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A new pterodactyloid pterosaur from the Santana Formation (Cretaceous) of Brazil
Abstract
A partial skull comprising fused maxilla/premaxilla and palate of a ctenochasmatoid pterosaur from the Santana Formation of the Araripe Basin in NE Brazil is named as the new genus and species Unwindia trigonus gen. et sp. nov. on account of its long slender rostrum, isodonty with raised dental alveoli and dentition of seven tooth pairs restricted to the portion of the rostrum anterior to the nasoantorbital fenestra. Unwindia is assigned to the Ctenochasmatoidea, and is probably basal within the clade.
... As noted by Vila Nova et al. (2014), many anatomical misinterpretations pertaining to Cearadactylus atrox are largely because the lower jaw was erroneously glued to the maxilla, and the premaxilla-maxilla attached to the dentary. Moreover, the enlarged teeth of the premaxilla were false and not representative of the true dental morphology (Martill, 2011;Vila Nova et al., 2014). A revised diagnosis of Cearadactylus atrox was presented by Vila Nova et al. (2014) noting that it is a valid taxon. ...
... One of the more recently described species of pterosaur from the Romualdo Formation is Unwindia trigonus (SMNK PAL 6597), erected by Martill (2011) and on the basis of the anterior end of a partial premaxilla-maxilla. Martill (2011) originally regarded Unwindia as a ctenochasmatoid, based on comparisons with Yixianopterus jingangshanensis (Lü et al., 2006b) and the presence of relatively few teeth confined to the anterior end of the skull. ...
... One of the more recently described species of pterosaur from the Romualdo Formation is Unwindia trigonus (SMNK PAL 6597), erected by Martill (2011) and on the basis of the anterior end of a partial premaxilla-maxilla. Martill (2011) originally regarded Unwindia as a ctenochasmatoid, based on comparisons with Yixianopterus jingangshanensis (Lü et al., 2006b) and the presence of relatively few teeth confined to the anterior end of the skull. Unwindia clearly differs from other pterodactyloids from the Santana Group, in that it has fewer teeth, with only seven tooth pairs present anterior to the anterior margin of the nasoantorbital fenestra, and less variation in tooth size (Martill, 2011). ...
The Gondwanan pterosaur record is scarce when compared with that of Laurasia and is reviewed here. The majority of Gondwanan pterosaur remains are derived from South America; however, the relative richness of the South American record compared with other Gondwanan continents is largely a result of the ‘Lagerstätten’ effect. Nevertheless, the South American pterosaur assemblage represents the most speciose and diverse known from Gondwana, with several lineages represented, including the Raeticodactylidae, Rhamphorhynchoidea, Darwinoptera, Ctenochasmatidae, Gnathosaurinae, Nyctosauridae, Ornithocheiridae, Tapejaridae, Thalassodromidae, Dsungaripteridae, Chaoyangopteridae and Azhdarchidae. Gondwanan pterosauromorphs are known only from South America. From Africa rhamphorhynchids, archaeopterodactyloids, pteranodontians, nyctosaurids, ornithocheirids, tapejarids, dsungaripteroids, chaoyangopterids, and azhdarchids have been reported. The Arabian Peninsula has produced nyctosaurids, an istiodactyliform, ornithocheirids and azhdarchids. Non-pterodactyloid pterosaurs have been reported from India. A possible azhdarchid has been reported from Madagascar and rhamphorhynchids are known from isolated teeth. The Antarctic pterosaur assemblage also comprises isolated remains of indeterminate pterodactyloids, and a possible indeterminate rhamphorhynchoid. The pterosaur record from East Gondwana comprises ornithocheirids, azhdarchids and a possible ctenochasmatoid from Australia, as well as azhdarchids from New Zealand. Although our understanding of Gondwanan pterosaurs has greatly improved within the last three decades, the discovery and description of more specimens, particularly from Antarctica and East Gondwana, will enhance our understanding of pterosaurian biodiversity and palaeobiogeography.
... Unwindia trigonus Martill, 2011. This taxon is based on a partial rostrum SMNK PAL 6597 from the Albian Romualdo Formation in Santana do Cari ri region, Ceará Province, Brazil (Martill 2011). The genus is characterized by reduced dentition with only seven tooth pairs ante rior to the nasoantorbital fenestra. ...
... The teeth are of similar size, in contrast with the heterodont dentition of ornithocheiroid taxa. Originally, the taxon was referred to the Ctenochasmatoidea (Martill 2011). Witton (2013: 211) cited personal communication from D.M. Unwin that "several features of its jaw and dental morphology are consistent with a lonchodectid identity." ...
... Семейство известно из середины мела (альб-турон) Англии (Lonchodectes compressi rostris, Lonchodraco giganteus, Ikrandraco machaerorhynchus), раннего мела (апт) Китая (Ikrandraco avatar) и позднего мела Европейской России (Lonchodraco (?) sp.). Рассмотрены также другие предполагаемые находки Lonchodectidae из раннего мела Англии (Serradraco sagittirostris (Owen, 1874), BEXHM 2015.18 и Palaeornis cliftii Mantell, 1844), Испании (Prejanopterus curvirostris Fuentes Vidarte et Meijide Calvo, 2010) и Бразилии (Unwindia trigonusMartill, 2011). Ни одна из этих находок не может быть отнесена к данному семейству. ...
The pterodactyloid family Lonchodectidae includes three genera, Lonchodectes Hooley, 1914, Lonchodraco Rodrigues et Kellner, 2013, and Ikrandraco Wang et al., 2014, and four species, Lonchodectes compressirostris (Owen, 1851), Lonchodraco giganteus (Bowerbank, 1846), Ikrandraco avatar Wang et al., 2014, and Ikrandraco machaerorhynchus (Seeley, 1870) comb. nov. [=Ornithocheirus microdon Seeley, 1870 syn. nov.]. The holotype of Lonchodectes compressirostris (NHMUK PV 39410) consists of two fragments of the anterior rostrum, not the mandibular and rostrum fragments as was supposed previously. The difference between Lonchodectes and Ikrandraco is not clear and the taxa could be synonyms. The diagnostic characters for the Lonchodectidae are the presence of the palatal ridge, elevated alveolar margin of the upper and lower jaws, small teeth that are not varying in size, and a prominent mandibular crest (unknown for Lonchodectes). The family includes taxa with long and low rostrum and prominent mandibular crest (Ikrandraco and, possibly, Lonchodectes), or with both premaxil lary and mandibular crests (Lonchodraco). Various phylogenetic analyses place the Lonchodectidae within the Ornithocheiroidea, frequently as a sister taxon to the Anhangueria. The family is known from the midCretaceous (AlbianTuronian) of England (Lonchodectes compressirostris, Lonchodraco giganteus, Ikrandraco machaero-rhynchus), the Lower Cretaceous (Aptian) of China (Ikrandraco avatar), and the Late Cretaceous (Cenomanian) of European Russia (Lonchodraco (?) sp.). The putative records of the Lonchodectidae from the Lower Cretaceous of England (Serradraco sagittirostris (Owen, 1874), BEXHM 2015.18, and Palaeornis cliftii Mantell, 1844), Spain (Prejanopterus curvirostris Fuentes Vidarte et Meijide Calvo, 2010), and Brazil (Unwindia trigonus Martill, 2011) are reviewed. None of them can be attributed to that group.
... According to some authors (e.g., Maisey 1991;Kellner and Campos 2007;Veldmeijer et al. 2009;Witton 2009;Martill 2011), the pterosaurs from the Chapada do Araripe (Cearà, Piauì and Pernambuco States of Brazil) come from the Romualdo and Crato Members of the Santana Formation. Other authors (e.g. ...
... Other authors (e.g. Martill et al. 2007;Vila Nova et al. 2014, 2011Aires et al. 2013;Kellner et al. 2013) follow the lithostratigraphical revision of Neumann and Cabrera (1999): the Santana Formation has changed its status becoming a Group (Santana Group) composed of the Rio da Batateira, Crato, Ipui, Romualdo, Arajara and Exu Formations, in stratigraphic order. We also follow here this last revision. ...
... The Romualdo Formation was once considered Aptian in age (e.g. Wellnhofer 1985Wellnhofer , 1991a, but later it has been reported as Aptian-Albian (Aires et al. 2013;Kellner et al. 2013), Albian (Kellner and Tomida 2000) and latest Albian (Martill 2011) based on the dating to the late Aptian -early Albian of the underlying Crato Formation based on its palynomorph content (Pons et al. 1990). ...
An apparently nearly complete and fairly well-articulated specimen of a large toothed pterosaur from the Lower Cretaceous of Brazil is exhibited at the CosmoCaixa – Museo de la Ciencia, Barcelona, Spain. The museum label refers it to Anhanguera piscator and reports a provenance from the Crato Formation. Its study shows that it is from the Romualdo
Formation and is an assembled and composite specimen over 45% reconstructed. The original skeletal elements show no
diagnostic feature of Anhanguera piscator, belong to non-tapejarid pterosaurs, and do not furnish further significative
scientific information. Several mistakes were made in the reconstruction and assemblage of the skeleton and some elements do not follow the proportions of Anhanguera piscator. Care is suggested when describing pterosaur material from the Romualdo Formation prepared and sold by private fossil dealers in the absence of an adequate documentation on the original material and its preparation.
... Lonchodraco giganteus was briefly described by Bowerbank (1846), and then in more detail by Owen (1851a). The lectotype, NHMUK PV 39412, includes the anterior parts of the rostrum and mandible preserved, and, contra Bowerbank (1846) and subsequent authors (Wellnhofer 1978;Martill 2011), does not include the anterior portion of the nasoantorbital fenestra because what appears to be the anterior margin of the fenestra is not present on both sides of the specimen and most likely represents breakage. The lectotype of Lonchodraco giganteus is readily distinguishable from pterosaurs from other Cretaceous deposits in Britain. ...
... In addition, Tropeognathus mesembrinus has been referred to Ornithocheirus (or Criorhynchus) (e.g., Unwin 2001Unwin , 2003Unwin , 2006. There is even one case where this species is regarded in three different genera within the same publication (Martill 2011): Criorhynchus (Introduction, first page), Tropeognathus (Table 1, second page), and Ornithocheirus (Affinities, fourth page). The holotype of Tropeognathus mesembrinus is a complete rostrum and mandible (BSP 1987 I 46), and there are two referred specimens: an almost complete mandible (SMNS 55414; Veldmeijer 2002) and a partial skeleton that includes an incomplete rostrum and mandible (MN 6594-V;Kellner et al. 2013). ...
