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A new species of Tupuxuara (Thalassodromidae, Azhdarchoidea) from the Lower Cretaceous Santana Formation of Brazil, with a note on the nomenclature of Thalassodromidae

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Abstract

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.

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... Indeterminate ornithocheirids reported from the Crato Formation include an isolated tooth (MN 4798-V; Sayão and Kellner, 2000;Unwin and Martill, 2007), a near complete postcranial skeleton (SMNK PAL 3854; Elgin and Frey, 2012), and incomplete wings (MPSC R-739 and MPSC R-779) which were originally interpreted as tapejarids by Nuvens et al. (2002) but were later regarded as Martill et al. (1993). Anhanguera and Ornithocheiridae redrawn from Wellnhofer (1991a), Conchostraca redrawn from Kozur and Weems (2010), Santanadactylus and Tropeognathus redrawn from Wellnhofer (1991c), Tapejara redrawn from Wellnhofer and Kellner (1991), Tupuxuara redrawn from Witton (2009) subadult ornithocheirids by Unwin and Martill (2007). Both partial wings have yet to be formally described. ...
... Before the first pterosaur remains were reported from the lagerstätten of the Romualdo Formation by Price (1971), this stratigraphic horizon was predominantly known for yielding many different species of fish (Brito and Yabumoto, 2011;Martill, 1988). The assemblage is dominated by ornithocheirids (Bantim et al., 2014;Kellner and Tomida, 2000;Pinheiro and Rodrigues, 2017;Vila Nova et al., 2014;Wellnhofer, 1985;Wellnhofer, 1991a), with tapejarids (Eck et al., 2011;Elgin and Campos, 2012;Wellnhofer and Kellner, 1991), and thalassodromids also present (Kellner et al., 1988;Kellner et al., 1994;Kellner et al., 2002;Veldmeijer et al., 2005b;Witton, 2009). ...
... Tupuxuara leonardii differs from Tupuxuara longicristatus based on the presence of a strong, medial ridge on the palate that does not extend to the anterior portion of the skull . Other cranial material referred to Tupuxuara leonardii includes a complete skull (IMCF 1052) and fragmentary yet almost complete skull of a reported juvenile (SMNK PAL 4330; Witton, 2009). Although Martill and Naish (2006) regarded Tupuxuara leonardii as a junior synonym of Tupuxuara longicristatus, on the basis that the palatal ridge varied because of individual variation in azhdarchoids, this claim was refuted by Kellner et al. (2007a). ...
Article
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.
... imperator and T. navigans [53]) from the Crato Formation (?Aptian) and Tupuxuara (T. longicristatus [54], T. leonardii and T. deliradamus [55]), Tapejara wellnhoferi and Thalassodromeus sethi from the younger Romualdo Formation (Albian) of the Araripe Basin. As is usual in Crato Formation fossils, the specimens referred to T. imperator are laterally compressed, with no information whatsoever on palatal anatomy. ...
... As can be observed in the holotypes of T. longicristatus and T. leonardii, a median keel (which is much more developed in T. leonardii) originates at the anterior part of the rostrum and broadens posteriorly, where the palate becomes increasingly convex [10,54] (Figure 3, B). The specimen IMCF 1052, illustrated in lateral aspect by Veldmeijer [60] and Witton [55], demonstrates that the palate of T. leonardii remains convex throughout its entire length, with the suborbital and subtemporal fenestrae being easily distinguishable in lateral view. This is also the condition described by Witton [55] for T. deliradamus. ...
... The specimen IMCF 1052, illustrated in lateral aspect by Veldmeijer [60] and Witton [55], demonstrates that the palate of T. leonardii remains convex throughout its entire length, with the suborbital and subtemporal fenestrae being easily distinguishable in lateral view. This is also the condition described by Witton [55] for T. deliradamus. Although the morphology of the palate where the palatal openings are located is still unknown for T. longicristatus, the holotype (MN 6591 V) shows strongly convex maxillary palatal plates below the nasoantorbital fenestrae (Figure 3, B), making it likely that the condition in T. longicristatus was similar to that observed in other Tupuxuara species. ...
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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.
... Tapejarids are represented by a single jaw fagment, BSP 1997 I 67, bearing the remains of a prominent midline crest [15]. This fragment has generally been interpreted as part of the mandibular symphysis [15,22], but might pertain to the rostrum, in which case it is worth noting that the jaw and crest morphology of the Kem Kem specimen is consistent with that reported for tapejarids ([23] fig. 3) and to some extent tupuxuarids ( [24] fig. 3). ...
... Alanqa saharica shares four unique features in common with neoazhdarchians (Thalassodromidae + Azhdarchidae): seen in lateral view the rostrum is always deeper than the mandibular symphysis at any corresponding point along the jaws ( [24,46,53], Habib pers. comm. ...
... comm. 2010); the mandibular symphysis is long, straight and low (Figures 2, 3; [24,45,53]); the mandibular symphysis has a low, elongate, blade-like ventral keel; and the cross-sectional profile of the mandibular symphysis is triangular near the tip of the jaw, but becomes Y-shaped posteriorly. ...
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The Kem Kem beds in South Eastern Morocco contain a rich early Upper (or possibly late Lower) Cretaceous vertebrate assemblage. Fragmentary remains, predominantly teeth and jaw tips, represent several kinds of pterosaur although only one species, the ornithocheirid Coloborhynchus moroccensis, has been named. Here, we describe a new azhdarchid pterosaur, Alanqa saharica nov. gen. nov. sp., based on an almost complete well preserved mandibular symphysis from Aferdou N'Chaft. We assign additional fragmentary jaw remains, some of which have been tentatively identified as azhdarchid and pteranodontid, to this new taxon which is distinguished from other azhdarchids by a remarkably straight, elongate, lance-shaped mandibular symphysis that bears a pronounced dorsal eminence near the posterior end of its dorsal (occlusal) surface. Most remains, including the holotype, represent individuals of approximately three to four meters in wingspan, but a fragment of a large cervical vertebra, that probably also belongs to A. saharica, suggests that wingspans of six meters were achieved in this species. The Kem Kem beds have yielded the most diverse pterosaur assemblage yet reported from Africa and provide the first clear evidence for the presence of azhdarchids in Gondwana at the start of the Late Cretaceous. This, the relatively large size achieved by Alanqa, and the additional evidence of variable jaw morphology in azhdarchids provided by this taxon, indicates a longer and more complex history for this clade than previously suspected.
... The posterior part of the palate of Dsungaripterus has more similarities with the azhdarchoids Caupedactylus ybaka (Kellner, 2013) and Tupuxuara leonardii IMCF 1052 (Witton, 2009;Pinheiro & Schultz, 2012) (Fig. 6). All show the ectopterygoid overlaying the pterygoid dorsally and fused with the medial pterygoid process posteriorly. ...
... A small foramen is present at the contact area between the medial pterygoid process and the posterior part of the ectopterygoid in Dsungaripterus, while in Tupuxuara and Caupedactylus there is a fenestra. The interpterygoid fenestra of Dsungaripterus differ from the 'heart-shape' outline of Caupedactylus ybaka (Kellner, 2013) and Tupuxuara leonardii (Witton, 2009;Pinheiro & Schultz, 2012), by being irregular with two symmetrical posterior notches. ...
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Pterosaur specimens with complete and well-preserved palatal region are rare. Here we describe new and previously collected specimens of the pterodactyloid pterosaur Dsungaripterus weii that are three-dimensionally preserved and provide new anatomical information for this species. Among the unique features is a lateral process of the pterygoid divided into two parts: an anterior thin, parabolic arc shaped element that separates the secondary subtemporal and the subtemporal fenestrae, followed by a dorsoventrally flattened portion that is directed inside the subtemporal fenestrae. The interpterygoid fenestrae join forming an irregular oval shape with two symmetrical posterior notches and a smooth anterior margin. Among all pterosaurs where the palate is known, the posterior configuration of the palate of D. weii is similar to some azhdarchoids, which is consistent with the suggested phylogenetic position of the Dsungaripteridae as closely related to the Azhdarchoidea. Furthermore, we identify symmetrical grooves on the lateral surface of the upper and lower jaws, that likely represent the impression of the edge of a keratinous sheath that would cover the upturned toothless rostrum during foraging activity, most likely consisting of hard elements, as has been previously assumed. Wear facets on the teeth also support this feeding mode.
... Apatorhamphus gyrostega displays several features found in Azhdarchoidea: the rostrum is deeper than the mandible in lateral view (Cai and Wei, 1994;Witton, 2009); and both the rostrum and mandible are relatively long and straight (Kellner and Langston, 1996;Witton, 2009), although there is a slight dorsal curvature of the holotype rostrum of A. gyrostega rostrum posteriorly. ...
