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The largest flying reptile from Gondwana: A new specimen of Tropeognathus cf. T. mesembrinus Wellnhofer, 1987 (Pterodactyloidea, Anhangueridae) and other large pterosaurs from the Romualdo Formation, Lower Cretaceous, Brazil

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A very large pterosaur (MN 6594-V) from the Romualdo Formation (Aptian/Albian), Santana Group, Araripe Basin, is described. The specimen is referred to Tropeognathus cf. T. mesembrinus mainly due to the presence of a low and blunt frontoparietal crest, the comparatively low number of teeth and the inclined dorsal part of the occipital region. Two distinct wingspan measurements for pterosaurs are introduced: the maximized wingspan (maxws), which essentially consists of doubling the addition of all wing elements and the length of the scapula or the coracoid (the smaller of the two), and the normal wingspan (nws), which applies a reducing factor (rfc) to the maximized wingspan to account for the natural flexures of the wing. The rfc suggested for pteranodontoids is 5%. In the case of MN 6594-V, the maxws and nws are 8.70 m and 8.26 m, respectively, making it the largest pterosaur recovered from Gondwana so far. The distal end of a larger humerus (MCT 1838-R) and a partial wing (MPSC R 1395) are also described showing that large to giant flying reptiles formed a significant part of the pterosaur fauna from the Romualdo Formation. Lastly, some comments on the nomenclatural stability of the Santana deposits are presented.
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... Azhdarchid pterosaurs were well represented in Gondwanan and Laurasian landmasses during the Aptian to Maastrichtian (Witton and Naish, 2008). Some species of this clade were giant forms among pterosaurs (see Kellner et al., 2013) and reached a wingspan of 9e11 m. This is the case of Quetzalcoatlus northropi from the Maastrichtian of United States (Lawson, 1975;Langston, 1981;Kellner and Langston, 1996;Andres and Langston, 2021), Arambourgiania philadelphiae from the Maastrichtian of Jordan (Arambourg, 1959;Frey and Martill, 1996), Hatzegopteryx thambema from the Maastrichtian of Romania (Buffetaut et al., 2002;Naish and Witton, 2017) and Cryodrakon boreas from the Campanian of Canada (Hone et al., 2019). ...
... Descriptions and comparisons were made following the terminol-ogyusedbyKellnerandTomida (2000), Bennett(2001a), Unwin(2003), Averianov (2010), Rodrigues et al. (2011) and Kellner et al. (2013). ...
... The estimation of the wingspan is complex due to two factors: (a) the fragmentary nature of the remains, especially in large-size pterosaurs that are preserved in three dimensions; and (b) the natural flexures of several parts of the wing. In particular, the elbow joint and wing phalanges must be considered in the wingspan estimations (WS), as this parameter influences wingspan (Eaton, 1910;Bennett, 2001b;Wang et al., 2010;Kellner et al., 2013). Kellner et al. (2013) suggest that, whenever possible, two wingspan values should be presented: (a) the maximized wingspan (maxws), which represents the addition of the maximum length of all wing elements, including the syncarpals and the length of the scapula or coracoid (considering the shortest element between them); and (b) the normal wingspan (nws), defined as the maxws reduced by a certain reducing factor (rfc), related with the different natural flexures of the wing. ...
Article
The largest pterosaur discovered in South America and one of the largest flying vertebrates in the world is described. Thanatosdrakon amaru gen. et sp. nov. is a new azhdarchid found in the upper-most levels of the Plottier Formation (upper Coniacian–lower Santonian, Neuquén Basin), Mendoza, Argentina. Two specimens were identified, the holotype (UNCUYO-LD 307) and the paratype, with an estimated wingspan of ∼ 7 m and ∼ 9 m, respectively. The cladistic analysis, based on 216 characters and 97 taxa, recovered Thanatosdrakon deeply nested within Azhdarchidae. Relationships within this clade are well resolved and the clade Quetzalcoatlinae presents two well-defined sister-groups: (Arambourgiania, Mistralazhdarcho, Aerotitan, Hatzegopteryx, Albadraco) + (Cryodrakon, Thanatosdrakon, Quetzalcoatlus ssp.). Thanatosdrakon is the oldest taxon of the clade Quetzalcoatlinae so far. It is represented by several well-preserved axial and appendicular bones in three dimensions. Some of these elements have never been described in giant azhdarchids (e.g. complete norarium, dorsosacral vertebrae and caudal vertebra) and this allows to expand the knowledge about the anatomy of this diverse group of pterosaurs. Finally, from a paleoecological point of view, Thanatosdrakon was found in floodplain deposits of ephemeral meandering systems indicating that this large flying species inhabited continental environments.