... A second dispute involves the use of postcranial material in the analyses of Unwin (2003) and Lü et al. (2010). As indicated previously (Hooley 1914;Kellner 1990;Unwin 2001;Martill 2011), pterosaur cranial and postcranial material have not been found in association in the Cambridge Greensand. In the case of specimens from the Chalk Formation, only the holotype of Lonchodraco giganteus [= Lonchodectes giganteus sensu Unwin 2001] has associated postcranial material. ...
Over a decade after the last major review of the Cambridge Greensand pterosaurs, their systematics remains one of the most disputed points in pterosaur taxonomy. Ornithocheiridae is still a wastebasket for fragmentary taxa, and some nomenclatural issues are still a problem. Here, the species from the Cretaceous of England that, at some point, were referred in Ornithocheirus, are reviewed. Investigation of the primary literature confirmed that Criorhynchus should be considered an objective junior synonym of Ornithocheirus. Taxonomic review of more than 30 species known from fragmentary remains showed that 16 of them are undiagnosable (nomina dubia): Palaeornis cliftii, Cimoliornis diomedeus, Pterodactylus compressirostris, Pterodactylus fittoni, Pterodactylus woodwardi, Ornithocheirus brachyrhinus, Ornithocheirus carteri, Ornithocheirus crassidens, Ornithocheirus dentatus, Ornithocheirus enchorhynchus, Ornithocheirus eurygnathus, Ornithocheirus oxyrhinus, Ornithocheirus scaphorhynchus, Ornithocheirus tenuirostris, Ornithocheirus xyphorhynchus, and Pterodactylus sagittirostris. Fourteen species are considered valid, and diagnoses are provided to all of them: Ornithocheirus simus, Lonchodraco giganteus comb. n., Lonchodraco machaerorhynchus comb. n., Lonchodraco(?) microdon comb. n., Coloborhynchus clavirostris, 'Ornithocheirus' capito, Camposipterus nasutus comb. n., Camposipterus(?) sedgwickii comb. n., Camposipterus(?) colorhinus comb. n., Cimoliopterus cuvieri comb. n., 'Ornithocheirus' polyodon, 'Ornithocheirus' platystomus, 'Pterodactylus' daviesii, and 'Ornithocheirus' denticulatus. These species are referred in the genera Ornithocheirus, Lonchodraco gen. n., Coloborhynchus, Cimoliopterus gen. n., and Camposipterus gen. n., but additional genera are probably present, as indicated by the use of single quotation marks throughout the text. A cladistic analysis demonstrates that Anhangueridae lies within a newly recognized clade, here named Anhangueria, which also includes the genera Cearadactylus, Brasileodactylus, Ludodactylus, and Camposipterus. The anhanguerian 'Cearadactylus' ligabuei belongs to a different genus than Cearadactylus atrox. Lonchodraconidaefam. n. (more or less equivalent to LonchodectidaesensuUnwin 2001) is a monophyletic entity, but its exact phylogenetic position remains uncertain, as is the case of Ornithocheirus simus. Therefore, it is proposed that Ornithocheiridae should be constricted to its type species and thus is redundant. Other taxa previously referred as "ornithocheirids" are discussed in light of the revised taxonomy.
... The unit is characterized by sandstones and conglomerates at the base, thick shale layers, and shell beds (coquinas) near the top (Beurlen, 1971;Mabesoone and Tinoco, 1973;Sales, 2005;Custódio et al., 2017;Fürsich et al., 2019). Also, it is widely recognized for its fossil content, characterizing a critical Konservat-Lagerstätte of the Early Cretaceous (Mabesoone and Tinoco, 1973;Maisey, 1991;Martill, 1997Martill, , 2007Martill, , 2011Kellner and Campos, 1999;Kellner, 2002;Fara et al., 2005;Martill et al., 2012). Although the influence of marine waters is clear, the seaway route and its impact on the paleoenvironment remain controversial. ...
... The first cycle is mainly characterized by concretion-bearing black shales, subordinated sandstones, and conglomerates at the base. Fossiliferous concretion-bearing black shales are approximately 40 m thick, being recognized as a Konservat-Lagerstätte (Mabesoone and Tinoco, 1973;Martill, 1988Martill, , 1997Martill, , 2007Martill, , 2011Maisey, 1991;Kellner and Campos, 1999;Fara et al., 2005;Martill et al., 2012). Indeed, such interval is interpreted as the unit's first marine incursion and marks its maximum flooding zone (Custódio et al., 2017;Fürsich et al., 2019). ...
The Aptian Romualdo Formation in the Araripe Basin represents the establishment of an epeiric sea in the Brazilian Northeast during the Early Cretaceous. This unit is composed by a succession of limestones, shales, sandstones, and coquinas, which constitute shell beds near the top. This last interval strongly suggests a marine influence in the whole basin based on the fossil assemblage. However, even though the establishment of marine waters during the deposition of the Romualdo Formation is a consensus, paleogeographical and paleoenvironmental aspects still need to be better understood. This study aims to perform a sedimentological analysis of the coquina interval, based on petrographic and taphonomic analysis of field data and in a new fully recovered well-core (2-AB-1-CE) drilled in the Serra do Mãozinha at Abaiara county in the Ceará State. The described rocks were subdivided into eight facies constituted by bivalves, gastropods, ostracods, peloids, micrite, siliciclastic grains, punctual foraminifers, and plant debris. Based on facies' vertical stacking at the well 2-AB-1-CE, a main regressive pattern was observed. It was also possible to subdivide the coquina interval into high-frequency transgressive-regressive cycles. The regional distribution of the facies permitted the identification of a shallow epeiric sea throughout the basin, with variations in the paleoenvironment energy and siliciclastic input. These variations were mainly controlled by paleotopography, fluvial discharge, and sea-level oscillations. The data allowed the identification of two subsequent paleogeographical configurations controlled by sea-level change. The first one, related to a sea-level high, was characterized by a protected bay on the western border and a gulf in the rest of the basin. Then a sea-level fall led to the formation of closed and open lagoons associated with higher terrigenous supply. As such, this study adds new data for a better understanding of the Araripe Basin and the impacts of sea-level fluctuations in South America during the Aptian.
... This is recorded in the Araripe Basin as the late Aptian Romualdo Formation (Fig. 1). This unit encompasses one of the most important fossil Konservat-Lagerstätten of the world (Mabesoone and Tinoco, 1973;Maisey, 1991;Martill, 1997Martill, , 2007Martill, , 2011Kellner and Campos, 1999;Kellner, 2002;Fara et al., 2005;Vila Nova et al., 2011;Martill et al., 2012), an impressive archive of the palaeobiota that lived and flourished in Gondwana during the early Cretaceous (Martill, 2007). Due to the high preservation quality of several fossil groups, especially 3D-preserved fishes (Martill, 1988;Maldanis et al., 2016), scientific studies have focused mainly on the systematic paleontology. ...
... Santos and Valença, 1968;Martill, 1988;Maisey, 1991;Fara et al., 2005), pterosaurs (e.g. Vila Nova et al., 2011 ;Martill, 2011;Aires et al., 2014), turtles (e.g. Meylan, 1996;Hirayama, 1998;Kellner and Campos, 1999;Romano et al., 2013), dinosaurs (Kellner, 1997;Kellner and Campos, 2000), and crustaceans (Santana et al., 2013;Pinheiro et al., 2014). ...
Geologic events related to the opening of the South Atlantic Ocean deeply influenced the sedimentary record of the Araripe Basin. As consequence, upper stratigraphic units of the basin record a marine ingression in northeastern Brazil during the late Aptian. The timing and stratigraphic architecture of these units are crucial to understand the paleogeography of Gondwana and how the proto-Atlantic Ocean reached interior NE Brazil during the early Cretaceous. This marine ingression is recorded in the Araripe Basin as the Romualdo Formation, characterized by a transgressive-regressive cycle bounded by two regional unconformities. In the eastern part of the basin, the Romualdo depositional sequence comprises coastal alluvial and tide-dominated deposits followed by marine transgressive facies characterized by two fossil-rich intervals: a lower interval of black shales with fossil-rich carbonate concretions (Konservat-Lagerstätten) and an upper level with mollusk-dominated shell beds and shelly limestones. Following the marine ingression, an incomplete regressive succession of marginal-marine facies records the return of continental environments to the basin. The stratigraphic framework based on the correlation of several sections defines a transgressive-regressive cycle with depositional dip towards southeast, decreasing in thickness towards northwest, and with source areas located at the northern side of the basin. The facies-cycle wedge-geometry, together with paleocurrent data, indicates a coastal onlap towards NNW. Therefore, contrary to several paleogeographic scenarios previously proposed, the marine ingression would have reached the western parts of the Araripe Basin from the SSE.
... The pattern in pre-metamorphic larval dentition closely resembles the dentition observed in some flying pterosaurs. Anhanguera blittersdorfi and Cearadactylus atrox (FIGURE 7.2A) (Martill, 2011), like several other pterosaur species (FIGURE 7.2B, C) (Unwin, 2002, Witmer et al., 2003, had anterior protruding teeth that were used to grab slippery fish. Their posterior teeth were short and perpendicular oriented (FIGURE 7.2A, C) and they were hypothesized to kill the grabbed fish and to partially process it before ingestion (Martill, 2011). ...
... Anhanguera blittersdorfi and Cearadactylus atrox (FIGURE 7.2A) (Martill, 2011), like several other pterosaur species (FIGURE 7.2B, C) (Unwin, 2002, Witmer et al., 2003, had anterior protruding teeth that were used to grab slippery fish. Their posterior teeth were short and perpendicular oriented (FIGURE 7.2A, C) and they were hypothesized to kill the grabbed fish and to partially process it before ingestion (Martill, 2011). Therefore, it can now be hypothesized that leptocephali trap soft or gelatinous prey organisms in their mouth cavity with their protruding teeth, after which the trapped organism can be transported towards to oesophagus using a tongue-like structure (observed during reconstruction, data not shown). ...