... Apatorhamphus gyrostega displays several features found in Azhdarchoidea: the rostrum is deeper than the mandible in lateral view (Cai and Wei, 1994;Witton, 2009); and both the rostrum and mandible are relatively long and straight (Kellner and Langston, 1996;Witton, 2009), although there is a slight dorsal curvature of the holotype rostrum of A. gyrostega rostrum posteriorly. ...
Article
A new genus and species of edentulous pterodactyloid pterosaur with a distinctive partial rostrum from the mid-Cretaceous (?Albian/Cenomanian) Kem Kem beds of southeast Morocco is described. The taxon is assigned to Chaoyangopteridae based upon its edentulous jaws, elongate rostrum and slightly concave dorsal outline. The rostral cross-section is rounded dorsally and concave on the occlusal surface. The lateral margins are gently convex dorsally becoming slightly wider toward the occlusal border, and a row of small lateral foramina parallel to the dorsal margin determines it as a taxon distinct from other chaoyangopterids. Apatorhamphus gyrostega gen et sp. nov. is a pterosaur of medium to large size (wingspan likely somewhere between ~3 m and ~7 m). This new species brings the number of named Kem Kem azhdarchoids to three, and the number of named Kem Kem pterosaurs to five, indicating a high pterosaur diversity for the Kem Kem beds.
... In contrast, the Thalassodrominae are so far exclusively known from the Romualdo Formation, comprising only two genera: Thalassodromeus and Tupuxuara Kellner and Campos, 1988, the latter with two valid species, Tupuxuara longicristatus Kellner and Campos, 1988, and Tupuxuara leonardii Campos, 1994 (see Kellner, 2013). These three taxa have been suggested to represent a single species (Martill and Naish, 2006), but this proposal was solidly refuted (Kellner and Campos, 2007), and the valid status of these taxa is currently favored (e.g., Witton, 2009Witton, , 2013Kellner, 2013;Andres et al., 2014;Vremir et al., 2015). A third species of Tupuxuara has also been proposed, Tupuxuara deliradamus Witton, 2009, later considered a nomen dubium (Kellner, 2013). ...
... In some areas, particularly at the dorsal region, the bone is only about 1 mm in thickness. Some fragmented areas show signs of pneumatization, with a trabecular internal structure (for further details on this structure, see Campos, 2002b, 2007), similar to the holotypes of 'Tupuxuara deliradamus' and Caupedactylus ybaka (Witton, 2009;Kellner, 2013). Only the left lateral side of the crest is exposed, with the right side obscured by the calcareous matrix. ...
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Thalassodromeus sethi (Pterodactyloidea, Tapejaridae, Thalassodrominae) is a unique pterosaur from the Romualdo Formation, Araripe Basin (Early Cretaceous, Albian). A large sagittal cranial crest (the largest known ossified crest of any pterosaur) and unusual toothless bladed jaws constitute some of its outstanding anatomical features. Its holotype (DGM 1476-R, almost complete skull and mandible in three dimensions) represents one of the largest pterosaur skulls ever found. Here, we provide a detailed osteological redescription of the holotype, which has only been preliminarily described. We also describe and depict for the first time some skeletal regions of Th. sethi, including the occiput, the palatal openings, and a dentary fragment. Finally, in the light of new information concerning this species, we revisited the specimen NMSG SAO 251093 (an incomplete mandible), also from the Romualdo Formation and originally referred to Th. sethi but recently redescribed as a new species of dsungaripterid pterosaur named Banguela oberlii. Here, the analysis of a cast (MN 4703-V) and its inclusion in a phylogenetic analysis recovered NMSG SAO 251093 within the Thalassodrominae, as a sister taxon of Th. sethi and indeed different from it at the species level. We hereby rename it Thalassodromeus oberlii, comb. nov. These considerations provide new data for discussions concerning the morphology of the Tapejaridae, the diversity of the Araripe pterosaur fauna, and the complex evolution of the pterodactyloid palatal region, as well as data for future morphofunctional studies of Thalassodromeus. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP Citation for this article: Pêgas, R. V., F. R. Costa, and A. W. A. Kellner. 2018. New information on the osteology and a taxonomic revision of the genus Thalassodromeus (Pterodactyloidea, Tapejaridae, Thalassodrominae). Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2018.1443273.
... Here I accept the synonymy of Tupuxuara longicristatus, Tupuxuara leonardi, and Thalassodromeus sethi proposed by Martill and Naish (2006). Tupuxuara deliradamus, described by Witton (2009), is also added to this list of synonyms. T. longicristatus is known from several specimens, including mostly undescribed skeletons and skulls, from the Albian Romualdo Formation in Ceará, Brazil (Kellner and Hasegawa 1993;Kellner and Campos 1994Kellner 2004;Veldmeijer et al. 2005;Martill and Naish 2006;Witton 2009;Aires et al. 2014). ...
... Tupuxuara deliradamus, described by Witton (2009), is also added to this list of synonyms. T. longicristatus is known from several specimens, including mostly undescribed skeletons and skulls, from the Albian Romualdo Formation in Ceará, Brazil (Kellner and Hasegawa 1993;Kellner and Campos 1994Kellner 2004;Veldmeijer et al. 2005;Martill and Naish 2006;Witton 2009;Aires et al. 2014). The two cervical vertebrae from the Romualdo Formation, GIUA 4895, the "paratype" of Santanadactylus brasilensis (Buisonje 1980), likely belong to this species as well. ...
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Citation: Averianov A (2014) Review of taxonomy, geographic distribution, and paleoenvironments of Azhdarchidae (Pterosauria) ZooKeys 432: 1–107. doi: 10.3897/zookeys.432.7913 Abstract The taxonomy, geographic distribution, and paleoenvironmental context of azhdarchid pterosaurs are reviewed. All purported pteranodontid, tapejarid, and azhdarchid specimens from the Cenomanian Kem Kem beds of Morocco are referred to a single azhdarchid taxon, Alanqa saharica. The four proposed autapomorphies of Eurazhdarcho langendorfensis from the lower Maastrichtian Sebeş Formation of Ro-mania are based on misinterpretations of material and this taxon is likely a subjective junior synonym of Hatzegopteryx thambema. Among 54 currently reported azhdarchid occurrences (51 skeletal remains and 3 tracks) 13% are from lacustrine deposits, 17% from fluvial plain deposits, 17% from coastal plain depos-its, 18% from estuarine and lagoonal deposits, and 35% from costal marine deposits. Azhdarchids likely inhabited a variety of environments, but were abundant near large lakes and rivers and most common in nearshore marine paleoenvironments. Copyright Alexander Averianov. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. A peer-reviewed open-access journal
... Tupuxuara spp. (Witton 2009), Quetzalcoatlus sp. (Kellner and Langston 1996)) or down-turned (Thalassodromeus, Figure 3(J), as in Tapejara). ...
... Rhamphorhynchus sp. (Witmer et al. 2003) as well as in Pteranodon longiceps (Bennett 2001), Tapejara wellnhoferi (Kellner 1996) and Tupuxuara deliradamus (Witton 2009), but the diagnostic value of this has not received much study. ...
Article
Numerous taxa make up the Early Cretaceous fauna of Brazil, including Ornithocheiroidea, Tapejaridae, Thalassodromidae, Chaoyangopteridae and a purported member of Azhdarchidae. Dsungaripteridae has only been tentatively assumed to be present in the form of ‘Santanadactylus’ spixi. New study of NMSG SAO 251093 (a specimen referred to Thalassodromeus sethi) suggests it is a previously unknown species of dsungaripterid, Banguela oberlii, tax. nov., differing from Thalassodromeus and other pterosaurs from the Early Cretaceous of Brazil by a unique combination of characters, including an upturned jaw tip, a short dorsal mandibular symphyseal shelf (dmss), and an autapomorphic thin crest placed halfway along the fused mandibular symphysis without a keel along the ventral margin of the jaw. B. oberlii, tax. nov., is referred to Dsungaripteridae based on a dmss no longer than the ventral shelf, U-shaped caudal margin of the ventral shelf and lateral margins of the mandibular symphysis concave in dorsal view. B. oberlii, tax. nov., is the youngest known dsungaripterid, and expands known morphological diversity in the clade as well as the Early Cretaceous pterosaur fauna of South America.
... The phylogenetic placement of M. altivolans within the Azhdarchoidea therefore remains uncertain and while postcranial characteristics support the erection of a new genus within either the Thalassodromidae (Witton 2009) or Chaoyangopteridae (Lü et al. 2008), no more specific a diagnosis can, or should, be reliably made at this time. ...
