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Biostratigraphic Reappraisal of the Lower Triassic Sanga Do Cabral

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Biostratigraphic Reappraisal of the Lower Triassic Sanga Do Cabral Journal of South American Earth Sciences 79 (2017) 281e296 Contents lists available at ScienceDirect Journal of South American Earth Sciences journal homepage: www.elsevier.com/locate/jsames Biostratigraphic reappraisal of the Lower Triassic Sanga do Cabral Supersequence from South America, with a description of new material attributable to the parareptile genus Procolophon * Sergio Dias-da-Silva a, , Felipe L. Pinheiro b, Atila Augusto Stock Da-Rosa c, Agustín G. Martinelli d, Cesar L. Schultz d, Eduardo Silva-Neves a, e, Sean P. Modesto f a Centro de Apoio a Pesquisa Paleontologica da Quarta Colonia,^ Universidade Federal de Santa Maria, Rua Maximiliano Vizotto, 598, Sao~ Joao~ do Pol^esine, Rio Grande do Sul, CEP: 97-230-000, Brazil b Laboratorio de Paleobiologia, Universidade Federal do Pampa, Av. Antonio Trilha, 1847, Sao~ Gabriel, Rio Grande do Sul, Brazil c Laboratorio de Estratigrafia e Paleobiologia, Departamento de Geoci^encias, Centro de Ci^encias Naturais e Exatas, Universidade Federal de Santa Maria, Santa Maria, RS, CEP: 97.105-900, Brazil d Departamento de Paleontologia e Estratigrafia, Universidade Federal do Rio Grande do Sul, Porto Alegre, Rio Grande do Sul, CEP: 91540-000, Brazil e Programa de Pos-Graduaç ao~ em Biodiversidade Animal, Universidade Federal de Santa Maria, RS, CEP: 97.105-900, Brazil f Department of Biology, Cape Breton University, Sydney, Nova Scotia, B1P 6L2, Canada article info abstract Article history: The Sanga do Cabral Supersequence (SCS), comprises the Brazilian Sanga do Cabral Formation (SCF) and Received 8 June 2017 the Uruguayan Buena Vista Formation (BVF). So far, the SCS has yielded temnospondyls, parareptiles, Received in revised form archosauromorphs, putative synapsids, and a number of indeterminate specimens. In the absence of 20 July 2017 absolute dates for these rocks, a biostratigraphic approach is necessary to establish the ages of the SCF Accepted 26 July 2017 and the BVF. It is well established that the SCF is Early Triassic mainly due to the presence of the Available online 29 July 2017 widespread Gondwanan reptile Procolophon trigoniceps. Conversely, the age of the BVF is subject of great controversy, being regarded alternatively as Permian, Permo-Triassic, and Early Triassic. The BVF has Keywords: fi Gondwana yielded the de nite procolophonid Pintosaurus magnidentis. Procolophonoidea is one of the most diverse Sanga do Cabral Supersequence and conspicuous terrestrial tetrapod groups of the Lower Triassic Lystrosaurus Assemblage Zone in the Triassic Karoo Basin of South Africa, which preserves tetrapods from the aftermath of the end-Permian extinction Temnospondyli event. Based on a previous interpretation that the fauna of the BVF is Permian, and in the reinterpretation Procolophonidae of disarticulated vertebrae from SCF with ‘swollen’ neural arches as belonging to either seymour- Archosauromorpha iamorphs or diadectomorphs, it was recently suggested that at least part of the SCF is Permian in age, which prompted this comprehensive reevaluation of both SCS's faunal content and geology. Moreoever, new, strikingly large procolophonid specimens (skull, vertebra, and a mandibular fragment) from the SCF are described and referred to the genus Procolophon. The large procolophonid vertebra described here contradicts the recent hypothesis that similar specimens from the SCF belong to seymouriamorphs or diadectomorphs, because its morphology is consistent with that found in Procolophon. There is not a single diagnostic specimen that supports the inference of Permian levels in the SCS. Accordingly, because all diagnostic and biostratigraphically informative fossils from the SCF and the BVF are either Early Triassic or restricted to the Triassic, we conclude that the available biostratigraphic data reinforce an Early Triassic age assignment to the SCS. © 2017 Elsevier Ltd. All rights reserved. 1. Introduction Uruguayan Buena Vista Formation (BVF), is crucial to the under- standing of the biotic recovery that followed the greatest biological The Western Gondwanan Sanga do Cabral Supersequence (SCS), crisis in the history of Earth, the end-Permian extinction event. The including the Brazilian Sanga do Cabral Formation (SCF) and the SCF is traditionally considered Early Triassic, an age assignment mainly due to the presence of the widespread Gondwanan reptile * Corresponding author. Procolophon trigoniceps (Dias-da-Silva et al., 2006a; Cisneros, E-mail address: [email protected] (S. Dias-da-Silva). http://dx.doi.org/10.1016/j.jsames.2017.07.012 0895-9811/© 2017 Elsevier Ltd. All rights reserved. 