... Pterosaurs underwent dramatic diversification in morphology in the Cretaceous (Prentice et al., 2011), and that includes their teeth -some clades, notably pteranodontids, lost their teeth entirely, whereas others (pterodactylids, germanodactylids, dsungaripterids, ornithodesmids) reduced the dentition to small numbers of short teeth. Only pterodaustrids and ctenochasmatids adopted long, slender teeth, and those of the former clade are hair-like or baleen-like (Wellnhofer, 1978;Martill, 2011). This suggests that the new teeth can be identified as ctenochasmatid, and in particular as gnathosaurine ctenochasmatid, an identification followed in analogous cases for long, slender, needle-like teeth in Early Cretaceous continental sedimentary deposits (e.g. ...
... Sánchez-Hernández et al., 2007, Fig. 5;Sweetman and Martill, 2010, Fig. 5). Ctenochasmatoid pterosaurs are well known mainly from the Early Cretaceous period in Asia, with numerous taxa from the Yixian Formation (early Aptian) of NE China (Lü et al., 2011), but with additional reports from South America (Martill, 2011) and Europe. Previous European reports include specimens from the Late Jurassic Solnhofen Limestone Formation (Tithonian) of southern Germany and equivalent-aged rocks of northern France (Bennett, 2007), the Purbeck Limestone Group (Berriasian) of southern England (Howse and Milner, 1995), and the Camarillas Formation (lower Barremian) of Burgos, Spain (Sánchez-Hernández et al., 2007) and the Leza Formation (Lower Cretaceous) of La Rioja, Spain (Pereda Suberbiola et al., 2012b). ...
New vertebrate remains reported from the Papo-Seco Formation (Lower Barremian, Lower Cretaceous) of Areias do Mastro, in Cabo Espichel, SW Portugal, south of Lisbon. The marine, lagoonal, and estuarine limestones, marls, sands and gravels have yielded remains of dinosaurs and other reptiles since the 19th century. Recent paleontological prospecting produced several vertebrate remains, including turtle shell fragments, crocodilian teeth, fish and pterosaurs. Research identified both bones and teeth of fish, crocodiles, dinosaurs Baryonyx and iguanodontian, as well as a ctenochasmatoid pterosaur, and a possible ornithocheirid pterosaur. These new disclosures are an important contribution to the knowledge of vertebrate diversity from the Portuguese Cretaceous. Faunal species combination proven to be similar to other faunal associations of Barremian formations in the Iberian Peninsula. © 2015, Universidad Complutense de Madrid. All Rights Reserved.
... Kellner & Campos (1988) concluded that there was not sufficient evidence to support the erection of a new taxon and considered "Pricesaurus megalodon" a nomen vanum. After this, the taxon was considered "of dubious validity" by Martill (2011). ...
... Premaxillary crests restricted to the rostrum are well spread among toothed pterosaurs from the Romualdo Formation (Campos & Kellner, 1985;Wellnhofer, 1987;Wellnhofer, 1991;Kellner & Tomida, 2000;Veldmeijer, 2003a). until now, only Brasileodactylus, Barbosania and unwindia securely lacked this structure (Veldmeijer 2003b;Veldmeijer et al., 2009;Elgin & Frey, 2011;Martill, 2011). The genus Brasileodactylus is also found in the older Crato Formation (sayão & Kellner, 2000). ...
The specimens attributed to “Pricesaurus megalodon” are re-examined in order to determine whether they constitute a distinct taxon of pterodactyloid pterosaur. After careful preparation and study, the material (rostral and middle parts of two probably distinct skulls) revealed affinities with the Anhangueridae, especially with the genus Anhanguera Campos & Kellner. The taxon “Pricesaurus megalodon” is considered here a nomen nudum for lacking distinctive diagnostic features and for being published in noncompliance with the standards of the International Code of Zoological Nomenclature.
... Posterior teeth of ctenochasmatids also exhibit some similarities to NNPM2588-65 in having a circular base, albeit they are often more elongated and sometimes possess defined carinae (Soto et al., 2021). Teeth of Unwindia trigonus, a pterosaur of uncertain affinities from the Santana formation, Brazil, were only slightly labiolingually compressed (Martill, 2011). The anterior teeth of anhanguerians sometimes had circular bases, with more posterior tooth morphologies being more labio-lingually compressed (Solonin et al., 2021). ...
Early Cretaceous was a time of great diversity for pterosaurs with numerous taxa described from around the world. However, pterosaur record from Eastern Europe, especially in the Early Cretaceous, is scarce. In this study we describe several isolated pterosaur teeth from the Albian deposits of marine Burim formation in Kaniv Natural Reserve, Cherkasy region, Ukraine. Pterosaur fossils are uncommon in this fauna that is dominated by cartilaginous fish and actinopterygians. Pterosaur material is represented by at least two distinct morphotypes corresponding to Anhangueria and an indeterminate pterosaur taxon. Despite their fragmentary nature, these findings are significant as they represent the first described pterosaur fossils from Ukraine and the first described pterosaur teeth from the Early Cretaceous of Eastern Europe. Additionally, we discuss the potential for terrestrial vertebrate material being found in Mesozoic marine deposits which are abundant across Ukraine but remain poorly studied.
... Most of these remains are currently under study, and among these, four pterosaur manus tracks are the main goal of this contribution. The Gondwanan pterosaur ichnological record is still poorly documented [27][28][29][30][31][32], whereas the bone record is very abundant, especially in South America, and particularly in Argentina and Brazil, with some records in Chile, Peru and Venezuela [33][34][35][36][37][38][39][40][41][42][43][44][45][46][47][48][49][50][51][52]. The aims are to describe, compare and analyze the new pterosaur tracks found at the ANPM Paso Córdoba and then to discuss the important role that they have in the reconstruction of the paleocommunity of the Anacleto Formation and for the latest Cretaceous of Gondwana. ...
The Campanian Anacleto Formation holds an abundant and diverse ichnofossil and body-fossil vertebrate record. Despite the striking diversity of this record, pterosaur fossils had never been described from the unit. Here, we report four pterosaur manus tracks from fluvial red beds cropping out in the Área Natural Protegida Municipal Paso Córdoba (Río Negro Province, northern Patagonia, Argentina). Tracks are longer than wide, tridactyl with digit impressions of different lengths (I < II < III), anteriorly directed and laterally asymmetrical. Being on loose slabs and lacking direct examination of pes morphology, the material is classified as undetermined pterosaur tracks. The new find represents the first occurrence of pterosaurs from the lower–middle Campanian of Argentina and one of the few evidences from South America for this time interval. In addition, it is one of the few ichnological pterosaur records from Gondwana, thus shedding light on the palaeobiogeography of this clade during the latest Cretaceous. Pterosaur tracks from the Anacleto Formation allow us to integrate the body-fossil record from the unit and to add a new component, along with birds, to the flying archosaur fauna coexisting with non-avian dinosaurs, notosuchians, chelonians, squamates and mammals in the Campanian of northern Patagonia.
... This fact, together with the absence of reported exportation permits, leads us to consider that these fossils may have been purchased (figure 5). Some publications state that the fossils were 'obtained from a quarry workman' [92] or 'from a fossil digger' [93] and eight publications [28,30,[94][95][96][97][98][99] directly acknowledge that the specimen was purchased. ...
Scientific practices stemming from colonialism, whereby middle- and low-income countries supply data for high-income countries and the contributions of local expertise are devalued, are still prevalent today in the field of palaeontology. In response to these unjust practices, countries such as Mexico and Brazil adopted protective laws and regulations during the twentieth century to preserve their palaeontological heritage. However, scientific colonialism is still reflected in many publications describing fossil specimens recovered from these countries. Here, we present examples of ‘palaeontological colonialism’ from publications on Jurassic–Cretaceous fossils from NE Mexico and NE Brazil spanning the last three decades. Common issues that we identified in these publications are the absence of both fieldwork and export permit declarations and the lack of local experts among authorships. In Mexico, access to many fossil specimens is restricted on account of these specimens being housed in private collections, whereas a high number of studies on Brazilian fossils are based on specimens illegally reposited in foreign collections, particularly in Germany and Japan. Finally, we outline and discuss the wider academic and social impacts of these research practices, and propose exhaustive recommendations to scientists, journals, museums, research institutions and government and funding agencies in order to overcome these practices.
... The Araripe Basin in northeastern Brazil -a region of low socioeconomic status -where 'Ubirajara' was found has suffered from illicit fossil trafficking for decades 4,5 . In fact, some foreign researchers openly admit to having obtained Araripe fossils from commercial sources or fossil dealers (for example, the dinosaur Irritator challengeri, the crocodilian Susisuchus anatoceps and the pterosaurs Ludodactylus sibbicki, Lacusovagus magnificens, Unwindia trigonus and Tupandactylus navigans [6][7][8][9][10][11] ). In one notorious case, researchers even conceded they were "deceived by the fossil dealers" 9 . ...
To the Editor — The dinosaur fossil ‘Ubirajara jubatus’ was allegedly exported from Brazil in 1995 and is currently held by State Museum of Natural History Karlsruhe (SMNK) in Germany. It has attracted the attention of the scientific community not only because of its intrinsic interest but also for the controversy surrounding its export from Brazil, which has put into question the legal and ethical context of this work and led to the removal of the publication. Note that ‘Ubirajara jubatus’ is no longer a valid scientific name due to this retraction, hence its appearance in quotation marks here. ‘Ubirajara’ is far from a unique case of an illegally acquired and exported fossil residing in a foreign museum. The Araripe Basin in northeastern Brazil — a region of low socioeconomic status — where ‘Ubirajara’ was found has suffered from illicit fossil trafficking for decades. In fact, some foreign researchers openly admit to having obtained Araripe fossils from commercial sources or fossil dealers (for example, the dinosaur Irritator challengeri, the crocodilian Susisuchus anatoceps and the pterosaurs Ludodactylus sibbicki, Lacusovagus magnificens, Unwindia trigonus and Tupandactylus navigans). In one notorious case, researchers even conceded they were “deceived by the fossil dealers”. Many of the illicit fossils, some of which are holotypes (specimens used to designate new species), are stored in German museums. Like many other Latin American countries, Brazil adopted rigid laws...
... Ikrandraco avatar and Haopterus gracilis also exhibit a certain degree of labiolingual compression of the teeth, at least in the distal part of the dentition (Wang & Lü, 2001;Wang et al., 2015), though not to the same degree seen in the istiodactylids and Hongshanopterus. The last two mandibular alveoli preserved in the holotype of Lonchodraco giganteus (the sister-taxon of Ikrandraco avatar in our analysis) are also labiolingually narrow (see Martill, 2011;Rodrigues & Kellner, 2013). ...