... Members of the Chaoyangopteridae such as Shenzhoupterus chaoyangensis and Chaoyangopterus zhangi from the Jiufotang Formation of Liaoning Province, are dated as Early Aptian, while Eopteranodon was uncovered from Barremian-Early Aptian deposits (Swisher et al. 1999). Members of the Thalassodromidae are known chiefly from the NE of Brazil, the two major fossiliferous deposits are both regarded as Early Cretaceous in age (Kellner and Campos 2002;Unwin and Martill 2007;Witton 2009). An isolated rostrum and mandible from the Javelina Formation of North America (Wellnhofer 1991;Kellner 2004) therefore appears to represent the sole member of the Thalassodromidae known from the Late Cretaceous (Martill and Naish 2006). ...
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A new pterosaur, Microtuban altivolans gen. et sp. nov., is described from the Sannine Formation of northern Lebanon. The specimen is the first pterosaur from the Early Cenomanian (Late Cretaceous) locality of Hjoûla and is regarded as the most complete pterosaur fossil discovered from Africa. While postcranial characters indicate a possible relationship with members of the Thalassodromidae or Chaoyangopteridae, the specimen possesses an exceptionally short wing-finger phalanx 4, forming only 1.1% of the total length of the wing-finger. Its appearance along with an unnamed ornithocheiroid from the slightly younger locality of Hâqel suggests that a number of pterosaur taxa existed within the local area, perhaps living on exposed carbonate platforms.
... A atividade aqui proposta tem uma seleção de representantes de pterossauros com particularidades referentes ao processo histórico de coleta e estudo, além da publicação dos materiais, etimologia de seus nomes, e apelo visual que evocam. As características e curiosidades de cada uma das cinco espécies selecionadas, a saber: (Witton, 2009) (Fig. 4E); e Tupuxuara deliradamus (Witton, 2008) (Fig. 4F). Todas elas se encontram detalhadas no Material Suplementar ("Material do Aplicador"). ...
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Introdução. A Paleontologia integra conceitos de outras áreas das Ciências da Natureza para tratar da evolução e mudanças ambientais no tempo Geológico. A despeito da sua importância, o rico registro fossilífero brasileiro é raramente tratado em sala de aula. Objetivos. Este trabalho propõe o jogo de tabuleiro “Qual é o Pterossauro?” como ferramenta adicional para o ensino de Paleontologia no Ensino Básico. Metodologia. As regras e as ilustrações são autorais e inéditas, e se basearam na literatura científica. Após a exposição ao tema em formato de uma aula, o jogo foi aplicado com alunos e posteriormente avaliado por formulário. Resultados. O jogo foi considerado como boa experiência educativa por 92 crianças de duas escolas públicas da Zona Leste da Cidade de São Paulo. Conclusão. A originalidade do jogo e de todas as ilustrações que incluem representações de pterossauros brasileiros se destacam por se tratarem de material original e autoral. O jogo se demonstrou viável para apresentar conceitos de Paleontologia e pterossauros brasileiros, como ferramenta para o ensino de Geociências no Ensino Básico e valorização do patrimônio fossilífero do nosso país.
... nov. and "Tupuxuara" deliradamus are hindered by the poor preservation of the holotype of the latter (SMNK PAL 6410), precluding the estimation of nasoantorbital fenestrae height and shape (Witton 2009). We should note that "Tupuxuara" deliradamus is a highly problematic taxon, the validity of which will be assessed in a future contribution. ...
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The Tapejaridae were an apparently worldwide distributed clade of edentulous pterosaurs, being a major component of several Lower Cretaceous terrestrial faunas. Despite their distribution across Gondwana and Laurasia, the oldest tapejarid remains were found in Barremian units from Europe, what led to the assumption that the clade originated in Eurasia and later dispersed southwards. Here we present a new tapejarid pterosaur species (Kariridraco dianae gen. et sp. nov.) from the Lower Cretaceous Romualdo Formation of Brazil. The holotype (MPSC R 1056) comprises an incomplete, three dimensionally preserved skull, lower jaw, and cervical vertebrae. It shows a unique combination of features such as unusually tall and comparatively short nasoantorbital fenestrae, as well as a premaxillary crest forming an angle of about 45° with respect to main skull axis. Phylogenetic analyses recover the new taxon as a Tupuxuara-related Thalassodrominae, a clade of early-diverging tapejarids that were apparently indigenous to central Gondwana. The inclusion of the new taxon in current phylogenetic frameworks, in addition to similarity cluster analyses of Early Cretaceous tapejarid-bearing pterosaur faunas, indicate Gondwana as the most parsimonious origin center for Tapejaridae, and show that pterosaur communities were affected by large scale tectonic-driven vicariant events.
... This latter extends parallel to the quadrate, as in Ta. wellnhoferi [33] and "Tupu." deliradamus (SMNK PAL 6410, [34]). ...
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A remarkably well-preserved, almost complete and articulated new specimen (GP/2E 9266) of Tupandactylus navigans is here described for the Early Cretaceous Crato Formation of Brazil. The new specimen comprises an almost complete skeleton, preserving both the skull and post-cranium, associated with remarkable preservation of soft tissues, which makes it the most complete tapejarid known thus far. CT-Scanning was performed to allow the assessment of bones still covered by sediment. The specimen can be assigned to Tupa. navigans due to its vertical supra-premaxillary bony process and short and rounded parietal crest. It also bears the largest dentary crest among tapejarine pterosaurs and a notarium, which is absent in other representatives of the clade. The new specimen is here regarded as an adult individual. This is the first time that postcranial remains of Tupa. navigans are described, being also an unprecedented record of an articulated tapejarid skeleton from the Araripe Basin.
... 5C). By contrast other edentulous pterosaurs either have a low mandibular crest (thalassodromeids) or none at all (pteranodontians, chaoyangopterids, azhdarchids) (Witton 2009, Vullo et al. 2012). Identification of BSP 1997 I 67 as tapejarid is supported by the recent discovery, in the Kem Kem group, of additional tapejarid material recently described by Martill et al. (2020). ...
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The geological and paleoenvironmental setting and the vertebrate taxonomy of the fossiliferous, Cenomanian-age deltaic sediments in eastern Morocco, generally referred to as the “Kem Kem beds”, are reviewed. These strata are recognized here as the Kem Kem Group, which is composed of the lower Gara Sbaa and upper Douira formations. Both formations have yielded a similar fossil vertebrate assemblage of predominantly isolated elements pertaining to cartilaginous and bony fishes, turtles, crocodyliforms, pterosaurs, and dinosaurs, as well as invertebrate, plant, and trace fossils. These fossils, now in collections around the world, are reviewed and tabulated. The Kem Kem vertebrate fauna is biased toward largebodied carnivores including at least four large-bodied non-avian theropods (an abelisaurid, Spinosaurus, Carcharodontosaurus, and Deltadromeus), several large-bodied pterosaurs, and several large crocodyliforms. No comparable modern terrestrial ecosystem exists with similar bias toward large-bodied carnivores. The Kem Kem vertebrate assemblage, currently the best documented association just prior to the onset of the Cenomanian-Turonian marine transgression, captures the taxonomic diversity of a widespread northern African fauna better than any other contemporary assemblage from elsewhere in Africa. Keywords Africa, Cretaceous, dinosaur, Gara Sbaa Formation, Douira Formation, paleoenvironment, vertebrate
... Several other fossils have been recorded in this region along the years (e.g., Kellner 1987;Maisey 1991), but flying reptiles predominate among tetrapods (e.g., Maisey 1991;Saraiva et al. 2014). Since the first specimen reported by Price (1971), over twenty-seven pterosaur species have been described up to now (e.g., Wellnhofer 1987;Frey and Martill 1994;Campos 2002, 2007;Frey et al. 2003aFrey et al. , 2003bWitton 2009;Kellner 2013;Bantim et al. 2014;Pêgas et al. 2016) some having exceptionally well preserved soft tissue (e.g., Kellner 1996). Despite the controversy of taxonomic assignment of some taxa (e.g., Fastnacht 2001;Veldmeijer 2003;Rodrigues and Kellner 2008;Martill and Unwin 2012;Pinheiro and Rodrigues 2017), two pterosaur groups, the toothed anhanguerids and the toothless tapejarids, predominate (e.g., Campos and Kellner 1985;Kellner 1989;Vila Nova and Sayão 2012). ...
