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Paleogeographic Distribution of the Terrestrial Squamate Reptiles from the Cretaceous of Brazil

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Paleogeographic Distribution of the Terrestrial Squamate Reptiles from the Cretaceous of Brazil 65 Review Article PALEOGEOGRAPHIC DISTRIBUTION OF THE TERRESTRIAL SQUAMATE REPTILES FROM THE CRETACEOUS OF BRAZIL DISTRIBUIÇÃO PALEOGEOGRÁFICA DOS RÉPTEIS ESQUAMATAS TERRESTRES DO CRETÁCEO DO BRASIL Carlos Roberto A. CANDEIRO 1 1. Paleontologist, doctorate student at Departamento de Geologia, Federal University of Rio de Janeiro. [email protected] ABSTRACT: The terrestrial squamates from Brazil are the Early Cretaceous Olindalacerta brasiliensis and Tijubina pontei from Ceará State; the Late Cretaceous Pristiguana brasiliensis from Minas Gerais State, Ophidia indet. and Aniloidea from São Paulo State. Squamates were widespread in western Gondwana and Laurasia landmasses, and iguanids and mainly snakes widely distributed in southern South America. KEYWORDS: Squamata. Paleogeographical distribution. Cretaceous. Brazil. INTRODUCTION paleogeographical and chronostratigraphical distribution of this group in Brazil during the The Early Cretaceous terrestrial basal Cretaceous. squamate species Tijubina pontei and Olindalacerta brasiliensis from the Crato METHODOLOGY Formation in Araripe Basin (Ceará State, Brazil) are among the earliest known squamate lepidosaurs The data on Squamata taxa for the present from South America. Accordingly, their review are mostly based in literature sources as paleogeographic position in Gondwana during the well as in the direct observation of the specimens. Early Cretaceous indicates a widespread Taxonomical treatment - In order to evaluate the Gondwanan distribution of squamates. Their taxonomic status of Squamata, I use information phylogenetic position places them at or near the obtained from Reynoso; Callison (2000). ancestry of squamates (BONFIM; MARQUES, Institutional abbreviations : Brazil - DNPM, 1997; EVANS; YABUMOTO, 1998). The Late Departamento Nacional da Produção Mineral, Rio Cretaceous squamate record consists only of de Janeiro, Rio de Janeiro State; MPSC, Museu de Pristiguana brasiliensis from late Mastrichtian Paleontologia de Santana, Universidade Regional Marília Formation of Minas Gerais State, which do Cariri, Ceará State. Japan : KMNH, Kitakyushu has been regarded as a possible iguanid by Museum and Institute of Natural History, Fukuoka, Apesteguía; Agnolin; Lio (2005), Estes (1983), Kyushu, Japan Province. Estes; Price (1973) and Candeiro et al. (2007). Geology - For the stratigraphic units in the Araripe Recent studies have reported an expanded Basin I follow the arrangement proposed by Martill and considerably detailed record of Late (1993) and for the Bauru Basin used Fernandes; Cretaceous continental squamate discoveries Coimbra (1996). (APESTEGUÍA, 2005; APESTEGUÍA; AGNOLIN; LIO, 2004, 2005; MARTINELLI; SQUAMATA BACKGROUND FORASIEPI, 2004). The recently reported Ophidia The current record of squamates in the from Turonian-Santonian Adamantina Formation Cretaceous beds of Brazil is extremely limited: a of the São Paulo state (BERTINI, 1994). Other, few lizards and possibly two snakes. The first more recently reported (ZAHER et al., 2003) formally described Cretaceous lizard from this squamate Anguimorpha from Adamantina Brazil was Pristiguana brasiliensis from the Bauru Formation, São Paulo State, confirm serpent Group (ESTES; PRICE, 1973). In addition, a well- paleogeographic distribution in Brazil. This preserved basal squamate was described from the contribution reviews the paleogeographic position Crato Formation (Ceará State; BONFIM; of squamate Brazilian Cretaceous records and the MARQUES, 1997; EVANS; YAMUBOTO, 1998). remarks of this group in Brazil. Unstudied materials of lizards and serpents are In this contribution, I present an up-to-date known from the Adamantina Formation of São list of Squamata taxa recognized in the Cretaceous Paulo State (BONAPARTE, 1978; BERTINI; of Brazil. Furthermore, I evaluate the BONFIM, 1998; ESTES; PRICE, 1973). Biosci. J., Uberlândia, v. 23, n. 1, p. 65-74, Jan./Mar. 2007 Received: 08/006/06 Accepted: 27/10/06 66 Paleogeographic distribution… CANDEIRO, C. R. A. correlations. This age assignment is supported by GEOLOGICAL SETTING stratigraphic studies (DE LIMA, 1978; DOYLE, Ceará State 1992; PONS et al., 1990). Araripe Group, Crato Formation The Araripe Group (Crato, Ipubi, Minas Gerais and São Paulo States Batateiras, and Santana formations; Araripe Basin) Bauru Group (Adamantina and Marília is located in the west-central region of Brazil formations) (Figure 1), covering Ceará and Piauí states. This Within the Bauru Basin, is the Bauru group is an important paleontological area of Group (Figure 1), the most extensively studied unit. Gondwana landmass due to its richness of The Bauru Group is