A new istiodactylid pterosaur, Nurhachius luei sp. nov., is here reported based on a complete skull with mandible and some cervical vertebrae from the lower part of the Jiufotang Formation of western Liaoning (China). This is the second species of Nurhachius , the type-species being N. ignaciobritoi from the upper part of the Jiufotang Formation. A revised diagnosis of the genus Nurhachius is provided, being this taxon characterized by the presence of a slight dorsal deflection of the palatal anterior tip, which is homoplastic with the Anhangueria and Cimoliopterus. N. luei sp. nov. shows an unusual pattern of tooth replacement, with respect to other pterodactyloid species. The relationships within the Istiodactylidae and with their closest taxa are investigated through a phylogenetic analysis by parsimony.
... This specimen was originally described as a tapejarid by Veldmeijer et al. (2005), an interpretation followed recently by Pêgas et al. (2016). Martill (2011) assigned a partial rostrum from the Romualdo Formation to the Ctenochasmatoidea, although this has been disputed by Witton (2013). Finally, Lacusovagus magnificens from the Crato Formation was originally interpreted as the first non-Chinese chaoyangopterid (Lü et al. 2008;Witton 2008), which was later challenged (Kellner 2013). ...
The Brazilian Crato Formation (Lower Cretaceous, Aptian) is well known for its rich pterosaur fauna. This paper deals with a new find represented by four articulated mid-cervical vertebrae.
The vertebrae show a morphology consistent with that seen in the Chaoyangopteridae, especially the relative elongation, low neural spines, lack of pneumatic foramina on the lateral face of
the centra and the presence of well-developed postexapophyses. Chaoyangopterids are, so far, represented with confidence only in Chinese deposits; the only record outside the Jehol Group is the
Crato Formation form Lacusovagus magnificens, a partial skull whose assignment to the Chaoyangopteridae has been disputed. Given this controversy, we review the phylogenetic position of Lacusovagus,
and discuss the nesting of our new specimen among theChaoyangopteridae, providing some comments concerning the composition of the group. We conclude that our new specimen provides further support for the presence of chaoyangopterids in the Early Cretaceous of Brazil.
... Character 45, state 1 (variation in size of the anterior teeth, with the 5 th and 6 th lower than the 4 th and 7 th ) is an unambiguous synapomorphy of the Anhanguera genus, corroborating previous proposals (Kellner 2003;Rodrigues and Kellner 2008) that used this feature to rule out the presence of such taxa as the European forms Coloborhynchus and Ornithocheirus from the Araripe Basin (Unwin 2003;Veldmeijer 2003Veldmeijer , 2006Fastnach 2001;Martill 2011;Martill and Unwin 2012). In M. kellneri, the 5 th , 6 th and 7 th alveoli are smaller than the 4 th and 8 th , and in Tropeognathus mesembrinus the 2 nd and 3 rd pairs are the largest in size. ...
A new species of pterosaur, Maaradactylus kellneri gen. nov., sp. nov. (Archosauria: Pterosauria) from the Romualdo Formation (Aptian/Albian), is herein described. The specimen (MPSC R 2357) was found at Sítio São Gonc¸alo, Santana do Cariri city (State of Ceará, northeast Brazil) and consists of the skull, atlas and axis, and represents one of the largest skulls of the Anhangueridae from the Araripe Basin described. The autapomorphies of the new pterosaur include the following characters: a premaxillary sagittal crest that is relatively long and high, beginning at the anterior part of the skull (rostrum) and extending to the 22ndpair of alveoli, not covering the nasoantorbital fenestra or the choanaes, and also the presence of 35 pairs of alveoli; smooth palatal ridge, which starts on the 5thpair of alveoli and ends on the 13thpair; palate is convex shaped in the anterior region; choanae not extending laterally; small and convex palatal elevation; the 5th, 6thand 7thalveoli smaller than the 4thand 8th; the alveoli decreasing in size from the 9thto the 12thand increasing from the 13thto 18th, and from the 18thto the 35ththey are arranged in triplets. Furthermore, the lateral surface of the premaxillary crest shows grooves and tridimensional structures that may have housed blood vessels.
... Nesov (1991: 19) previously suggested that this specimen might belong to an azhdarchid.Unwin 2001Unwin , 2003 Witton et al. 2009 ), because these postcranial elements (cervicals , humerus, and femur) bear certain azhdarchoid characters. However, the known cranial remains of Lonchodectes, exclusively consisting of toothed jaw fragments, do not show any specific resemblance with azhdarchoid pterosaurs (Unwin 2001; Martill 2011). It is more parsimonious to combine the cranial remains from the Cambridge Greensand, including the toothless holotype of O. sedgwicki and the skull fragment CAMSM B.54406 with Tapejara-like supraoccipital crest, with the postcranial elements from the same stratigraphic unit bearing certain azhdarchoid characters (moderately elongated middle cervicals, humerus with straight deltopectoral crest and pneumatic foramen on ventral side, and femur with low neck to shaft angle and large pneumatic foramen at the greater trochanter). ...
Ornithostoma sedgwicki Seeley, 1891 from the Lower Cretaceous (Albian) Cambridge Greensand of England is represented by edentulous jaw fragments, posterior skull fragment with the supraoccipital crest, and by several postcranial bones attributed previously to Lonchodectes. Ornithostoma is referred to the Azhdarchoidea based on a combination of derived characters (teeth absent, middle cervicals moderately elongated, pneumatic foramen on anterior side of humerus, large pneumatic foramen on posterior side of femur at greater trochanter) and plesiomorphic characters (deltopectoral crest not warped, femoral neck to shaft angle less than 145°). The structure of the supraoc-cipital crest and humerus resembles those in Tapejara, but Ornithostoma differs from that taxon by a strong median ridge on the occiput presumably associated with a more elongated rostrum. At least three taxa of basal azhdarchoids were present in the British Lower Cretaceous. РЕЗЮМЕ Ornithostoma sedgwicki Seeley, 1891 из нижнемелового (альб) кембриджского зеленого песчаника Англии представлен фрагментами беззубых челюстей, фрагментом задней части черепа с затылочным гребнем, и некоторыми костями посткраниального скелета, которые раньше относили к Lonchodectes. Ornithostoma от-носится к Azhdarchoidea на основании комбинации продвинутых признаков (отсутствие зубов, умеренно удлиненные среднешейные позвонки, отверстие пневматизации на передней стороне плечевой кости, круп-ное отверстие пневматизации на задней стороне бедренной кости у большого трохантера) и плезиоморфных признаков (дельтопекторальный гребень не загнут, угол между шейкой и телом бедренной кости меньше 145°). Строение затылочного гребня и плечевой кости сходно с таковыми у Tapejara, но Ornithostoma отлича-ется от последнего таксона мощным медиальным гребнем на затылке, предположительно связанным с более удлиненной ростральной частью черепа. В нижнем мелу Великобритании существовало, по крайней мере, три таксона базальных аждархоидов.
... While lacking elements comparable to MPSC R 859, the recently-described Unwindia trigonus [27], also from the Romualdo Formation, has teeth restricted to the rostral end of the skull, well anteriorly from the rostral margin of the nasoantorbital fenestrae. The incomplete nature of Unwindia's holotype avoids an accurate determination of the nasoantorbital opening's size for this taxon. ...
A new and unusual specimen of a probable azhdarchoid pterosaur is described for the Early Cretaceous (Albian) Romualdo Formation of Brazil. The specimen consists of a palate that, although fragmentary, has a unique morphology differing from all other known pterosaurs with preservation of palatal elements. The new specimen probably indicates the presence of a yet undescribed pterodactyloid taxon for Romualdo Formation and brings new information on pterosaur diversity of this sedimentary unity. Mainly due to the rarity of pterodactyloid specimens with palate preservation, this structure has been overlooked in this clade. Here, we reassess the palatal anatomy of Pterodactyloidea, revealing an intriguing variety of morphotypes and evolutionary trends, some of them described here for the first time. The morphological disparity displayed by different pterodactyloid taxa may be further evidence of the presence of diverse feeding strategies within the clade.
O Membro Romualdo (Formação Santana) corresponde a uma sucessão de arenitos,
folhelhos, calcários e coquinas depositados durante a idade Aptiano (estágio local
Alagoas) na Bacia do Araripe. Esta unidade, limitada por duas discordâncias regionais,
é reconhecida pelo seu vasto registro fossilífero que a caracteriza como um Konservat-
Lagerstätte. A deposição das rochas do Membro Romualdo está associada a eventos
de ingressão marinha, registrados em folhelhos e coquinas, responsáveis por alterar
aspectos fisiográficos no interior do nordeste brasileiro. Esses eventos, interpretados
por meio de dados palinológicos e paleontológicos, são pouco compreendidos em
termos sedimentológicos. Um melhor entendimento da evolução paleoambiental e
paleogeográfica do Membro Romualdo auxilia na caracterização geológica do Aptiano
no Brasil, momento esse em que importantes reservatórios de petróleo do intervalo
pré-sal estavam em desenvolvimento nas bacias da margem leste meridional
brasileira. Neste sentido, este trabalho apresenta um modelo paleoambiental evolutivo
das rochas desta unidade com base em uma análise faciológica, litogeoquímica e
petrofísica. De maneira similar é também apresentado um modelo paleogeográfico
utilizando aspectos bioestratinômicos de coquinas que ocorrem na porção superior
deste membro. Este trabalho é baseado em dados de testemunho de sondagem,
obtidos através do poço estratigráfico 2-AB-1-CE, descrições de afloramentos (com
ênfase no intervalo das coquinas), descrições petrográficas, análises de fluorescência
de raio X portátil (pFRX) e dados de raio gama (RG) e raio gama espectral (RGe).
Através da análise de fácies foram identificadas 19 fácies sedimentares e cinco
sucessões de fácies. A partir da integração dos aspectos faciológicos com os dados
geoquímicos (pFRX) e petrofísicos (RG e RGe) foram confeccionados cinco modelos
paleoambientais representando as fases lacustre, de planície aluvial e leques
terminais, de mar interior, de frente deltaica e fluvial costeiro. Ainda, a análise
bioestratinômica do intervalo das coquinas permitiu reconstruir a paleogeografia da
Bacia do Araripe durante os estágios finais do Aptiano. Por meio desta análise uma
conexão entre as bacias do Araripe e Potiguar é sugerida durante os momentos de
ingressão marinha no nordeste brasileiro.
