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Pterosauria, Pterodactyloidea, Tapejaridae) Historical Biology An International Journal of Paleobiology ISSN: 0891-2963 (Print) 1029-2381 (Online) Journal homepage: http://www.tandfonline.com/loi/ghbi20 New information on the postcranial skeleton of the Thalassodrominae (Pterosauria, Pterodactyloidea, Tapejaridae) Richard Buchmann, Taissa Rodrigues, Sabrina Polegario & Alexander W. A. Kellner To cite this article: Richard Buchmann, Taissa Rodrigues, Sabrina Polegario & Alexander W. A. Kellner (2018) New information on the postcranial skeleton of the Thalassodrominae (Pterosauria, Pterodactyloidea, Tapejaridae), Historical Biology, 30:8, 1139-1149, DOI: 10.1080/08912963.2017.1343314 To link to this article: https://doi.org/10.1080/08912963.2017.1343314 Published online: 29 Jun 2017. Submit your article to this journal Article views: 130 View Crossmark data Citing articles: 1 View citing articles Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=ghbi20 HISTORICAL BIOLOGY, 2018 VOL. 30, NO. 8, 1139–1149 https://doi.org/10.1080/08912963.2017.1343314 New information on the postcranial skeleton of the Thalassodrominae (Pterosauria, Pterodactyloidea, Tapejaridae) Richard Buchmanna,b , Taissa Rodriguesc, Sabrina Polegariod and Alexander W. A. Kellnere aLaboratório de Mastozoologia, Departamento de Zoologia, Universidade Federal do Estado do Rio de Janeiro, Rio de Janeiro, Brazil; bPós-graduação em Ciências Biológicas, Universidade Federal do Estado do Rio de Janeiro, Rio de Janeiro, Brazil; cLaboratório de Paleontologia, Departamento de Ciências Biológicas, Centro de Ciências Humanas e Naturais, Universidade Federal do Espírito Santo, Vitória, Brazil; dCentro de Ciências Agrárias, Universidade Federal do Espírito Santo, Alegre, Brazil; eLaboratório de Sistemática e Tafonomia de Vertebrados Fósseis, Departamento de Geologia e Paleontologia, Museu Nacional, Universidade Federal do Rio de Janeiro, São Cristóvão, Brazil ABSTRACT ARTICLE HISTORY The clade Tapejaridae is composed by pterosaurs commonly found in fossiliferous deposits in northeastern Received 19 May 2017 Brazil. It is constituted by two less inclusive clades: the smaller-bodied Tapejarinae and the larger Accepted 13 June 2017 Thalassodrominae. Here we describe the specimen MN 6566-V, from the Lower Cretaceous Romualdo KEYWORDS Formation of the Araripe Basin, Brazil. The specimen is overall well preserved tridimensionally, and consists Tapejaridae; of three posterior cervical vertebrae, incomplete right and left scapulocoracoids, and the proximal portion Thalassodrominae; of a right humerus. Comparisons to specimens described in the literature enable its identification as a Romualdo Formation; Lower thalassodromine, whose postcranial material is still poorly known despite the large amount of pterosaurs Cretaceous; Araripe Basin known from this unit. Introduction while Unwin (2003) recovered Tapejara as the sister group of The Tapejaridae is a group of cosmopolitan pterosaurs, known Tupuxuara + Azhdarchidae, i.e. a paraphyletic Tapejaridae, a from Cretaceous deposits in Brazil (Kellner 1989; Kellner and view that found support by Martill and Naish (2006), who also Campos 1994; Campos and Kellner 1997; Kellner and Campos recovered a tree with a paraphyletic Tapejaridae with respect 2002; Sayão and Kellner 2006; Eck et al. 2011; Kellner 2013; Aires to the Azhdarchidae. The discovery and further recognition of et al. 2014; Manzig et al. 2014; Vila Nova et al. 2015), China a related clade, the Chaoyangopteridae, known mostly from (Wang and Zhou 2003a; Lü and Yuan 2005), Spain (Vullo et al. Chinese deposits (Wang and Zhou 2003b; Lü et al. 2008) and 2012), Hungary (Ősi et al. 2011; Andres et al. 2014), Morocco allegedly one species from Brazil (Witton 2008), brought further (Wellnhofer and Buffetaut 1999), and possibly also the United discussion on their phylogenetic relationships. Analyses based States (Kellner 2004) (but see Averianov 2014; who interprets mostly on Kellner’s (2003) characters have consistently recov- the material from Hungary and Morocco as belonging to the ered tapejarines and thalassodromines as sister groups, closely Azhdarchidae). They are edentulous pterosaurs with, normally, related to a clade formed by chaoyangopterids and azhdarchids large sagittal cranial crests. These pterosaurs belong in the more (Vullo et al. 2012; Wang et al. 2012; Aires et al. 2014; Manzig inclusive clade Azhdarchoidea. According to Kellner (2003), et al. 2014). Lü et al. (2008) presented an analysis largely based tapejarids are diagnosed by the confluent naris and nasoantor- on Unwin’s (2003) characters, and recovered either thalasso- bital fenestra, which together make up for more than 45% of the dromines as a sister-group of azhdarchids, in