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The Early Cretaceous deposits of the Araripe Basin in northeast Brazil has yielded numerous vertebrate fossils, in which pterosaurs are the predominant tetrapods. Almost all specimens of this extinct group of flying reptiles recovered from this basin come from two stratigraphic units, the Crato and Romualdo Formations, with the pterosaurs from the former being usually small to middle-sized and large individuals (with a maximized wingspan over 5 m) being only found in the latter. Here we report on a new specimen (MPSC R 1221) composed of a partial right wing, which is the largest pterosaur discovered from the Crato Formation so far, having an estimated maximized wingspan of 5.5 m. Despite the incompleteness of this material, MPSC R 1221 can be referred to the Anhangueridae based on the length ratio between the metacarpal IV and the first wing phalanx. According to the osteohistological study and the degree of fusion, MPSC R 1221 represents a sub-adult individual, showing that the animal had not reached the maximum size before its death. The present study shows that large-sized pterosaurs were also present in the Crato Formation and that their rarity might be an artefact of preservation.
... A downward curvature of the mandibular symphysis appears to be a feature common, although not unique, to certain tapejarines (P egas et al. 2016), while an anteriorly deepening posterior symphysis is also seen in the thalassodromid Tupu-xuara (e.g. Witton 2009). In this latter taxon, however, the posterior symphysis is short, deeper than wide, and abruptly narrows anteriorly, bearing sharp dorsolateral margins. ...
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We describe and interpret a posterior mandibular symphysis of a very large azhdarchid pterosaur. The specimen LPB (FGGUB) R.2347 exhibits a series of morphological characters present in both azhdarchid and tapejarid pterosaurs, suggesting a more basal position within the clade Azhdarchidae. This fossil was collected from Maastrichtian continental deposits near Vălioara in the Hațeg Basin, Romania, but cannot be confidently referred to the contemporaneous giant Hatzegopteryx thambema, also from Vălioara, due to the absence of overlapping skeletal elements. Remarkably, this mandibular symphysis shares a number of features the smaller azhdarchoid Bakonydraco galaczi from the Santonian of Hungary. Additional comparisons with previously described large‐sized azhdarchid mandibles indicate a certain degree of morphological and probably ecological disparity within the group. This specimen represents the largest pterosaur mandible ever found and provides insights into the anatomy of the enigmatic giant pterosaurs.
... In Thalassodromidae the mandibles are elongate, straight and gently tapered, usually with a ventral keel of varying degrees of prominence, which does not extend far posteriorly, fading out at the posterior termination of the mandibular symphysis, as is the case for Tupuxuara (Witton, 2009). The tip of the mandible also bears a keeled dorsal surface, which is not seen in the new Kem Kem taxon. ...
Article
A new genus and species, Xericeps curvirostris gen. et sp. nov., is erected for a highly distinctive pterosaur mandible from the mid-Cretaceous (?Albian to lower Cenomanian) Kem Kem beds of south east Morocco. The new taxon is referred to Azhdarchoidea based on the absence of teeth, slenderness of its mandible with sulcate occlusal surface, presence on the posterior section of the mandibular symphysis of short paired ridges bounding a central groove, and the presence of elongate foramina on its occlusal and lateral surfaces. A slight dorsal curvature determines it as a distinct genus of azhdarchoid, as does an autapomorphy: the presence of a continuous longitudinal groove on the ventral midline of the mandibular symphysis. The new species brings to three the number of named pterosaurs from the Kem Kem beds and together with an unnamed tapejarid, points to a relatively diverse pterosaur assemblage in these deposits.
... The Romualdo Member, the upper unit in the Santana Formation, stands out for the quality and tridimensional preservation of its fossils in calcareous concretions, belonging to different taxonomic groups, such as Gymnosperms and Angiosperms leaves and trunks (Saraiva et al. 2003, Lima et al. 2012; Gastropods (Beurlen 1964); Crustaceans (Martins Neto 1987 andPinheiro et al. 2013); Conchostracans (Carvalho and Viana 1993); Ostracods (Carmo et al. 2004) and, especially, vertebrates: Chondrichthyes Actinopterygii (Agassiz 1841, Brito and Ferreira 1989); Testudines (Hirayama 1998, Oliveira and; Dinosauria (Kellner 1999, Kellner and Campos 1996, Martill et al. 1996, Naish et al. 2004; Pterosauria (Kellner 1984, Wellnhofer 1985, Kellner and Tomida 2000, Witton 2009, Bantim et al. 2014andCrocodilia (Price 1959, Kellner 1987) and no insect has yet been attributed to this unit in the literature, which limits paleo-ecological inferences, since insects (especially Orthoptera) are excellent bioindicators of paleoenvironments (Martins Neto 2006). Orthopteran constitutes the most diverse group of Polyneoptera, containing around 22.5 thousand species (both extant and fossils) (Grimaldi andEngel 2005, Heads andMartins Neto 2007). ...
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The Romualdo Member of the Santana Formation, a lithostatigraphic unit attributed to a marine intrusion, is famous for its preserved fossils in calcareous concretions, which stand out for their diversity and excellent preservation levels. This paper aims to record the first occurrence of the Class Insecta in the Romualdo Member of the Santana Formation of the Araripe Basin, as well as to describe and discuss the paleoecological implications of such finding. The first occurrence of the order Orthoptera (family Gryllidae) is presented for this unit. This new species is attributed to the genus Araripegryllus, that lasted throughout the deposition of the Crato Member, which is under the Romualdo Member. In reference to its statigraphic origin, the specimens was named Araripegryllus romualdoi sp. nov.
... The presence and distribution of postexapophyses among pterosaurs is also complex, with authors suggesting independent origin in some taxa (Andres and Ji 2008). In fact, the evolution of the neck structure of pterosaurs is still poorly understood, with only a few studies published so far [see Witton (2009) and Averianov (2013) for further comments on neck reconstruction and biomechanics]. ...
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The cervical series is poorly known in several pterosaur clades despite the fact that distinct morphotypes of individual elements have been recognised. Among the least known is the neck of the Tapejaridae that until recently has received little attention. In order to provide further data on the tapejarid cervical anatomy, we describe three specimens from the Romualdo Formation (Lower Cretaceous, Brazil): AMNH 22568 comprises the last five cervical elements of a large individual, AMNH 24445 and MN 4728-V, both including three meso-cervical elements of small individuals. Despite size differences, AMNH 22568 and MN 4728-V are anatomically more similar to each another, showing a pattern of high, laterally flattened (blade-like) neural spines, short centra pierced by two lateral pneumatic foramina. AMNH 24445 presents a distinct anatomy, with low (but still evident) neural spines, more elongated centra with concave ventral surface, and only one lateral pneumatic foramen. Based on comparative studies we refer AMNH 22568 and MN 4728-V to the Thalassodrominae and AMNH 24445 to the Tapejarinae, helping to differentiate the cervical elements of these clades. Thalassodrominae cervical vertebrae can be distinguished by the presence of two lateral pneumatic foramina, a broad centrum (length/width ratio lower than two), and a flat ventral surface. Tapejarinae cervical vertebrae show only one lateral pneumatic foramen, the length/width ratio higher than two, and a concave ventral surface. Thalassodromine and tapejarine cervical vertebrae differ from those of azhdarchids in the presence of lateral pneumatic foramens, developed neural spine, and length/width ratio lower than three, and by being comparatively shorter.
... 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. ...
Article
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.
... They assign this fossil to a new species of small pterosaur, Thalassodromeus sebesensis (a name first coined in a conference abstract published in 2013; Grellet-Tinner et al., 2013). GTC build a taxonomic argument on the basis of this single incomplete specimen that posits the presence of a major group of pterosaurs hitherto entirely restricted to the Early Cretaceous of South Americathalassodromines (Kellner and Campos, 2007) or thalassodromids (Witton, 2009)in the European Late Cretaceous. GTC note that "this important discovery doubles the thalassadromine fossil record and demonstrates a 42 million year temporal displacement between the Romanian species and its older Aptian Gondwanan congener Thalassodromeus sethi". ...
Article
In a recent Gondwana Research article Grellet-Tinner and Codrea (in press) (hereafter “GTC”) describe a single bone (UBB ODA-28, collections of Babeş-Bolyai University, Cluj Napoca, Romania) from the Upper Cretaceous Şard Formation (=middle section of the Sebeş Formation) (Transylvanian Basin, Romania) as a pterosaur premaxillary cranial crest. They assign this fossil to a new species of small pterosaur, Thalassodromeus sebesensis (a name first coined in a conference abstract published in 2013; Grellet-Tinner et al., 2013). GTC build a taxonomic argument on the basis of this single incomplete specimen that posits the presence of a major group of pterosaurs hitherto entirely restricted to the Early Cretaceous of South America – thalassodromines (Kellner and Campos, 2007) or thalassodromids (Witton, 2009) – in the European Late Cretaceous. GTC note that “this important discovery doubles the thalassadromine fossil record and demonstrates a 42 million year temporal displacement between the Romanian species and its older Aptian Gondwanan congener Thalassodromeus sethi”. If GTC are correct, this new fossil represents a remarkably unexpected and potentially very important discovery that could rewrite aspects of pterosaur evolutionary history. We have assembled a large international team who disagree with the arguments presented by GTC. As we demonstrate, the fossil fragment they describe is misidentified; it is, firstly, not from a pterosaur but is clearly a piece of the shell of the turtle Kallokibotion (Nopcsa, 1923a,b) and, secondly, is therefore not the groundbreaking discovery of an ‘anachronistic’ Gondwanan pterosaur in Europe as claimed. Because ODA-28 is not a pterosaur, yet alone a Thalassodromeus, GTC's conclusions are also unsupported.