A new and articulated specimen of a pterosaur wing including upper arm, forearm, parts of the carpus and metacarpus, and a wing phalanx from Maastrichtian phosphatic deposits of Morocco are assigned to Tethydraco cf. regalis Longrich et al., 2018.
The specimen comes from the village of Ouled Abdoun, close to the Oued Zem basin and its phosphatic mines (Morocco). The fossil is part of the collection of the Université Hassan II of Casablanca (ID Number FSAC CP 251).
In the first part, the thesis presents a synthetic introduction about the morphology, anatomy, physiology and evolution of pterosaurs in order to offer a comprehensive framework on this fascinating group of extinct flying tetrapods.
The main goal of this work is the taxonomic identification of the specimen, principally by morphological and morphometric/statistic analysis, based on the comparison with the most similar pterosaurs of the same epoch.
Aspect of the humerus morphology and dimensional ratios of the wing elements suggest that T. cf. regalis is an azhdarchid rather than pteranodontid, as originally proposed. A high abundance of azhdarchid remains in the open marine setting of the Moroccan phosphates casts doubt on suggestions that Azhdarchidae were largely terrestrial pterosaurs.
Fundado em 1985, o Museu de Paleontologia em Santana do Cariri (MPSC) tem armazenado fosseis provenientes da Bacia do Araripe, com o objetivo de guardar e proteger especimes importantes para a ciencia. Os fosseis que compoem o acervo sao, por exemplo, peixes, plantas, tartarugas, crocodilomorfos, insetos, crustaceos, anuros, conchas, bivalves, dinossauros e pterossauros. Este trabalho e a continuacao de outro que foi iniciado ha 13 anos atras, desde entao, a colecao de pterossauros aumentou consideravelmente, e hoje conta com 134 especimes, dotados de material pos-craniano (155), crânio (19), vertebra cervical (5), fragmentos desconhecidos (2), fosseis adulterados (8) e mais 38 pecas que foram classificadas erroneamente como pterossauros. Alguns especimes de pterossauros se encontram sob emprestimo em outras instituicoes ou em preparacao no Laboratorio de Paleontologia da Universidade Regional do Cariri (LPU).
The Cambridge Greensand, a remanié deposit that crops out in Cambridgeshire, eastern England, has yielded numerous, though fragmentary, late Early Cretaceous (Albian) vertebrate fossils including more than 2000 isolated pterosaur bones. So far, 32 species of pterosaur have been proposed in connection with the Cambridge Greensand material, but there has been and continues to be considerable confusion concerning the validity of these taxa, their relationships to each other and to other pterosaurs, and the material upon which they were established. A comprehensive systematic revision identified eleven valid species distributed among three families: the Ornithocheiridae (Ornithocheirus simus and possibly a second, as yet unnamed species of Ornithocheirus, Coloborhynchus capito, Coloborhynchus sedgwickii, Anhanguera cuvieri, and Anhanguera fittoni); the Lonchodectidae (Lonchodectes compressirostris, Lonchodectes machaerorhynchus, Lonchodectes microdon and Lonchodectes platystomus); and a species of edentulous pterosaur (Ornithostoma sedgwicki) that may represent the earliest record for the Pteranodontidae. It is possible that some of the taxa currently recognised represent sexual dimorphs (Coloborhynchus capito and Coloborhynchus sedgwickii, Lonchodectes compressirostris and Lonchodectes machaerorhynchus), or disjunct populations of a single species (Ornithocheirus simus and Ornithocheirus sp., Lonchodectes compressirostris and Lonchodectes microdon) and that there may be as few as seven valid species, but the Cambridge Greensand pterosaurs are too poorly known to demonstrate this at present. The Cambridge Greensand pterosaur assemblage is similar to a slightly younger, but much smaller assemblage from the Lower Chalk of England and shares some elements, such as ornithocheirids, in common with many other late Early and early Late Cretaceous assemblages. It is distinguished by the absence of tapejarids and the presence of Lonchodectes which, so far, is only known from the Cretaceous of England. The disparity in taxonomic composition is possibly related to ecological differentiation, and might also reflect some provincialism in late Early and early Late Cretaceous pterosaur faunas.
Der Cambridge Greensand, eine in Ostengland aufgeschlossene Remanié-Ablagerung, hat zahlreiche Wirbeltiere aus der oberen Unterkreide (Alb) geliefert. Darunter fanden sich mehr als 2000 isolierte Pterosaurierknochen. Insgesamt wurden aus dem Greensand bis zu 32 Flugsauriertaxa beschrieben, was zu einer beträchtlichen taxonomischen und nomenklatorischen Verwirrung geführt hat, die bis heute andauert. Eine vollständige Revision erkennt 11 Arten aus drei Familien an: (1) die Ornithocheiridae (Ornithocheirus simus und vielleicht eine zweite, bislang unbenannte Art von Ornithocheirus, sowie Coloborhynchus capito, Coloborhynchus sedgwickii, Anhanguera cuvieri und Anhanguera fittoni); (2) die Lonchodectidae (Lonchodectes compressirostris, Lonchodectes machaerorhynchus, Lonchodectes microdon und Lonchodectes platystomus); und schließlich einen zahnlosen Flugsaurier (Ornithostoma sedgwicki). der zu keiner der vorgenannten Familien gehört und sich als stratigraphisch ältester Nachweis der Pteranodontidae erweisen könnte. Es ist nicht auszuschließen, dass einige der gegenwärtig erkannten Taxa eher einen ausgeprägten Sexualdimorphismus illustrieren denn taxonomisch distinkte Arten darstellen (Coloborhynchus capito und Coloborhynchus sedgwickii, Lonchodectes compressirostris und Lonchodectes machaerorhynchus) oder sogar lediglich Endpunkte einer intraspezifisch variablen Population (Ornithocheirus simus und Ornithocheirus sp., Lonchodectes compressirostris und Lonchodectes microdon). In dieser strengeren Fassung bestünden nur sieben gültige Arten, doch leider sind die Flugsaurier des Cambridge Greensand zu schlecht bekannt, um diese Fragen zu beantworten. Die Flugsaurierfauna des Cambridge Greensand ähnelt jüngeren kreidezeitlichen Faunen aus dem Lower Chalk von England. Weiter-hin enthält sie Faunenelemente, wie etwa Ornithocheiriden. die auch für zahlreiche andere Faunen der hohen Unterkreide und tiefen Oberkreide charakteristisch sind. Das Fehlen von Tapejariden und das Auftreten des anscheinend endemischen Lonchodectes sind weitere Kennzeichen des Cambridge Greensand. Die Zusammensetzung der Pterosaurierfaunen folgte offenbar ökologischen Differenzierungen und illustriert einen gewissen Provinzialismus an der Grenze Unter-Oberkreide.
doi:10.1002/mmng.20010040112
A reassessment of the systematic relationships of pterosaurs from the Lower Cretaceous Yixian Formation of Liaoning Province, China, shows that Dendrorhynchoides should be reassigned to the Anurognathidae (“Rhamphorhynchoidea”) and that Eosipterus possibly belongs within Ctenochasmatidae (Pterodactyloidea). These pterosaurs formed an integral part of a diverse community that inhabited lowland terrestrial environments in the region of northeast China in the Early Cretaceous. A new compilation of data for the Lower Cretaceous hints at a broad differentiation between pterosaurs that lived in continental habitats (anurognathids, ctenochasmatoids, dsungaripteroids) and those that frequented marine environments (ornithocheiroids). Moreover, there is evidence of further differentiation within continental habitats, between pterosaurs living in lowland and coastal regions (anurognathids. ctenochasmatoids) and those living in more inland environments (dsungaripteroids). The temporal and geographical range extensions for high rank taxa that are implied by the Yixian pterosaurs further emphasise the incompleteness and unevenness of the pterosaur fossil record and its unreliability for biostratigraphic zonation.
Eine Neubewertung der systematischen Stellung der Flugsaurier von der unterkretazischen Yixian-Formation der Provinz Liaoning, China, zeigt, dass Dendrorhynchoides den Anurognathiden (“Rhamphorhynchoidea”) zugeordnet werden kann und dass Eosipterus vermutlich zu den Ctenochasmatiden (Pterodactyloidea) gehört. Diese beiden Flugsaurier bilden einen integralen Bestandteil einer diversen Fauna, die in der Unteren Kreide ein terrestrisches Flachland-Ökosystem im Nordosten Chinas bewohnte. Fasst man die für die Untere Kreide verfügbaren Daten zusammen, so zeigt sich eine weitgehende Differenzierung zwischen Flugsauriern, die überwiegend in kontinentalen Ökosystemen lebten (Anurognathidae, Ctenochasmatoidea, Dsungaripteroidea) und jenen, die auch oft in marinen Bereichen auftreten (Ornithocheiroidea). Darüber hinaus gibt es auch Hinweise auf eine Differenzierung innerhalb der kontinentalen Habitate, zwischen Pterosauriern, die sich in den Ebenen und küstennahen Bereichen aufhielten (Anurognathidae, Ctenochasmatoidea) und den Bewohnern von mehr küstenfernen Ökosystemen (Dsungaripteroidea). Die von den Taxa der Yixian-Formation angezeigte Erweiterung der stratigraphischen und geographischen Reichweite für Taxa höheren Ranges unterstreichen die Unvollständigkeit und Unausgewogenheit des Fossilberichtes der Flugsaurier und seine Unzulänglichkeit für biostratigraphische Zonierungen.
doi:10.1002/mmng.20000030109
ABSTRACT
Cearadactylus atrox nov. gen., nov. sp.: a new Pterosauria (Pterodactyloidea) from the Ararípe-Plateau, Ceará, Brazil.
A skull with mandible ascribed to a Pterosauria (Pterodactyloidea) is described. It originates from Lower Cretaceous, Santana formation, Ararípe-Plateau, Ceará, Brazil. A new genus and species is instituted.
Classis: Reptilia Laurenti, 1768 Subclassis: Archosauria Cope, 1869 Ordo: Pterosauria Kaup, 1874 Subordo: Pterodactyloidea Plieninger, 1901 Familia indet.
Cearadactylus nov. gen.