a polytomy with skull length, and the presence of a premaxillary crest beginning tapejarines and chaoyangopterids (in an Adam Consensus tree), at the anterior portion of the skull and extending posteriorly, or thalassodromines and tapejarines as successive sister-groups over the occipital region. of azhdarchids + chaoyangopterids (in a 50% majority-rule con- Both the monophyly and the composition of the Tapejaridae are sensus tree). Other analyses recovered thalassodromines as a disputed. The clades Thalassodrominae (or Thalassodromidae), sister-group to tapejarines + chaoyangopterids (i.e. proposing Tapejarinae, and Chaoyangopterinae (or Chaoyangopteridae) that the Chaoyangopteridae should be considered a subfamily are normally recovered as monophyletic in the literature, but within the Tapejaridae; Pinheiro et al. 2011), or even thalasso- there are several, disparate proposals on their phylogenetic dromines + dsungaripterids as the sister-group of chaoyangop- relationships to each other and to other pterosaurs, such as the terids + azhdarchids, and tapejarines as the sister-group of this Azhdarchidae and the Dsungaripteridae. Kellner (2003, 2004) clade (Andres et al. 2014) (Figure 1). Such disparate proposals, found tapejarids to be the sister group of the Azhdarchidae, besides methodological differences in the analyses, are mostly CONTACT Richard Buchmann [email protected]; Taissa Rodrigues [email protected] © 2017 Informa UK Limited, trading as Taylor & Francis Group Published online 29 Jun 2017 .R. BUCHMANN ET AL 1140 Figure 1. Most recent phylogenetic proposals of the ingroup relationships of the Azhdarchoidea. (A) Vullo et al. (2012); (B) Adams consensus tree from Lü et al. (2008); (C) Pinheiro et al. (2011); (D) Andres et al. (2014). 1. Clade Azhdarchoidea sensu Kellner (2003). due to the incompleteness of several fossils, including the ones Anatomical abbreviations from the Romualdo and Crato formations, which are known con, condyle; cor, coracoid; cot, cotyle; dpc, deltopectoral crest of mostly by skull material. As a discussion on this matter is beyond the humerus; fopn, foramen pneumaticum; fotr, foramen trans- the scope of the present work, the taxonomy presented by Kellner versarium; glfo, glenoid fossa; hyp, hypapophysis; mus, muscle and Campos (2007) is followed. scar; nc, neural canal; ns, neural spine; poex, postexapophysis; Among the tapejarids known from the Romualdo Formation, poz, postzygapophysis; prcor, coracoid process; prsca, scapular thalassodromines are comparatively larger than tapejarines, process; prz, prezygapophysis; sca, scapula; sglpr, supraglenoid and are represented by two genera typical from this deposit, process; tpr, transverse process; tucor, coracoid tuberculum; uc, Tupuxuara and Thalassodromeus (Kellner & Campos 2007). ulnar crest of the humerus. Postcranial material attributed to Thalassodrominae is not unknown but is nonetheless scarce; therefore the description of new material can contribute to the clarification of the phy- Geological setting logenetic position of the group. Here, we describe a specimen (MN 6566-V) referable to the clade consisting of three cervical The Santana Group pertains to the Araripe Basin, which crops out vertebrae, two partial scapulocoracoids, and a partial humerus. in the Brazilian northeast, between the states of Ceará, Pernambuco and Piauí (Neumann and Cabrera 1999; Assine 2007). It is divided in three formations, named, from bottom to top, Crato, Ipubi and Institutional abbreviations Romualdo. The Crato and Romualdo Formations stand out for AMNH, American Museum Natural History, New York, USA; their fossiliferous richness, being Aptian-Albian in age (see Kellner CMN, Canadian Museum of Nature, Ottawa, Canada; MN, et al. 2013 for a discussion of nomenclatural issues). Museu Nacional / Universidade Federal do Rio de Janeiro, Rio The material herein described comes from the Romualdo de Janeiro, Brazil; NSM, National Science Museum, Tokyo, Formation, which is composed of shales, marls and limestones Japan; SMNK, Staatliches Museum für Naturkunde Karlsruhe, (Assine 2007). In the shales, carbonate concretions containing Karlsruhe, Germany. well-preserved fossils are observed, normally in three dimensions. HISTORICAL BIOLOGY 1141 Table 1. Dimensions presented by the cervical vertebrae of MN 6566-V. lost most of its outer surface. It is short and projected posteriorly, Centrum forming a convex surface, resulting in a procoelous centrum, length in mm Centrum width as common in pterosaur vertebrae. Anteriorly, three adjacent (zygapophyses at mid-length Length:width pneumatic foramina (sensu Vila Nova et al. 2015) are present excluded) in mm ratio around the neural canal: one dorsally and two laterally, similar Cervical
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