... A Formação Romualdo é mundialmente reconhecida por uma diversa fauna de pterossauros excepcionalmente preservados, sendo a grande maioria tridimensionais (Kellner, 2004;Pinheiro & Shultz, 2012;Kellner et al., 2013;Aureliano et al., 2014;Bantim et al., 2014;2021;Headden & Campos, 2014;Vila Nova et al., 2014;Pinheiro & Rodrigues, 2017;Buchmann et al., 2021;Cerqueira et al., 2021). O registro de pterossauros da unidade é abundante, porém muito fragmentado e com identificações dúbias, estas últimas demonstradas por Pinheiro & Rodrigues (2017) para o gênero Anhanguera. ...
Chapter
O rifteamento do Gondwana resultou em mudanças importantes na composição química e distribuição de nutrientes dos oceanos, além de ter contribuído para modificações na circulação de massas de água, o que acarretou na abertura de novos espaços ecológicos. Com a reativação de sistemas de falhas do embasamento durante a ruptura, tem-se a formação de lagos de riftes perenes e temporários, por vezes alimentados por incursões fluviais. Além disso, os pulsos de incursões marinhas durante os eventos de transgressão no final do Cretáceo Inicial também geraram importantes modificações na biota da Bacia do Araripe. A gênese desses novos ecoespaços resultou em grandes transformações bióticas, criando novas possibilidades para o surgimento, a diversificação e a extinção de espécies em um curto espaço temporal. As rochas da Bacia do Araripe registram este momento em que ficam evidentes as relações intrínsecas entre as alterações dos espaços ecológicos e as consequentes biotas registradas em ambientes continentais e marinhos. A integração entre os dados sedimentológicos, estratigráficos e paleontológicos das rochas da Bacia do Araripe permite categorizar três biotas, Missão Velha, Crato e Romualdo, produtos das dinâmicas tectônicas globais associadas à ruptura do Gondwana e suas decorrentes variações climáticas e oceanográficas. Essas biotas apresentam registros paleobiológicos singulares, com animais e vegetais abundantes, diversificados e muitas vezes de caráter endêmico. Muitos dos fósseis identificados são espécies com preservação excepcional, que possibilitam o entendimento de aspectos-chave para a evolução biológica e a relação com as espécies viventes.
... A geologia da Bacia do Araripe é caracterizada por uma estratigrafia complexa formada por quatro sequências limitadas por discordâncias (ASSINE, 1992;1994; sendo: uma Sequência Paleozoica representada pela Formação Cariri; uma Sequência Juro-Neocomiana constituída pelas formações Brejo Santo, Missão Velha e Abaiara; uma Sequência Aptiano-Albiana que compreende as formações Barbalha, Crato, Ipubi e Romualdo e; uma Sequência Albiano-Cenomaniana representada pela Formação Exu (KELLNER et al., 2013;ASSINE et al., 2014;CUSTÓDIO et al., 2017). A deposição das formações pertencentes à Sequência Juro-Neocomiana estão associadas aos processos de rifteamento da Gondwana (ASSINE, 1992). ...
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A Bacia do Araripe é conhecida mundialmente por sua diversidade de fósseis em excelente estado de conservação, que registram a fauna e a flora do Paleozoico ao Mesozoico, depositados durante milhões de anos. A vasta literatura sobre a bacia é concentrada em sua maioria sobre o Mesozoico, com especial interesse ao Grupo Santana, cujo estratos aflorantes são do período Cretáceo, podendo conter lacunas entre seus diferentes sistemas deposicionais. O presente artigo tem como objetivo realizar uma revisão bibliográfica sobre a Formação Barbalha fornecendo o estado da arte e seus gargalos que corresponde a base deste grupo. As bibliografias analisadas indicaram uma predominância de estudos de natureza geológica e uma escassez de trabalhos paleontológicos. Dentre os estudos paleontológicos, a maioria ocorre com microfósseis e a minoria com paleoflora. A carência de estudos fossilíferos para a Formação Barbalha pode estar relacionada aos processos de fossilização, os quais estão diretamente relacionados aos interesses dos pesquisadores em estudar as demais formações, devido a maior qualidade de seu registro fóssil.