... Taxonomic reviews of the specimens from these European deposits led several authors to propose a reduction in the number of species present (Bennett, 1996;Unwin, 2001). Indeed, Araripe pterosaurs were first named based on very fragmentary or incomplete specimens, potentially creating a similar scenario (Price, 1971;Wellhofer, 1977;Kellner, 1984;Witton, 2009). Taking this into account, the holotype and only known specimen of C. atrox was compared with other toothed pteranodontoids. ...
Article
Based on one of the first cranial pterosaur specimens unearthed from the Romualdo Formation (Araripe Basin), Cearadactylus atrox has caused disagreement among paleontologists regarding its relationships. Ranging from an ornithocheirid, an indeterminated pterodactyloid, to a ctenochasmatid, some authors even regarded this species as representing a distinct suprageneric clade. Further preparation of the holotype that was transferred to the collections of the Museu Nacional/UFRJ (MN 7019-V) revealed several new features allowing a redescription and reevaluation of the phylogenetic position of this species. Among the new observations, it is clear that the rostral end of this specimen had been glued to the skull, rendering previous anatomical interpretations incorrect. There is no rostral gap, and the expanded rostral end of the premaxillae is larger than the dentary, rather than smaller. Cearadactylus atrox is here considered a valid taxon that can be diagnosed by a dentary groove that bifurcates at the rostral end, orbit and naris in a high position relative to the nasoantorbital fenestra, and a comparatively small number of teeth (32–36 maxillary, 22–26 mandibular), decreasing in size towards the posterior end. Phylogeneticaly, it is placed as the sister group of the Anhangueridae, forming a large clade of Brazilian forms (Tropeognathus and Anhanguera), which has a European taxon (‘Ornithocheirus’ compressirostris) as its sister group.SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP
... Brasileodactylus araripensis (Kellner, 1984) Cearadactylus atrox (Leonardi and Borgomanero, 1985) Cearadactylus? ligabuei (Dalla Vecchia, 1993) Coloborhynchus robustus (Fastnach, 2001) Coloborhynchus speilbergei (Veldmeijer, 2003) Coloborhynchus piscator (Kellner and Tomida, 2000) Pricesaurus megalodon (Martins-Neto, 1986) Santanadactylus pricei (Wellnhofer, 1985) Santanadactylus araripensis (Wellnhofer, 1985) Santanadactylus brasiliensis (Buisonjé de, 1980) Santanadactylus spixi (Wellnhofer, 1985) Tropeognathus mesembrinus (Wellnhofer, 1987) Tropeognathus robustus (Wellnhofer, 1987) Tapejara wellnhoferi (Kellner, 1989) Tupuxuara longicristatus (Kellner and Campos, 1988) Tupuxuara leonardi (Kellner and Campos, 1994) Tupuxuara deliridamus (Witton, 2009) Thalassodromeus sethi (Kellner and Campos, 2002) ...
Article
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.
... Up to date, 27 species from these deposits were formally introduced (Campos and Kellner 1985, Witton 2009, Eck et al. 2011), but the validity and the taxonomic assignment of some taxa have been subject of controversy (e.g., Fastnacht 2001, Veldmeijer 2003, Rodrigues and Kellner 2008, Martill and Unwin 2012. ...
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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.
... tween the occlusal and opposite side (~8°), by fewer and less regular vascular foramina, and by wineglass cross-section. All more or less complete rostrum fragments of Azhdarcho are short, suggesting that the rostrum and mandibular symphysis were distinctly shorter than those of Quetzalcoatlus and more approximating condition of Tupuxuara, Zhejiangopterus, or Bakonydraco (Kellner and Langston 1996; Ősi et al. 2005; Witton 2009). The dentary symphysis of Azhdarcho is generally similar to that of Bakonydraco (Ősi et al. 2005: fig. ...
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The osteology of the azhdarchid pterosaur Azhdarcho lancicollis Nessov, 1984 from the Late Cretaceous (Turonian) of Uzbekistan is described in detail based on more than 200 bone fragments representing several skull bones, cervical and dorsal vertebrae, pectoral girdle, and limb bones. Azhdarcho lancicollis is characterized by relatively short dentary symphysis and hyperelongated middle cervical vertebrae. The relative length of the cervicals is expressed by the formula I+II < III < IV < V > VI > VII > VIII > IX. The osteology in all azhdarchids is remarkable uniform but Azhdarcho can be distinguished from all other known azhdarchid genera. The phylogenetic analysis showed that the Turonian Azhdarcho and the Santonian Bakonydraco occupy a phylogenetic position basal to the Campanian Zhejiangopterus and the Maastrichtian Quetzalcoatlus.
... To date, recovered specimens include the ornithocheiroids: Araripesaurus castilhoi (Price 1971), Araripedactylus dehmi (Wellnhofer 1977), Anhanguera santanae (Wellnhofer 1985, 1991a), A. blittersdorffi (Campos and Kellner 1985), Brasileodactylus araripensis (Kellner 1984), Cearadactylus artox (Leonardi and Borgomanero 1985), C.? ligabuei (Dalla Vecchia 1993), Coloborhynchus robustus (Fastnacht 2001), C. spielbergi (Veldmeijer 2003), C. piscator (Kellner and Tomida 2000), Santanadactylus araripensis (Wellnhofer 1985; Kellner and Tomida 2000), Santanadactylus brasiliensis (De Buisonjé 1980), S. cf. pricei (Wellnhofer 1985), Ornithocheirus mesembrinus (Wellnhofer 1987), and the azhdarchoids: Tupuxuara longicristatus (Kellner 1989), T. leonardi (Kellner and Campos 1994b), T. deliradamus (Witton 2009), Thalassodromeus sethi (Kellner and Campos 2002), Tapejara wellnhoferi (Kellner 1989; Wellnhofer and Kellner 1991). The latter of these taxa, T. wellnhoferi, remains to date as one of the best represented of the Brazilian azhdarchoids, known from a collection of excellently preserved cranial material (AMNH 24440, UOSG 12891 Wellnhofer and Kellner 1991; MN 6595-V, Kellner 1989; MCT 1500-R Kellner 1996a, b). ...
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The postcranial elements of two similar sized and juvenile individuals, along with a partial skull, are attributed to the Early Cretaceous pterosaur Tapejara wellnhoferi. The remains, recovered from a single concretion of the Romualdo Member, Santana Formation, NE-Brazil, represent the first account of multiple specimens having settled together and allow for a complete review of postcranial osteology in tapejarid pterosaurs. A comparison of long bone morphometrics indicates that all specimens attributed to the Tapejaridae for which these elements are known (i.e. Huaxiapterus, Sinopterus, Tapejara) display similar bivariate ratios, suggesting that Chinese and Brazilian taxa must have exhibited similar growth patterns. An unusual pneumatic configuration, whereby the humerus is pierced by both dorsally and ventrally located foramina, is observed within T. wellnhoferi, while the pneumatic system is inferred to have invaded the hindlimbs via the femur in all members of the Azhdarchoidea. The partial preservation of the endocranial cavity allows for a reconstruction of the tapejarid brain, where despite a small orbit with respect to skull size, the presence of large flocculi and ocular lobes indicate that Tapejara possessed both excellent balancing and visual systems as a consequence of its aerial lifestyle. KeywordsBrazil–Lower Cretaceous–Santana Formation–Pterosauria–Tapejaridae–Osteology
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Pterosaurs, flying reptiles that lived during the Mesozoic era, were the first vertebrates to develop powered flight. Despite being patchy, the pterosaur fossil record extends worldwide, being quite diverse at the Araripe Basin, Brazil. The Romualdo Formation (Aptian-Albian), a Konservat Lagerstätten rich in calcareous concretions, has yielded several of these exceptionally well-preserved, three-dimensional fossils. MN 4727-V, the focus of the present study, consists exclusively of three-dimensionally preserved post-cranial axial and appendicular elements. The anatomy suggests that MN 4727-V is a pteranodontoid since it exhibits an enlarged, warped deltopectoral crest and a subtriangular distal articulation of the humerus. The phyloge-netic analysis recovered MN 4727-V as an anhanguerid by the combination of the following characters: scapula substantially shorter than the coracoid (sca/co < 0.80), and posterior expansion of the sternal articulation of the coracoid. The presence of elliptical-shaped, dorsomedially inclined depressions on the dorsal surface of the prezygapophyses of the preserved cervicals has never been reported for any other anhanguerid, although they might not be diagnostic to MN 4727-V. The tail bears unique middle caudals that exhibit ventral projections, a character previously reported only for Anhanguera piscator. The wingspan of MN 4727-V, a juvenile individual, was estimated to be about 3.9 metres.