composed of the Adamantina Cretaceous vertebrate faunas. It is composed of (Turonian-Santonian), Uberaba (Turonian- intercalating marine, littoral and continental Coniacian) and Marília (late Maastrichtian) sediments (NEUMANN et al., 2003). formations. The Caiuá Group is composed by the Within the Araripe Group, the fossils of Rio Paraná (Santonian-Maastrichtian), Goio Êre Tijubina pontei (BONFIM; MARQUES, 1997) and (Santonian-Maastrichtian), and Santo Anastácio Olindalacerta brasiliensis (EVANS; (Aptian-Cenomanian) formations. Dias-Brito et al. YABUMOTO, 1998) were collected in the Crato (2001) discussed the age of Santo Anastácio, Formation (Figure 1). Adamantina, Uberaba, and Marília formations, The Crato Formation was previously Fernandes (1998) the for Goio Êre and Rio Paraná included in the Santana Formation (MARTILL, formations. The Bauru Group has a complex 1993). The Santana Formation was subsequently nomenclatural history (BARCELOS, 1984; subdivided into the three members Crato, Ipubi, FERNANDES; COIMBRA, 1996), partially due to and Romualdo (BERTHOU, 1990), later elevated the artificial division of the Upper Cretaceous strata to formations rank: the Crato, Ipubi and Santana in the flat areas of the Goiás, Mato Grosso do Sul, Formations (the later being equivalent to the Minas Gerais (“Triângulo Mineiro”), Paraná and Romualdo Member) (MARTILL, 1993). Together São Paulo states. with the Exu Formation, which consists mainly of Both the Adamantina and Marília coarse clastic sediments, these formations were formations are chronologically correlated to the included in the Araripe Group (MARTILL, 1993). Upper Cretaceous strata from Argentina (HUENE, The Crato Formation is the most basal of these 1927, 1939). Within the Paraná Basin, the formations and consists of organic rich mudstones Adamantina and Marília formations have been and laminated micritic plattenkalk limestones described as Grez (GONZAGA DE CAMPOS, possibly of marginally marine origin (BECHLY, 1905), Series (FREITAS, 1955), Formation (ARID, 1998). The limestone sequence is about 30 m in 1967; DIAS-BRITO et al., 2001; MEZZALIRA, thickness, showing considerable variations in the 1974) and Group (SOARES et al., 1980; SUGUIO, underlying topography (MARTILL, 1993). The age 1980). Other workers subdivided further these of the sedimentary sequence of the Araripe Group formations into more units (e.g., FERNANDES, is uncertain due to lack of good stratigraphic 1998). marker fossils and lack of other geological The recognition and description of the evidence. It is generally cited as Early to mid- stratigraphical relationships among the various Cretaceous (Aptian, and/or Albian, and perhaps formations of the Bauru Basin allowed to better Cenomanian) (MARTILL, 1993). Berthou (1990) understanding of the extensive fossil collections in suggested a late Aptian to early Albian age based a chronostratigraphic context. on sedimentological and palaeontological Biosci. J., Uberlândia, v. 23, n. 1, p. 65-74, Jan./Mar. 2007 67 Paleogeographic distribution… CANDEIRO, C. R. A. Figure 1. Distribution of Cretaceous continental Araripe and Bauru basins in Brazil (Adapted from MARTILL, 1993 and FERNANDES; COIMBRA, 1996. PHYLOGENY OF SQUAMATA Locality, geology, and age . Nova Olinda municipality, Ceará State, Brazil, Crato Formation, The phylogenetic relationships of the Aptian/Albian age. Cretaceous squamates of Brazil are subjects of Remarks: The remains of the primitive squamate considerable interest. Recent studies on lepidosaur Tijubina pontei from Early Cretaceous sediments relationships have focused on traditional was briefly described by Bonfim; Marques (1997) osteological comparisons (e.g. ALBINO, 1996; who referred it as a possible member of Teiidae. BONAPARTE, 1996) and cladistic analyses (e.g. More recently, Bonfim (2001) and Bonfim; Avilla CALDWELL, 1999; ESTES; DE QUEIROZ; (2002) suggested Tijubina represents a basal GAUTHIER, 1988; EVANS, 2003; ESTES, 1983; squamate closely related to Huehuecuetzpalli LEE; 1998; REYNOSO; 1998; REYNOSO; mixtectus Reynoso, 1998 from the Aptian of CALLISON, 2000). The Brazilian Tijubina and Mexico. The results of a cladistic analysis by Olindalacerta and the Early Cretaceous Bonfim; Marques (op. cit) places Tijubina and Huehuecuetzpalli from Mexico all retain certain Huehucuetzpalli in a basal position within osteological features that indicate relationship with Squamata where they comprise a sister clade to the Lepidosauria, with which they were crown-group squamates. contemporaneous. The other Early Cretaceous Olindalacerta brasiliensis Evans; Yabumoto, 1998 basal squamate (EVANS; 2003) are not known well Holotype : KMNH VP 400, 001 an articulated enough to afford confident comparisons. specimen. Recent taxonomic assignments and Locality, geology and age : Ceará State, the precise cladistic analyses have consistently recognized provenance-can you
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