282 S. Dias-da-Silva et al. / Journal of South American Earth Sciences 79 (2017) 281e296 2008a). The age of the BVF, however, has been subject of great Federal do Pampa (UNIPAMPA), Sao~ Gabriel, RS, Brazil; UFSM, controversy because it has been alternatively regarded as Permian, Laboratorio de Estratigrafia e Paleobiologia, Universidade Federal Permo-Triassic, and Early Triassic (Pineiro~ et al., 2003, 2004, 2007a, de Santa Maria (UFSM), Santa Maria, RS, Brazil. b, c; Dias-da-Silva et al., 2006a; Modesto and Botha-Brink, 2010). In line with a previous interpretation that the fauna of the BVF is 2. Geology and biostratigraphic reappraisal Permian, together with a reinterpretation of disarticulated verte- brae from SCF with ‘swollen’ neural arches as attributable to either 2.1. Geology and geochronology seymouriamorphs or diadectomorphs, it was suggested recently that at least part of the SCF is Permian in age (Pineiro~ et al., 2015). If The Sanga do Cabral Supersequence (SCS) comprises the Bra- correct, it implies that the Permo-Triassic boundary lies within the zilian Sanga do Cabral Formation (SCF) and the Uruguayan Buena SCF, which would make the SCF fauna critical to analyses of Vista Formation (BVF) (Zerfass et al., 2003). Based upon its tetrapod tetrapod survivorship of the end-Permian extinction event. The content, the SCF has long been regarded as Early Triassic, and hypothesis that part of the SCF is Permian, which contradicts all correlated with the Lower Triassic Katberg Formation of the Karoo previous work supporting an Early Triassic age for that formation, Basin in South Africa (for a complete up-to-date set of SCS's pub- has prompted our comprehensive reevaluation of both SCS's faunal lished vertebrate specimens, see Table 1). Together with other content and geology, in order to provide a biostratigraphic reas- Lower Triassic units (Fig. 1), this Western Gondwanan super- sessment for this supersequence. sequence crucially documents biotic recovery following the end- Whereas we re-examine all biostratigraphic evidence for the age Permian Extinction Event, which is traditionally regarded as the of the SCS, procolophonoid fossils remain central to any biostrati- most severe of the ‘Big Five’ mass extinctions. Accordingly, the SCS graphic assessment of this supersequence. Procolophonoidea is the helps to understand the decisive worldwide biotic turnover from only parareptile clade that survived the end-Permian extinction Paleozoic to Mesozoic eras (see Benton et al., 2004; Smith and event (Modesto et al., 2001, 2003; Modesto et al., 2010) and became Botha, 2005; Botha and Smith, 2006; Sahney and Benton, 2008). one of the most diverse and conspicuous group of tetrapods of the The SCF was proposed by Andreis et al. (1980) for rocks cropping out Lower Triassic Lystrosaurus Assemblage Zone in the Karoo Basin of around the city of Rio Pardo, which is located centrally in Rio Grande South Africa (Smith and Botha, 2005; Botha and Smith, 2006; Botha do Sul State (RS) (Fig. 2). There, orange-coloured, fine-grained et al., 2007; Macdougall and Modesto, 2011). In South America, sandstones with localized intraformational conglomerates occur procolophonoids are known from both the Sanga do Cabral and over the pinkish to white, fine sandstones of the Rio do Rasto Buena Vista formations (see Zerfass et al., 2003). The Brazilian Formation. The conglomerates represent shallow braided streams, material comprises several specimens, including fairly complete in an approximate 1:100 thickness/width scale, whereas the orange skulls (e.g. Lavina, 1983; Langer and Lavina, 2000; Cisneros and fine sandstones are interpreted as massive, or presenting horizontal Schultz, 2002; Dias-da-Silva et al., 2006a). Originally ascribed to stratification, and denote a broad semiarid plain. The intraforma- two species, Procolophon pricei (Holotype UFRGS-PV231T; Lavina, tional conglomerates are the source of the majority of vertebrate 1983) and P. brasiliensis (Holotype MCN-PV-1905; Cisneros and fossils, which consist mainly of isolated, fragmentary bones. Schultz, 2002), both have been synonymized under P. trigoniceps Vertebrate remains are rarely preserved in the sandstones. The (Cisneros, 2008a). Although those best preserved and most diag- most commonly represented groups are procolophonids and tem- nostic specimens (i.e. skull and lower jaws) are unequivocally nospondyls (see discussion on the fossiliferous content, below). The referred to P. trigoniceps, the real diversity of procolophonians in the SCF occurs in an EeW outcrop belt in RS, with many fossiliferous SCS is unknown because several other specimens are fragmentary outcrops (Lavina, 1983; Lavina and Barberena, 1985; Da-Rosa et al., and poorly preserved (Langer and Lavina, 2000). Procolophonoid 2009; Dias-da-Silva and Da-Rosa, 2011), and have a probable material from the BVF so far comprises a single specimen of the physical continuation in Uruguay (Andreis
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