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Tupuxuara deliradamus' was originally described as a tapejarid pterosaur from the Lower Cretaceous Romualdo Formation (Albian) of the Araripe Basin, Northeastern Brazil, and was regarded later by other authors as an indeterminate azhdarchoid. Whereas its taxonomic status remains controversial, its phyloge-netic affinities were never investigated under computational analyses. Moreover, after its original description , other putative coeval pterosaurs have been described from Brazil. In the present study, these data are combined and we reconstruct the phylogenetic affinities of 'Tu. deliradamus' for the first time. It is recovered within Tapejarinae, nesting as the sister taxon to Caupedactylus ybaka. This result challenges the former assignation to Thalassodrominae. The holotype of 'Tu. deliradamus' shares a suit of traits of tapejarines, such as the lacrimal process of jugal being posteriorly inclined, the posterodorsal margin of the nasoantorbital fenestra forming an abrupt bend level with the dorsal margin of the orbit, and inclination of the quadrate with relation to the jugal bar is approximately 150°. Unfortunately, the fragmentary condition of SMKN PAL 6410 hampers a reliable taxonomic assignment at species level. Nevertheless, the new data supports its assignation to Tapejarinae indet.
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Tapejarids are edentulous pterosaurs particularly abundant in the Chinese Jiufotang Formation, counting with over 10 described specimens and dozens of undescribed ones. A total of seven nominal tapejarid species (within two genera) have been proposed, though it is disputed how many of those are valid instead of sexual or ontogenetic morphs of fewer, or a single, species. However, detailed revisions of the matter are still lacking. In the present work, we provide a specimen-level survey of anatomical variation in previously described Jiufotang tapejarid specimens, as well as of six new ones. We present qualitative and morphometric comparisons, aiming to provide a basis for a taxonomic reappraisal of the complex. Our results lead us to interpret two Jiufotang tapejarid species as valid: Sinopterus dongi and Huaxiadraco corollatus (gen. et comb. nov.). Our primary taxonomic decisions did not rely around cranial crest features, which have typically been regarded as diagnostic for most of these proposed species albeit ever-growing evidence that these structures are highly variable in pterosaurs, due to ontogeny and sexual dimorphism. However, a reassessment of premaxillary crest variation in the Sinopterus complex reveals that while much of the observed variation (crest presence and size) can easily be attributed to intraspecific (ontogenetic and sexual) variation, some of it (crest shape) does seem to represent interspecific variation indeed. A phylogenetic analysis including the species regarded as valid was also performed.
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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.
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A historic specimen described for the first time revealed important autapomorphic characters, permitting the definition of a new species, Javelinadactylus sagebieli gen. n. et sp. n., which represents the second toothless species from the Javelina Formation, Big Bend National Park of West Texas (United States of America). The remains of J. sagebieli (Azhdarchoidea: Tapejaridae) were found in 1986, but were never properly studied, and its taxonomic affinity remains undefined. The description is based on a partially articulated skull and mandible, which offer information on the anatomy of a single azhdarchoid pterosaur. J. sagebieli exhibit a large nasoantorbital fenestra, a rostral index of medium value and is assigned to the clade Thalassodrominae, a group of tapejarid pterosaurs that were reported exclusively from the Romualdo Formation of Brazil, with only two genera known. Thalassodromines are characterized by a typical cranial configuration with toothless jaws and a high and wide premaxilla bar, formed by sub-parallel or parallel borders. The new specimen described here represents the first record of the Tapejaridae group in the Maastrichtian of North America, and the cranial morphology of the new taxon increase the richest of the diversity of the azhdarchoid pterosaurs during the end of the Late Cretaceous, suggesting that the tapejarids were still diversifying in the Maastrichtian.
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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).
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A three-dimensional and almost complete pterosaur mandible from the Crato Formation (Early Cretaceous of Northeastern Brazil), Araripe Basin, is described as a new species of a tapejarine tapejarid. Tapejarines are a particular group of toothless pterosaurs, characterized by well-developed cranial crests, downturned rostra, and have been proposed to represent frugivorous flying reptiles. Though comparatively well represented and distributed, the evolutionary history of the group is still poorly known, and the internal relationships of its members are not well understood. The new species here reported, named Aymberedactylus cearensis gen. et sp. nov., adds new data concerning the evolution of the group, concerning their morphology and geographical origin. It differs from known tapejarids due to its unusually elongate retroarticular process and a shallow fossa on the splenial exhibiting distinctive rugose texture. Furthermore, it exhibits a suite of basal and derived conditions within the Tapejaridae, demonstrating how their morphological traits probably evolved and that these forms were even more diverse than already acknowledged. The discovery of Aymberedactylus cearensis sheds new light on the evolutionary history of the Tapejarinae.
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Tapejarids are edentate pterosaurs recovered mainly from Early Cretaceous deposits. They are diagnosed by five synapomorphies, among which only one is postcranial: a broad and well-developed tubercle at the ventroposterior margin of the coracoid. Regarding the clade Thalassodrominae, most phylogenetic studies are based on cranial elements, as postcranial skeletons of these pterosaurs are rare. Here, new postcranial material from the Romualdo Formation (Aptian–Albian) from the Araripe Basin is described. The material comprises the three posteriormost cervical vertebrae, the first seven dorsal vertebrae (fused into a notarium), both scapulocoracoids, a fragment of a sternum, a partial right humerus, a small fragment of a 4th phalanx of the wing finger, a distal extremity of the right femur and the proximal portions of both tibia and fibula. Comparisons with other specimens and morphological features examined in a phylogenetic context, such as the presence of three foramina lateral and dorsal to the neural canal of the cervical vertebrae, the presence of a notarium and a pneumatic foramen on the ventral side of the proximal portion of the humerus, allow the assignment of this specimen as Thalassodrominae indet. Regarding palaeobiogeographical aspects, to date, this clade is exclusively found in the Romualdo Formation. It is the most complete postcranial material assigned to the Thalassodrominae described so far.
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A new unusual tapejarid pterosaur from the Early Cretaceous Romualdo Formation (Araripe Basin, Brazil) is described, based on a skull, lower jaw and some postcranial elements. Caupedactylus ybaka gen. et sp. shows the typical high nasoantorbital fenestra of the Thalassodrominae but lacks a palatal ridge, and shares with the Tapejarinae several features, including a downturned rostral end, allowing its allocation to that clade. Furthermore, the new species differs in having an anteriorly and posteriorly expanded premaxillary sagittal crest, the lacrimal process of the jugal strongly inclined, and a slit-like postpalatine fenestra, among other characters. The region of the left jugal-quadratojugal-quadrate shows a pathology that is likely the result of an infection. The lateral surface of the premaxillary crest presents grooves that were interpreted in other pterosaurs as impressions of blood vessels, corroborating growing evidence that cranial crests could have been involved in thermoregulation. Also, the new species has a well-preserved palate with a large palatine forming the anterior region of the choanae and the postpalatine fenestra and a secondary subtemporal fenestra. Since the latter has been regarded as unique to non-pterodactyloids, its occurrence in Caupedactylus demonstrates that the evolution of palatal region in pterosaurs is more complex than previously thought.
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A partial skull of the pterodactyloid pterosaur Tupuxuara from the Santana Formation (Cretaceous, ?Albian) of Brazil displays catastrophic damage to the anterior rostrum and mandible. The damage is not a consequence of diagenetic fracturing and is thought to represent a violent experience inflicted on the skull prior to deposition. The skull fracture patterns are consistent with a single, high-energy impact occurring at the jaw tips with a caudodorsal force vector. This impact resulted in telescoping of the rostrum into the nasoantorbital fenestra, buckling of the mandible and subsequent destruction of basal cranial elements from the caudal transmission of impact forces. However, the lack of clear event signatures on the specimen prevents identification of the specific cause of this damage.
