of the Second Gondwanan Dinosaur Hotton N. 1955. A survey of adaptive relationships of Symposium, edited by Tomida Y, Rich T.H., Vickers- dentition to diet in the North American Iguanidae. Rich P., National Science Museum Monographs. American Midle Nature, 53:88-114. Kellner A.W.A. 1996. Remarke on Brazilian dinosaurs. Manuscrito ID 21309 Memoirs of the Queensland Museum, 39:611-626. Recebido em: 25/04/2011 Aprovado em: 07/03/2012

330 Revista Brasileira de Geociências, volume 42(2) 2012

Top View