Diagnose of the genus: large pterosaurian, but not a giant one, predator. Skull long and low. Anterior ends of both skull and man¬dible spatulated. The alveolar borders of the premaxillae do not occlude with the jaw, thus leaving a wide gap. Premaxillary and anterior dentary teeth considerably longer than the other ones cro¬oked, stretched outwards. The antero-superior border of the antorbital fenestra (fused with the external naris) is formed only by the premaxilla.
Derivatio nominis: from the state of Ceará from where the skull proceeds; and from the Greek δάχτυλος = digit, with reference to the long IV digit of the Pterosauria.
Type-species: Cearadactylus atrox nov. gen. nov. sp.
Type by monotypy
Cearadactylus atrox nov. gen. nov. sp.
Figs. 1 and 2: table.
Holotype: skull with jaw; tabular region and braincase lacking.
Depositorium: Borgomanero collection — Rua Almirante Tamandaré 915. 80000 Curitiba PR, Brazil. N° F-PV-93. [NB: the holotype is now conserved in the collection of the Departamente Nacional de Produção Mineral-DNPA, Seção de Paleontologia, Urca, Praia Vermelha, Av. Pasteur, 404 — Rio de Janeiro, Brazil]
Plastotype: cast of the holotype deposited in the Paleontological Section of the "Departamento Nacional de Produção Mineral", Av. Pasteur, 404 — Rio de Janeiro, RJ, Brazil.
Horizon, type and age: Member Romualdo, Santana Formation, Araripe Group, Lower Cretaceous, Aptian.
Type locality: Araripe-Plateau, Ceará, Brazil. The precise locality is unknown, out the specimen may possibly originate from the eas¬tern end of the Plateau.
Derivatio nominis: atrox: from the Latin, means cruel; with refe¬rence to the undoubtedly carnivorous diet s. l. (probably psicivorous), as well as to the long teeth which represented a veritable trap for the prey.
Diagnose of the species: large pterosaurian, but not a giant one (wing-span about 4 m in the adult), predator; skull showing the follo¬wing features: skull long and low, general outline, in zenithal view, elongated, birsoyd, with rostral end spatulated. The alveolar bor¬ders of the premaxillary do not occlude with the jaw, leaving a wide gap. Premaxillary dentition very long and, except the most anterior teeth, obliquely outward stretched and curved; maxillary teeth spaced, short, conical, not very sharp and slightly stretched back¬ward. The teeth of the dentary show a strictly similar disposition and pattern. All teeth alternate in the upper and inferior rows, so that they interfinger, while in the occlusion the lower ones lie exter¬nal to the maxillary, and the upper ones lie external to the mandi¬ble. Antorbital fenestra (fused with the external naris) very long, with the posterior border forming an angle of about 90° with the lower border; the maxilla does not bound its upper margin. Long groove along almost the entire length of the suture between premaxillae. Sagittal crest lacking, at least until the first third part of the antorbital opening. Base of the temporal region (between the quadrate and the posterior border of the antorbital fenestra) narrow and funnel shaped. Length of the symphysis equal to nearly one third of the mandible. Mandibular ramus deeper than the rostral portion of the skull. Anterior end of the mandible spatulated.
Out of curiosity, this pterosaur was mentioned, as animal protagonist, in the novel "Jurassic Park" by Michael Crichton (M. Crichton, Jurassic Park, Arrow, London, 1991, p. 278).
A partly preserved skull of a large pterosaur from the Santana Formation (Aptian) of the Chapada do Araripe, northeastern Brazil, is described. The skull is assigned to a new species, Cearadactylus? ligabuei nov. sp. The new species is characterized, with respect to the other described pterosaurs from Santana Formation, by a spatulate tip of the snout with 4 very large premaxillary teeth followed by much smaller maxillary teeth, premaxillae with a sharp, gibbous dorsal margin in the middle-anterior part of the snout but without a sagittal crest, and a relatively elongate jugal plate. -Author
Cearadactylus atrox, a large pterodactyloid pterosaur represented by an incomplete skull and lower jaw from the Lower Cretaceous Santana Formation of Brazil, is a valid species. Diagnostic characters include a mandibular symphysis with a transversely expanded ‘spatulate’ anterior end that is considerably wider than the rostral spatula, and a third rostral tooth that has a basal diameter more than three times that of the fifth tooth. Additional diagnostic characters, contingent upon assignment of Cearadactylus atrox to the Ctenochasmatidae, include: anterior ends of jaws divaricate and containing 7 pairs of rostral teeth and 6 pairs of mandibular teeth; marked dimorphodonty, with an abrupt change in tooth morphology; and a “high check”. ‘Cearadactylus? ligabuei’ Dalla Vecchia, 1993, based on an incomplete skull, also from the Santana Formation, is not related to Cearadactylus atrox, exhibits several ornithocheirid synapomorphies and is referred, tentatively, to Anhanguera. Cearadactylus atrox exhibits various synapomorphies of the Ctenochasmatidae (rostrum anterior to nasoantorbital fenestra greater than half total skull length, teeth in anterior part of dentition relatively elongate and pencil-shaped, premaxilla has at least 7 pairs of teeth), the defining synapomorphy of the Gnathosaurinae (rostrum with dorsoventrally compressed laterally expanded spatulate anterior expansion), and shares two synapomorphies with the Chinese gnathosaurine Huanhepterus quingyangensis (anterior tips of jaws divaricate, teeth restricted to anterior half of mandible). Two elongate cervical vertebrae, also from the Santana Formation and previously assigned to ‘Santanadactylus brasilensis’, are tentatively referred to Cearadactylus. Reconstruction of the temporal history of the Ctenochasmatidae suggests that while ctenochasmatines became increasingly specialised for filter feeding, gnathosaurines changed from sieve feeding to piscivory, acquiring several cranial characters that are similar to those of ornithocheirids, a group that also includes large aerial piscivores that used a terminal tooth grab for prey capture.
Cearadactylus atrox aus der Santana-Formation (Unterkreide, NO-Brasilien) ist eine valide Art. Eine Revision des Taxons, von dem ein unvollständiger Schädel mit Unterkiefer vorliegt, ergab folgende diagnostische (autapomorphe) Merkmale. Die Symphyse hat ein transversal verbreitertes spatelförmiges Vorderende, das deutlich breiter ist als das Schnauzenende. Der dritte rostrale Zahn erreicht einen basalen Durchmesser, der jenen des fünften Zahns um das Dreifache übertrifft. Hinzu kommen Merkmale, die C. atrox mit der Ctenochasmatidae gemein hat, darunter die vorn auseinanderklaffenden Kieferränder, sieben rostrale Zahnpaare, sechs Unterkieferzahnpaare, eine ausgeprägte Dimorphodontie sowie eine hohe Wangenregion. ‘Cearadactylus ? ligabuei’ Dalla Vecchia 1993, ebenfalls mit einem unvollständigen Schädel belegt, ist nicht näher mit C. atrox verwandt. Im Gegensatz zu letzterem zeigt ‘C. ? ligabuei’ signifikante Ähnlichkeiten mit den Ornithocheiridae. Unter Vorbehalt wird er hier der Gattung Anhanguera zugeordnet. C. atrox hat neben eindeutigen Synapomorphien der Ctenochasmatidae, z. B. erreicht das Rostrum anterior des nasoantorbitalen Fensters mehr als die halbe Schädellänge, die vordersten Zähne sind verlängert und stiftförmig und die das Prämaxillare trägt mindestens sieben Zahnpaare. Daneben besitzt C. atrox auch noch die entscheidende Synapomorphie der Gnathosaurinae, nämlich ein Rostrum mit dorsoventral komprimierter vorderem Auswuchs. Außerdem ist C. atrox gekennzeichnet durch zwei Autapomorphien des Gnathosaurinen Huanhepterus quingyangensis aus China: divergierende Schnauzenenden und Zähne begrenzt auf vordere Kieferhälfte. Schließlich werden zwei lange Halswirbel, die auch aus der Santana Formation stammen und bislang zu Santanadactylus brasiliensis gerechnet wurden, unter Vorbehalt zu Cearadactylus gestellt. Die Evolutionsgeschichte der Ctenochasmatidae ist durch eine zunehmende Spezialisierung auf filternde Ernährungsweise gekennzeichnet. Die Gnathosaurinen dagegen stellten sich von der filternden auf eine piscivore Ernährung um, wobei sie eine Reihe von Schädelmerkmalen erworben haben, die den Ornithocheiriden konvergent ähnlich ist.