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The largest known flying organisms are the azhdarchid pterosaurs, a pterodactyloid clade previously diagnosed by the characters of their extremely elongate middle-series cervical vertebrae. The named species of the Azhdarchidae are from the Late Cretaceous. However, isolated mid-cervical vertebrae with similar dimensions and characters have been referred to this group that date back to the Late Jurassic, implying an almost 60 million year gap in the fossil record of this group and an unrecorded radiation in the Jurassic of all the major clades of the Pterodactyloidea. A new pterosaur from the Early Cretaceous of Liaoning Province of China, Elanodactylus prolatus gen. et sp. nov., is described with mid-cervical vertebrae that bear these azhdarchid characters but has other postcranial material that are distinct from the members of this group. Phylogenetic analysis of the new species and the Pterodactyloidea places it with the Late Jurassic vertebrae in the Late Jurassic–Early Cretaceous Ctenochasmatidae and reveals that the characters of the elongate azhdarchid vertebrae appeared independently in both groups. These results are realized though the large taxon sampling in the analysis demonstrating that the homoplastic character states present in these two taxa were acquired in a different order in their respective lineages. Some of these homoplastic characters were previously thought to appear once in the history of pterosaurs and may be correlated to the extension of the neck regions in both groups. Because the homoplastic character states in the Azhdarchidae and Ctenochasmatidae are limited to the mid-cervical vertebrae, these states are termed convergent based on a definition of the term in a phylogenetic context. A number of novel results from the analysis presented produce a reorganization in the different species and taxa of the Pterodactyloidea.
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We report on two juvenile specimens of Pterodaustro guinazui, a medium-sized, filter-feeding pterodactyloid pterosaur from the Lower Cretaceous (Albian) lacustrine deposits of the Lagarcito Formation of central Argentina (Sierra de Las Quijadas, San Luis Province). Both specimens lack the skull, but one is otherwise nearly complete and articulated, and the other is more fragmentary. Their small size (300 mm of wingspan), the lack of fusion of several postcranial bones, the minimal epiphyseal ossification, and the porous appearance of the periosteal surfaces indicate the specimens died at a very early stage of postnatal development. The presence of derived characters unique, within Pterodactyloidea, to P. guinazui (e.g., caudal vertebrae exceeding 16 elements) supports the proposed specific identification of the specimens. The new material provides anatomical information previously unknown for P. guinazui and sheds light on allometric transformations during the growth of this pterosaur. Comparisons with more mature and much larger individuals support two major allometric trends: (i) the negative allometric growth of the proximal portion of the forelimb, and (ii) the positive allometric growth of the metacarpal IV.
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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.
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Author Summary Just because a component of an extinct animal resembles that of a living one does not necessarily imply that both were used for the same task. The lifestyles of pterosaurs, long-extinct flying reptiles that soared ancient skies above the dinosaurs, have long been the subject of debate among palaeontologists. Similarities between the skulls of living birds (black skimmers) that feed by skimming the water surface with their lower bill to catch small fish, and those of some pterosaurs have been used to argue that these ancient reptiles also fed in this way. We have addressed this question by measuring the drag experienced by model bird bills and pterosaur jaws and estimating how the energetic cost of feeding in this way would affect their ability to fly. Interestingly, we found that the costs of flight while feeding are considerably higher for black skimmers than previously thought, and that feeding in this way would be excessively costly for the majority of pterosaurs. We also examined pterosaur skulls for specialised skimming adaptations like those seen in modern skimmers, but found that pterosaurs have few suitable adaptations for this lifestyle. Our results counter the idea that some pterosaurs commonly used skimming as a foraging method and illustrate the pitfalls involved in extrapolating from living to extinct forms using only their morphology.
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Life-history parameters of pterosaurs such as growth and ontogenetic development represent an enigma. This aspect of pterosaur biology has remained perplexing because few pterosaur taxa are represented by complete ontogenetic series. Of these, Pterodaustro is unique in that besides being represented by hundreds of individuals with wing spans ranging from 0.3 to 2.5m, it includes an embryo within an egg. Here we present a comprehensive osteohistological assessment of multiple skeletal elements of a range of ontogenetic sizes of Pterodaustro, and we provide unparalleled insight into its growth dynamics. We show that, upon hatching, Pterodaustro juveniles grew rapidly for approximately 2 years until they reached approximately 53% of their mature body size, whereupon they attained sexual maturity. Thereafter, growth continued for at least another 3-4 years at comparatively slower rates until larger adult body sizes were attained. Our analysis further provides definitive evidence that Pterodaustro had a determinate growth strategy.
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Azhdarchid pterosaurs were among the most widespread and successful of pterosaur clades, but their paleoecology remains controversial. Morphological features common to all azhdarchids include a long, shallow rostrum; elongate, cylindrical cervical vertebrae that formed a long and unusually inflexible neck; and proportionally short wings with an abbreviated fourth phalanx. While azhdarchids have been imagined as vulture-like scavengers, sediment probers, swimmers, waders, aerial predators, or stork-like generalists, most recent authors have regarded them as skim-feeders, trawling their lower jaws through water during flight and seizing aquatic prey from the water's surface. Although apparently widely accepted, the skim-feeding model lacks critical support from anatomy and functional morphology. Azhdarchids lack the many cranial specialisations exhibited by extant skim-feeding birds, most notably the laterally compressed lower jaw and shock absorbing apparatus required for this feeding style. Well-preserved azhdarchid skulls are rare, but their rostra and lower jaws appear to have been sub-triangular in cross-section, and thus dissimilar to those of skim-feeders and sediment probers. Taphonomic data indicates that azhdarchids predominately inhabited inland settings, and azhdarchid morphology indicates that they were poorly suited for all proposed lifestyles bar wading and terrestrial foraging. However, azhdarchid footprints show that their feet were relatively small, padded and slender, and thus not well suited for wading. We argue that azhdarchids were stork- or ground hornbill-like generalists, foraging in diverse environments for small animals and carrion. Proficient terrestrial abilities and a relatively inflexible neck are in agreement with this interpretation.
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The specimen consists of a complete skull, the first one reported from the Crato Member, of a new species, Tapejara imperator n.sp. It displays a remarkable sagittal crest that doubles the length and increases in about six times the height of the skull. The upper and main portion of the crest is formed by soft tissue that is supported anteriorly and posteriorly by long strips of bone. This crest, never before reported in any vertebrate, most likely was a display structure, but due to its size it must also had some aerodynamic effect during flight.
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The morphology of jaws, neck, notarium, ribs, sacrum, pectoral girdle, wing and hind limbs of the type genus of azhdarchids are described and analyzed. Diagnosis of the family and genera Tepuxuara, Bennettazhia gen. nov., Azhdarcho, Bogolubovia, Quetzalcoatlus and Arambourgiania are given. -English summary
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A phylogenetic analysis indicates that the Tapejaridae is a monophyletic group of pterodactyloid pterosaurs, diagnosed by the following synapomorphies: premaxillary sagittal crest that starts at the anterior tip of the premaxilla and extends posteriorly after the occipital region, large nasoantorbital fenestra that reaches over 45% of the length between premaxilla and squamosal, lacrimal process of the jugal thin, distinct small pearshaped orbit with lower portion narrow, and broad tubercle at the ventroposterior margin of the coracoid. Several cranial and postcranial characters indicate that the Tapejaridae are well nested within the Tapejaroidea, in sister group relationship with the Azhdarchidae. A preliminary study of the ingroup relationships within the Tapejaridae shows that Tupuxuara is more closely related to Thalassodromeus relative to Tapejara. At present tapejarid remains have been found in the following deposits: Crato and Romualdo members of the Santana Formation (Aptian-Albian), Araripe Basin, Brazil; Jiufotang Formation (Aptian), Jehol Group of western Liaoning, China; and in the redbeds (Cenomanian) of the Kem Kem region, Morocco. An incomplete skull found in the Javelina Formation (Maastrichtian), Texas also shows several tapejarid features and might be a member of this clade. Although information is still limited, the present distribution of the Tapejaridae indicates that this clade of pterosaurs was not exclusive of Gondwana, and was more widespread than previously known.
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Introduction The first significant account of a pterosaur from the Crato Formation was published over a decade ago (Frey and Martill, 1994). In the short intervening period between then and now, more than 30 individuals have come to light. This is a modest total, especially when compared to the 1000+ individuals recovered, for example, from the Solnhofen Limestones of southern Germany or the Niobrara Chalk of Kansas. Still, several finds notable for their completeness, or for the exceptional preservation of soft-tissue structures, such as extensions to cranial crests or of the integument associated with the foot, have already demonstrated the importance of this lagerstätte for our understanding of pterosaur palaeobiology (Frey and Martill, 1994; Campos and Kellner, 1997; Frey and Tischlinger, 2000; Frey et al., 2003c). The Crato pterosaur assemblage also contains a number of genera (e.g. Arthurdactylus, Ludodactylus, Ingridia gen. nov.) that are unique to this deposit (Frey and Martill, 1994; Frey et al., 2003b; this chapter). They represent several Lower Cretaceous pterosaur lineages and throw some much-needed light on the ecology, palaeobiogeography and evolutionary history of pterosaurs during an interval when they appear to have reached their highest levels of global diversity (Unwin, 2005). Fragments of several limb bones from the Upper-Triassic Caturrita Formation of southern Brazil (Bonaparte et al., 2006) may represent the earliest occurrence of pterosaurs in South America.