... In order to establish the wingspan of the holotype of M. zhuiana, length measurements of published ctenochasmatids (Wellnhofer, 1970(Wellnhofer, , 1978Jiang et al., 2016) were used to calculate a simple regression equation for the wingspan versus skull length (Table 3), and the assignment of Ctenochasma specimens follows Bennett (2007). Two types of wingspans are present in pterosaur researches: the differences are a result of either excluding or including the length of the carpus and coracoid, also known as normal and maximized wingspan, respectively (Bennett, 2001;Kellner et al., 2013). Here, the normal wingspan is used. ...
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Moganopterus zhuiana Lü et al., 2012 was erected as a member of the Boreopteridae, which was questioned by different researchers shortly after the publication. Although the new assignment to the Ctenochasmatidae is widely accepted by pterosaur researchers, some characteristics still require a detailed description. Here, the holotype of this taxon is restudied, and some ambiguous characteristics are re-identified. The diagnosis of this taxon has been revised as the following: a large ctenochasmatid pterosaur, which can be distinguished from other members of this clade by a single autapomorphy: an elongated rod-like parietal crest that extends posterodorsally, forming an angle of about 15° with the ventral margin of the skull. This taxon can be further distinguished from other ctenochasmatids on the basis of the following combination of characteristics: straight occlusal surfaces of the upper and low jaws; presence of a low premaxillary crest confined anterior to the nasoantorbital fenestra; rostrum about two thirds of the skull length; nasoantorbital fenestra occupying slightly more than 20% of the skull length; about 100 slender teeth; and a mid-cervical length/width ratio of about 7. The wingspan of M. zhuiana has been re-estimated according to a simple regression equation for wingspan versus skull length in ctenochasmatids. It confirms that M. zhuiana, although smaller than previous thought, is still the largest known ctenochasmatid. When comparing the sizes of ctenochasmatids in the Jurassic and Cretaceous, ctenochasmatids showed a rough tendency to increase their sizes.
... Moreover, the mesiodistal length between each alveolus (i.e., interalveolar spacing) tends to increase posteriorly (Table 2). Based on the photographs published by Kellner et al. (2013), the sizes of the alveoli are more variable in Coloborhynchus clavirostris than in Ferrodraco and the interalveolar spaces are shorter. The alveolar borders in Ferrodraco are more inflated relative to the jawline and are prominently scalloped. ...
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Ferrodraco lentoni, an anhanguerid from the Upper Cretaceous Winton Formation of northeast Australia, is the most complete Australian pterosaur described to date, represented by a partial cranium, incomplete cervical series and wing elements. Herein we present a comprehensive osteological description of Ferrodraco, as well as an emended diagnosis for this taxon. In addition, we compare Ferrodraco with other isolated pterosaur remains from Australian Cretaceous deposits. Subtle, yet salient, differences indicate that at least three of these specimens, all derived from the upper Albian Toolebuc Formation, are distinct from Ferrodraco. However, we are uncertain whether these specimens are attributable to Mythunga camara, Aussiedraco molnari, Thapunngaka shawi, or an as yet un-named taxon. Detailed description of the postcranial material of Ferrodraco also provides an opportunity to reassess its phylogenetic position. In one analysis, Ferrodraco and Mythunga are resolved as sister taxa within Tropeognathinae, whereas in another, Ferrodraco, Mythunga, and Tropeognathus form a polytomy within Coloborhynchinae. Either way, these slight differences notwithstanding, a close relationship between Ferrodraco and Mythunga is evident, supporting the interpretation that they form a clade. By contrast, Aussiedraco molnari is resolved as a member of Targaryendraconia, a clade with a cosmopolitan distribution. The presence of several anhanguerian taxa or lineages in the late Early and early Late Cretaceous of northeast Australia is suggestive of even greater diversity in the Australian pterosaur fauna.
... Among these are pterosaurs such as Tapejara wellnhoferi Kellner, 1989, andCaupedactylus ybaka Kellner, 2013, among several other findings (e.g. Campos and Kellner, 1985;Kellner et al., 2013;Cerqueira et al., 2021). Additionally, in Upper Cretaceous rocks of the Goio-Erê Formation, Manzig et al. (2014) reported the discovery of a bonebed composed of hundreds of bones of pterosaurs three-dimensionally preserved, most belonging to the tapejarid Caiuajara dobruskii Manzig et al., 2014. ...