The Cambridge Greensand, a remanié deposit that crops out in Cambridgeshire, eastern England, has yielded numerous, though fragmentary, late Early Cretaceous (Albian) vertebrate fossils including more than 2000 isolated pterosaur bones. So far, 32 species of pterosaur have been proposed in connection with the Cambridge Greensand material, but there has been and continues to be considerable confusion concerning the validity of these taxa, their relationships to each other and to other pterosaurs, and the material upon which they were established. A comprehensive systematic revision identified eleven valid species distributed among three families: the Ornithocheiridae (Ornithocheirus simus and possibly a second, as yet unnamed species of Ornithocheirus, Coloborhynchus capito, Coloborhynchus sedgwickii, Anhanguera cuvieri, and Anhanguera fittoni); the Lonchodectidae (Lonchodectes compressirostris, Lonchodectes machaerorhynchus, Lonchodectes microdon and Lonchodectes platystomus); and a species of edentulous pterosaur (Ornithostoma sedgwicki) that may represent the earliest record for the Pteranodontidae. It is possible that some of the taxa currently recognised represent sexual dimorphs (Coloborhynchus capito and Coloborhynchus sedgwickii, Lonchodectes compressirostris and Lonchodectes machaerorhynchus), or disjunct populations of a single species (Ornithocheirus simus and Ornithocheirus sp., Lonchodectes compressirostris and Lonchodectes microdon) and that there may be as few as seven valid species, but the Cambridge Greensand pterosaurs are too poorly known to demonstrate this at present. The Cambridge Greensand pterosaur assemblage is similar to a slightly younger, but much smaller assemblage from the Lower Chalk of England and shares some elements, such as ornithocheirids, in common with many other late Early and early Late Cretaceous assemblages. It is distinguished by the absence of tapejarids and the presence of Lonchodectes which, so far, is only known from the Cretaceous of England. The disparity in taxonomic composition is possibly related to ecological differentiation, and might also reflect some provincialism in late Early and early Late Cretaceous pterosaur faunas.Der Cambridge Greensand, eine in Ostengland aufgeschlossene Remanié-Ablagerung, hat zahlreiche Wirbeltiere aus der oberen Unterkreide (Alb) geliefert. Darunter fanden sich mehr als 2000 isolierte Pterosaurierknochen. Insgesamt wurden aus dem Greensand bis zu 32 Flugsauriertaxa beschrieben, was zu einer beträchtlichen taxonomischen und nomenklatorischen Verwirrung geführt hat, die bis heute andauert. Eine vollständige Revision erkennt 11 Arten aus drei Familien an: (1) die Ornithocheiridae (Ornithocheirus simus und vielleicht eine zweite, bislang unbenannte Art von Ornithocheirus, sowie Coloborhynchus capito, Coloborhynchus sedgwickii, Anhanguera cuvieri und Anhanguera fittoni); (2) die Lonchodectidae (Lonchodectes compressirostris, Lonchodectes machaerorhynchus, Lonchodectes microdon und Lonchodectes platystomus); und schließlich einen zahnlosen Flugsaurier (Ornithostoma sedgwicki). der zu keiner der vorgenannten Familien gehört und sich als stratigraphisch ältester Nachweis der Pteranodontidae erweisen könnte. Es ist nicht auszuschließen, dass einige der gegenwärtig erkannten Taxa eher einen ausgeprägten Sexualdimorphismus illustrieren denn taxonomisch distinkte Arten darstellen (Coloborhynchus capito und Coloborhynchus sedgwickii, Lonchodectes compressirostris und Lonchodectes machaerorhynchus) oder sogar lediglich Endpunkte einer intraspezifisch variablen Population (Ornithocheirus simus und Ornithocheirus sp., Lonchodectes compressirostris und Lonchodectes microdon). In dieser strengeren Fassung bestünden nur sieben gültige Arten, doch leider sind die Flugsaurier des Cambridge Greensand zu schlecht bekannt, um diese Fragen zu beantworten. Die Flugsaurierfauna des Cambridge Greensand ähnelt jüngeren kreidezeitlichen Faunen aus dem Lower Chalk von England. Weiter-hin enthält sie Faunenelemente, wie etwa Ornithocheiriden. die auch für zahlreiche andere Faunen der hohen Unterkreide und tiefen Oberkreide charakteristisch sind. Das Fehlen von Tapejariden und das Auftreten des anscheinend endemischen Lonchodectes sind weitere Kennzeichen des Cambridge Greensand. Die Zusammensetzung der Pterosaurierfaunen folgte offenbar ökologischen Differenzierungen und illustriert einen gewissen Provinzialismus an der Grenze Unter-Oberkreide.
A reassessment of the systematic relationships of pterosaurs from the Lower Cretaceous Yixian Formation of Liaoning Province, China, shows that Dendrorhynchoides should be reassigned to the Anurognathidae (“Rhamphorhynchoidea”) and that Eosipterus possibly belongs within Ctenochasmatidae (Pterodactyloidea). These pterosaurs formed an integral part of a diverse community that inhabited lowland terrestrial environments in the region of northeast China in the Early Cretaceous. A new compilation of data for the Lower Cretaceous hints at a broad differentiation between pterosaurs that lived in continental habitats (anurognathids, ctenochasmatoids, dsungaripteroids) and those that frequented marine environments (ornithocheiroids). Moreover, there is evidence of further differentiation within continental habitats, between pterosaurs living in lowland and coastal regions (anurognathids. ctenochasmatoids) and those living in more inland environments (dsungaripteroids). The temporal and geographical range extensions for high rank taxa that are implied by the Yixian pterosaurs further emphasise the incompleteness and unevenness of the pterosaur fossil record and its unreliability for biostratigraphic zonation.
A well-preserved pterosaur with nearly complete skull is described from the Lower Cretaceous Yixian Formation at Sihetun in
western Liaoning. It is characterized by a low and long crestless skull, slender and pointed teeth, long metacarpal, nearly
equal length of metatarsals I–III and short pedal digit V. It is referred to a new genus and species of the family Pterodactylidae:Haopterus gracilis gen. et sp. nov. This is the first pterosaur with a nearly complete skull from the Jehol Biota; it also represents the first
non-controversial fossil record of Pterodactylidae in Asia.Haopterus is more derived thanPterodactylus from the Late Jurassic Solnhofen in Germany. This discovery extends the distribution of the family Pterodactylidae from Europe
and Africa to Asia and its latest occurrence from the Late Jurassic to the Early Cretaceous. The discovery ofHaopterus gracilis provides further evidence for the study of the origin and radiation of the Jehol Biota; it also sheds new light on the evolution
and distribution of pterosaurs in the late Mesozoic.
Most of the toothed pterosaurs recovered from the Araripe Basin in Brazil (Santana Formation) have premaxillary sagittal and dentary sagittal crests. Some clear differences (and various less clear features) between the crested taxa have been used to classify the fossils, resulting in much scientific debate. On the other hand, a few potentially important features have been largely neglected so far. The present work presents an update of one of these crested taxa, Coloborhynchus, discussing the dentition and other previously unnoticed features in order to evaluate the systematic position of this taxon.
The discovery of a previously undescribed pterosaur, Thalassodromeus sethi, yields information on the function of cranial crests and the feeding strategy developed by these extinct flying reptiles.
The material consists of a large skull (length: 1420 millimeters, including the crest) with a huge bony crest that was well
irrigated by blood vessels and may have been used for regulation of its body temperature. The rostrum consists of two bladelike
laminae, the arrangement of which is analogous to the condition found in the bird Rynchops, which skims over the water to catch food, indicating that T. sethialso may have been a skimmer.
A large number (> 1 per cent.) of fossil fish from the Santana Formation (?Albian, Lower Cretaceous) of north-east Brazil, occurs at a high angle, or even vertical, to bedding. Petrographic and taphonomic analyses of such occurrences indicate that the laminated argillaceous sediments were soupy and allowed sinking dead fish to penetrate the sediment to depths greater than the total body length of the fish. Density contrasts between the dead fish and the surrounding sediment were negligible, and penetration was largely a result of acceleration due to the high density contrast between the fish and the water column. A commonly recurrent angle of fish to bedding of approximately 45° reflects the gliding angle achieved during forward descent with pectoral fins extended.
A statistical and morphological study of the pterosaur Rhamphorhynchus from the Solnhofen Limestone shows that specimens fall into discrete size-classes that are year-classes resulting from seasonal mortality or preservation of specimens. Taxonomic characters used in the past to separate species of Rhamphorhynchus are all related to size and ontogeny, and all specimens belong to a single species, R. muensteri. Thus, the collections of Rhamphorhynchus provide an unequaled record of the growth and development of a Jurassic rhamphorhynchoid pterosaur. Rhamphorhynchus did not have rapid determinate growth; the growth rate was comparable to that of extant crocodilians, and growth continued over the course of at least three years after individuals began to fly. -Author
A new species of pterosaur, Coloborhynchus spielbergi sp. nov. (Pterodactyloidea), from the Romualdo Member (Albian) of the Santana Formation is described. The type consists of the skull, mandible and many of the post-cranial bones. The specimen displays a high degree of co-ossification indicating that the animal was an adult and likely quite old when it died. The wingspan is reconstructed at nearly 6 m. Among the characteristic features are a large anteriorly positioned premaxillary sagittal crest and a smaller, also anteriorly positioned dentary sagittal crest, a flat anterior aspect of the skull from which two teeth project and a ventrally fused pelvis. Comments on Brazilian pterosaurs are made in connection with the classificiation of the Leiden specimen.
The Romualdo Member of the Santana Formation (Araripe Basin, northeastern Brazil) is famous for the abundance and the exceptional preservation of the fossils found in its early diagenetic carbonate concretions. However, a vast majority of these Early Cretaceous fossils lack precise geographical and stratigraphic data. The absence of such contextual proxies hinders our understanding of the apparent variations in faunal composition and abundance patterns across the Araripe Basin.We conducted controlled excavations in the Romualdo Member in order to provide a detailed account of its main stratigraphic, sedimentological and palaeontological features near Santana do Cariri, Ceará State.We provide the first fine-scale stratigraphic sequence ever established for the Romualdo Member and we distinguish at least seven concretion-bearing horizons. Notably, a 60-cm-thick group of layers (“Matracão”), located in the middle part of the member, is virtually barren of fossiliferous concretions.Moreover, a sample of 233 concretions shows that (i) the stratigraphic distribution of the concretions is very heterogeneous and their density varies from 0.8 to 15 concretions/m3; (ii) concretions have a preferential, bimodal orientation pattern (major NW–SE axis and secondary ≈N–S axis) throughout the section, suggestive of permanent palaeocurrents of low energy; (iii) few concretions yield the well-preserved vertebrates that have made the Romualdo Member so famous, and those are mainly restricted to four stratigraphic horizons; (iv) only six fish taxa were recovered, the most common being Vinctifer and Tharrhias, followed by Cladocyclus, whereas Brannerion, Calamopleurus (=Enneles) and Notelops are rare. No tetrapod was found in the sample; (v) there is a strong stratigraphic control in the distribution of these taxa and one can distinguish at least three major assemblages at the same locus. These are, from older to younger, a Tharrhias-dominated assemblage, an assemblage dominated by Tharrhias and by Cladocyclus, and a Vinctifer-dominated assemblage. The stratigraphic sequence of these assemblages also corresponds to their ranking in terms of diversity (richness and evenness); (vi) previous accounts on the taxonomic composition and relative abundance of fossils from the Romualdo Member were severely biased toward well-preserved and exotic fossils. They are therefore inappropriate for drawing palaeoecological inferences.The factors responsible for the variations in faunal composition and abundance patterns across the Araripe Basin remain largely unknown, and we hypothesize that climate and/or palaeogeography might be the major forcing agents. Only fine-scale stratigraphic and palaeontological investigations have the potential to solve this issue. In turn, this work marks the first step of an expanded research program that aims at explaining the spatio-temporal relationships between palaeocommunities and their palaeoenvironment in the Araripe Basin during Aptian/Albian times.
Statistical analyses of measurement data and morphological studies of pterosaur fossils from the Upper Jurassic Solnhofen Limestone of Germany reveal the presence of year-classes resulting from repeated seasonal sampling of pterosaur populations. Year-classes of Rhamphorhynchus had previously been named as separate species. Similar year-classes of nominal species of Pterodactylus indicate that size-classes of some pterodactyloid species also had been named as separate species or genera. Lumping the nominal species reduces the Solnhofen pterosaur fauna to a small number of large species, and suggests that some long-held ideas about pterosaur evolution are incorrect.
I have recently obtained from the upper chalk of Kent some remains of a large species of Pterodactylus. The bones consist of—
• 1. The fore part of the head as far as about the middle of the cavitas narium, with a corresponding portion of the under jaws,—many of the teeth remaining in their sockets. (See Plate, fig. 1.)
• 2. A fragment of a bone of the same animal, apparently a part of the coracoid. (Fig. 2.)
• 3. A portion of what appears to be one of the bones of the auricular digit, from a chalk-pit at Halling. (Fig. 3.)
• 4. A portion of a similar bone, from the same locality as No. 1. (Fig. 4.)
• 5. The head of a long bone, probably the tibia, belonging to the same animal as the head No. 1. (Fig. 5.)
• 6. A more perfect bone of the same description, not from the same animal, but found at Halling. (Fig. 6.)
The latter specimen appears to me to be the same description of bone as that described by Professor Owen in the Geological Transactions (2nd Ser. vol. vi. p. 411, and pl. 39. fig. 1). The mutilated condition of the figured specimen would not allow Professor Owen to speak of its identity with the bird tribe with great certainty, and he at the same time points out its discrepant characters.
This paper is concerned with the famous fossil-bearing carbonate concretions of the Romualdo Member of the Santana Formation Konservat Lagerstätten of north-east Brazil. This palaeontologically important horizon was first dated as Cretaceous by the French palaeoichthyologist Louis Agassiz on the basis of fish fossils obtained by Bavarian explorers Spix and Martius between 1817 and 1820 and Scottish botanist and explorer George Gardner between 1836 and 1841. Gardner equated the concretion level with the English Albian ‘Upper Greensands’ on the basis of an imagined similarity of stratigraphic sequence with that of the Isle of Wight, southern England. Since then high precision dating of this remarkable deposit has proved elusive and the concretion-bearing part of the Santana Formation has been variously dated as early Late Cretaceous or late Early Cretaceous. Attempts at greater precision over the last 30 years have cited its age variously as Aptian, Albian or possibly Cenomanian, but few reliable data have been presented to support these dates.
FOSSILIZED impressions of pterosaur wing membranes are known from a number of localities1,2, but true soft-tissue preservation is extremely rare3,4. We present the first description of the internal anatomy of the wing membrane, based on exceptionally well preserved soft tissues from the forearm of a Lower Cretaceous, Brazilian pterosaur. A thin epidermis overlies a dermis composed successively of a 'stratum vasculosum', 'stratum spongiosum' and a layer of striated muscle. This exceptional specimen provides important new insights into pterosaur biology. Incipient wrinkles and an apparent lack of stiffening fibres suggest the proximal region of the wing membrane was thin, extensible and tensioned by the hind limb5,6. Excess metabolic heat resulting from rigorous activity could be lost by vasodilation of the vascular layer, a mechanism consistent with active flight in pterosaurs7-9.
Pterosaur remains from Wealden strata of southern England have largely been referred to the Ornithocheiroidea, with only a solitary controversial claim of a lonchodectid providing evidence of heightened diversity. A reappraisal of a historic Wealden specimen suggests that “Palaeornis” cliftii Mantell, 1844, an isolated humerus from the Hastings Beds Group of West Sussex, is not an ornithocheiroid as previously reported but instead confirms the presence of lonchodectid pterosaurs in the British Wealden. The diversity of British Wealden pterosaurs is heightened further by a recently-discovered pterosaur humerus from the Wealden Group of the Isle of Wight, providing the first record of azhdarchoid pterosaurs in the British Lower Cretaceous. This specimen is thought to represent a non-azhdarchid neoazhdarchian and, being from Barremian deposits, represents the earliest known occurrence of such a pterosaur.
Abstract Based on a nearly complete skeleton with skull from the Early Cretaceous of Liaoning Province, a new ornithocheirid pterosaur: Boreopterus cuiae gen. et sp. nov. is erected. Boreopterus cuiae is different from other pterosaurs preserved with skulls known from the western Liaoning Province and its neighboring areas. This new pterosaur has more and larger teeth than those in other ornithocheirids. Its anterior nine pairs of teeth are larger than other teeth. The fourth pair of upper and lower teeth are slightly larger than the third pair. Overall, Boreopterus cuiae shows much small range of tooth size variation than Anhanguera piscator and Coloborhynchus robustus. The new taxon shares with other ornithocherids in having a relatively large size of the third and fourth pairs of teeth.
The anterior tips of associated upper and lower jaws of a pterosaur from the Lower Cretaceous of Brazil are described and
assigned to the taxonColoborhynchus in the family Ornithocheiridae. It is characterized by the shape and position of the sagittal crest on the upper and lower
jaw, the arrangement and length of the teeth and the spoon-like lateral expansion of the anterior parts of the jaws. It closely
resemblesColoborhynchus wadleighi from North America andColoborhynchus clavirostris from England. Diagnostic anatomical characteristics permit a revision of the genusTropeognathus, which is shown here to be a junior synonym of other described taxa.Tropeognathus mesembrinus is referred toCriorhynchus andT. robustus toColoboryhnchus. Consistent anatomical features enable the new jaw fragment to be assigned toColoborhynchus robustus.
Das anteriore Ober- und Unterkieferfragment eines Pterosauriers aus der Unteren Kreide von Brasilien wird beschrieben und
zur GattungColoborhynchus in die Familie Ornithocheiridae gestellt. Die charakteristischen antomischen Eigenschaften sind der sagitale Kamm am Ober-
und Unterkiefer, die Anordnung und Länge der Zähne und die löffeiförmige laterale Verbreiterung der anterioren Kieferenden.
Das Stück ähnelt starkColoborhynchus wadleighi aus Nordamerika undColoborhynchus clavirostris aus England. Die diagnostischen morphologischen Eigenschaften des beschriebenen Stückes ermöglichen eine Revision der GattungTropeognathus, die synonym mit anderen Gattungen ist.Tropeognathus mesembrinus wird in die GattungCriorhynchus, T. robustus in die GattungColoborhynchus gestellt. Aufgrund der starken anatomischen Übereinstimmung kann das beschriebene Fragment alsColoborhynchus robustus identifiziert werden.
A new species of the sail-crested pterosaur Tupuxuara is described from the Santana Formation of Brazil, Tupuxuara deliradamus sp. nov. The holotype, a partial skull, and a larger, partial skull referred to the same taxon differs from Tupuxuara leonardii by having a nasoantorbital fenestra with an acutely-angled posterior border with a long, straight posterodorsal margin, a reclined cranium, and an orbit situated entirely in the ventral half of the nasoantorbitral fenestra. Unfortunately, neither specimen is comparable with the fragmentary rostrum representing Tupuxuara longicristatus. In addition, resolution of a recent nomenclatural problem over the correct name for the clade containing Tupuxuara and its sister taxon, Thalassodromeus, is provided. Both genera are used by different authors as the nomenclatural basis for the group, but “Tupuxuaridae” has never been explicitly erected as a new taxon, and therefore fails to meet ICZN criteria that new taxa are only valid if authors clearly indicate their intention to establish new names. By contrast, “Thalassodrominae” was explicitly erected as a name for the Thalassodromeus + Tupuxuara clade, thereby fulfilling all ICZN requirements for naming of a new taxon and making Thalassodromeus stand as the type genus for this group.
A portion of pterosaur skull from the Romualdo Member of the Santana Formation (?Albian–?Turonian, Cretaceous) of north-east Brazil provides new data on the morphology and ontogeny of azhdarchoid pterosaur cranial crests. The specimen consists of parts of the cranial bones posterodorsal to the nasoantorbital fenestra, including partial nasals, lacrimals, frontals and possibly the parietals. A posterodorsally directed premaxillary crest with a concave posterior border is located dorsal to the posterior border of the nasoantorbital fenestra. A well-defined suture indicates overlapping, posterodorsally directed growth of the premaxilla over the skull roof, suggesting that the generation of the premaxillary crest is a late ontogenetic feature and thus probably related to sexual display. The systematics of Tupuxuara and its relationship to other azhdarchoids is reviewed and a cladistic analysis of the group is presented. Tupuxuara is found to be the sister-taxon to Azhdarchidae. Tupuxuara longicristatus Kellner and Campos, 1988 is argued to be the only valid named species in this genus and Thalassodromeus Kellner and Campos, 2002 is considered a junior subjective synonym of this taxon. As originally conceived, Tapejaridae is paraphyletic: a new, more restrictive version of Tapejaridae (including Tapejara and Sinopterus dongi) might exist, but its monophyly is weakly supported. Furthermore, Tapejara was found to be paraphyletic in all trees.
Previous cladistic studies of pterosaur relationships suffer from restricted numbers of taxa and characters, incomplete data sets and absence of information on characters, tree structure and the robustness of trees. Parsimony analysis of a new character data set (60 characters, 20 terminal taxa, 93.75% complete) yielded six trees. In the strict consensus tree Preondactylus is the most basal taxon followed, stepwise, by the Dimorphodontidae and the Anurognathidae. Beyond this basal group, more derived pterosaurs (Campylognathoididae (Rhamphorhynchidae + Pterodactyloidea)) share a suite of characters principally associated with elongation of the rostrum. The Pterodactyloidea consists of four major clades. The Ornithocheiroidea is the most basal taxon consisting, stepwise, of Istiodactylus, the Ornithocheiridae, Nyctosaurus and the Pteranodontidae. The remaining taxa, Ctenochasmatoidea, Dsungaripteroidea and Azhdarchoidea, are weakly united in a clade of non-ornithocheiroid pterodactyloids, but their inter-relationships are difficult to resolve. Cycnorhamphus is the basal-most ctenochasmatoid, while the remaining taxa (Pterodactylus, Lonchodectidae, Ctenochasmatidae) form an unresolved trichotomy. The Dsungaripteroidea (Germanodactylus + Dsungaripteridae) is strongly supported by features of the skull and dentition. The Azhdarchoidea (Tapejara [Tupuxuara + Azhdarchidae]) is united by cranial characters such as elevation of the antorbital region, and relative shortening of the wing finger. The pattern of pterosaur evolution suggested by the results of this analysis is broadly similar to traditional ideas, but has greater resolution, more complexity and reveals several previously unrecognized 'events'.