Article
We describe a new locality with ctenochasmatid pterosaurs found in a tidal estuarine paleoenvironment of the Quebrada Monardes Formation (Lower Cretaceous). The new locality, which is named "Cerro Tormento", is in Cerros Bravos in the northeast Atacama region, Northern Chile. Here, we describe four cervical vertebrae, one of them belonging to a small individual, the impression of a right scapulocoracoid, a left coracoid, an impression of a left humerus, an incomplete left humerus, a distal fragment of the right humerus, and impressions of a left femur and tibiotarsus. The presence of three humeri and a cervical vertebra belonging to a small pterosaur indicate that these materials represent more than one individual. The cervical vertebrae present diagnostic traits shared with ctenochasmatid pterosaurs, such as elongated vertebral centra, with integrated neural arch, low neural spines, and dorsally located neural canal. It is currently not possible to determine if there are one or more species represented. This finding is the second geographic occurrence of pterosaurs of the clade Ctenochasmatidae in the Atacama region, and although it is currently uncertain if ctenochasmatids from both locations were contemporaneous. This suggests that at least one species of the clade Ctenochasmatidae was widespread in what is now northern Chile. In addition, the presence of bones belonging to more than one individual preserved in Cerro Tormento suggest that pterosaur colonies were present at the southwestern margin of Gondwana during the Lower Cretaceous.
... Assine (2007) e Assine et al. (2014) apontaram que as concreções calcárias exibem em seu interior um total de 18 espécies de peixes, compondo uma paleoictiofauna considerada marinha (Castro-Leal et al., 1998;Fara et al., 2005;Silva-Santos e Valença, 1968). Conforme Kellner e Campos (2000), ainda se encontram vertebrados, dinossauros (Kellner, 1999;Kellner e Campos, 2000;Leonardi e Borgomanero, 1981), tartarugas (Oliveira e Kellner, 2007;Price, 1973) e pterossauros (Aureliano et al., 2014;Campos e Kellner, 1985, 1996Kellner e Campos, 2002;Kellner et al., 2013;Price, 1971). Foram evidenciadas fibras estruturais em uma membrana alar de pterossauro (Campos et al., 1984), tecido mole de peixes preservado em três dimensões (Martill, 1988) e em outros tetrápodes, que incluem parte da pele (dermes e epiderme), dos músculos e dos vasos sanguíneos em pterossauros (Kellner, 1996b(Kellner, , 1996cMartill e Unwin, 1989), dinossauros (Kellner, 1996a) e crocodilianos (Campos et al., 2001;Figueiredo e Kellner, 2009). ...
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The Araripe Basin is a rift basin originated by the rupture of the Gondwana paleocontinent, known as the most complete sedimentary record Interior Basin in the northeastern Brazil. The manuscript presents the state of art of the stratigraphy of the Araripe Basin, focusing on its evolution over time with respect to stratigraphic sequences based on unconformity surfaces, presenting new results and also making a critical review of stratigraphic classifications proposals aiming to elaborate an integrated classification. The first stratigraphic proposals were elaborated in a very comprehensive and simplified form. Over time, more detailed studies have been carried out, corroborating for a better and more refined compartmentalization of the present sedimentary packages. The 1980s were essential for the geoscientific production inherent to the geology of the basin due to the interest in the oil potential of the Brazilian basins. Nevertheless, even today the basin keeps motivating many researches that lead to a better understanding of local and regional geology. The Araripe Basin has five major sequences: Paleozoic, represented by the Cariri formation; Early Rift, consisting of the Brejo Santo formation and the bottom of the Missão Velha formation; Rift, formed by the upper part of the Missão Velha formation and throughout the Abaiara formation; and post- Rift, separated in two sequences: post-rift I constituted by the Barbalha, Crato, Ipubi and Romualdo formations, and post-rift II, characterized by the Araripina and Exu formations. New results are incorporated into the Missão Velha and Abaiara formations, whose distinction is shown here; also, in relation to the Barbalha and Cariri formations new data were gathered. This work focuses on the feasibility of the data, on the interpretations proposed according to the stratigraphic norms and the proposition of a sequential stratigraphic classification.
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