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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
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Immature specimens of the Late Cretaceous pterosaur Pteranodon were identified using three size-independent criteria: (1) fusion of various cranial and postcranial elements; (2) degree of epiphyseal ossification; and (3) bone grain or degree of ossification of limb-bone shafts. Immature individuals make up 15% of available specimens of Pteranodon and do not differ significantly in size from mature individuals. This and the extensive fusion of the mature skeleton suggests that Pteranodon had determinate growth. The bone of limb-bone shafts of immature individuals is fibro-lamellar bone, which suggest that they grew rapidly to adult size. The size-independent criteria can also be used to identify immature and mature individuals of other pterosaur taxa, and other large pterodactyloids also probably exhibited rapid determinate growth. -Author
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Through the kindness of Prof. E. L. Packard, of the University of Oregon, I have recently received for study a fossil specimen found by him in the marine Cretaceous rocks of eastern Oregon. This specimen clearly belongs to the Pterosauria, and, as this reptilian group has not previously been known to occur in the Pacific coast region of North America, the discovery is of much scientific interest. In North America pterosaurian remains have been found in the marine Niobrara Chalk of -western Kansas and in the fresh-water Morrison deposits of Wyoming. Three genera are recognized Pteranodon and Nyctodactylus from the Niobrara, and Dermodacty-lus from the Morrison formation. The first two mentioned genera are adequately defined from well-preserved specimens ; but the latter, founded on a single incomplete and poorly preserved skeleton, is at this time inadequately characterized. Well-preserved pterosaurian specimens are among the rarest of American reptilian fossils, and when this pterosaurian fauna is con-trasted with those of Great Britain and Europe, with their great number of genera and species of wide geological range, the paucity of our rocks in pteryodactyle remains becomes strikingly apparent. This comparison serves also to accentuate the importance of this latest discovery, in greatly extending their Imown geographical range as well as furnishing a representative of the order that is intermediate in geological position between the earliest and latest known American members. In regard to the geological occurrence of this specimen. Professor Packard, under date of September 19, 1927, writes me as follows: "These specimens were found in Cretaceous shales associated with a determinable ammonite fauna of Lower Chico, or possibly Upper Horsetown age." The specimens referred to in the above citation are the pterosaur and an ichthyosaur,^ the first and only vertebrate re-mains so far found in this formation.
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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.
Article
Numerous remains of the azhdarchid pterosaur Quetzalcoatlus sp., have been recovered over the last twenty years from the Late Cretaceous (Maastrichtian) rocks in Big Bend National Park in Trans-Pecos Texas. Among more than 200 bones found at one locality are four incomplete skulls and mandibles, which provide the most complete information about cranial structures in the Azhdarchidae. What is currently known indicates that the Azhdarchidae is the sister group of the Tapejaridae from Early Cretaceous deposits in northeastern Brazil.
Article
A cladistic analysis based on 39 terminal taxa and 74 characters (several multistate) using PAUP (Phylogenetic Analysis Using Parsimony) (3.1.1 for MacIntosh and 4.0b10 for Microsoft Windows) presents a new hypothesis of pterosaur inter-relationships. This study suggests that the most primitive taxon is the Anurognathidae, followed by Sordes and all remaining pterosaurs. Dendrorhynchoides is confirmed as a member of the Anurognathidae, being closely related to Batrachognathus. Preondactylus occupies a more derived position than Sordes , which questions its previous assignment as the most primitive pterosaur. The hypothesis of rhamphorhynchoid paraphyly is confirmed, with the Rhamphorhynchidae more closely related to the Pterodactyloidea than to more basal forms. The Pterodactyloidea shows a basal dichotomy: the Archaeopterodactyloidea and the Dsungaripteroidea. The Archaeopterodactyloidea is formed by Pterodactylus + Germanodactylus and a clade formed by Gallodactylidae + Ctenochasmatidae. The Nyctosauridae occupies the basal position within dsungaripteroids and is followed by the Pteranodontoidea and the Tapejaroidea. Pteranodontoids have Pteranodon at the base, followed stepwise by Istiodactylus, Ornithocheirus and the Anhangueridae. Tapejaroids are composed of the Dsungaripteridae at the base followed by the Tapejaridae and the Azhdarchidae. Major trends within pterosaur evolutionary history are: general increase in size (wing span and body); increase of wing metacarpal and pteroid; decrease of proportional length of the second and third wing phalanx relative to the first; gradual increase of rostrum (anterior to external nares); and anterior shift of the skull-mandible articulation. Cranial crests are present in most pterodactyloids, but markedly in the Ornithocheiroidea, where all taxa show some sort of crest on the skull. The loss of teeth, previously assumed to have occurred independently in several lineages, seems to be a general trend among dsungaripteroids. Several nodes recovered by this analysis are supported by very few characters, a result at least partially attributable to the limited available information from several taxa due to poor preservation and/or preparation.
Article
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.
Article
The vomeronasal (VN) system is a pheromone-processing sensory system of tetrapods. Tetrapods use pheromones to communicate territorial boundaries, reproductive status, sex, and species identity. Presumed impressions of VN bulbs on phytosaur frontals led to a claim that phytosaurs possessed the VN system. However, in extant crocodilians, which lack the VN system, the corresponding impressions are associated not with cerebral tissue but with the ophthalmic nerves. Phytosaur head morphology was not conducive to pheromone collection. The extant phylogenetic bracket suggests that all extinct archosaurs, including phytosaurs, lacked the VN system. Without the pheromonal sense, they would not have used chemical means to convey territorial boundaries, reproductive status, sex, and species identity. Instead, they would have used visual, acoustic, and tactile cues, as in extant archosaurs and other tetrapods in which the VN sense is reduced or absent.
Article
Metric data from the large collection of the Late Cretaceous pterosaur Pteranodon are bimodally distributed with a more numerous small size-class and a less numerous large size-class. The size-classes differ in cranial crest and pelvic structure. The small size-class has small cranial crests and deep puboischiadic plates that produce a large pelvic canal, while the large size-class has large cranial crests and shallow puboischiadic plates that produce a small pelvic canal. The difference in pelvic structure suggests that the small size-class is female. The various functions proposed for cranial crests are reviewed, and it is concluded that none can account for the difference in crest size. The conclusion is that the large crests of males are display structures.
Article
Two specimens of a tapejarid pterosaur (Pterodactyloidea. Tapejaridae) are described as representing a new species. Both specimens show evidence for soft tissues preserved in association with a sagittal bony cranial crest. Both specimens are from the Nova Olinda Member Konservat Lagerstätte of the Crato Formation of the Araripe Basin, northeastern Brazil. They represent the second tapejarid species from this formation. Comparisons are made with other crested pterosaurs and comments on the utility and aerodynamics of pterosaurian head crests are made.
Article
Abstract  A new species of tapejarid pterosaur, Huaxiapterus corollatus sp. nov. is erected on the basis of a nearly complete skull and postcranial skeleton from the Lower Cretaceous Jiufotang Formation of Liaoning Province, China. Huaxiapterus corollatus sp. nov. is characterized by a hatchet-shaped rectangular process on the premaxilla, whose short axis is perpendicular to the anterior margin of the premaxillae. Except for this process, other characters of the skull such as the breadth of the snout between the anterior margin of the nasoantorbital fenestra and the anterior margin of the premaxilla are similar to that of Huaxiapterus jii. Huaxiapterus and a second Chinese tapejarid, Sinopterus, share several unique cranial characters in common with Tapejara and these three genera appear to be more closely related to each other than to other azhdarchoids. The Chinese tapejarids (Sinopterus and Huaxiapterus) have relatively elongate skulls and weakly developed cranial crests and seem to be less derived than Tapejara, with its shorter, deeper skull and large cranial crest. Tupuxuarids (Tupuxuara and Thalassodromeus) have often been associated with tapejarids in the family Tapejaridae, but this relationship is controversial because some phylogenetic analyses have supported the pairing of tupuxuarids with Azhdarchidae. We propose that Tapejaridae be restricted to Tapejara, Sinopterus and Huaxiapterus.
Article
Two recently collected specimens of the Late Cretaceous pterosaurNyctosaurus differ from all previously known specimens in the possession of a large branching cranial crest. The crest extends upward and backward from the posterior skull roof and is nearly three times the length of the skull proper. Despite the large crest, the specimens do not differ significantly in morphology from previously known specimens ofNyctosaurus, and do not seem to represent a new species ofNyctosaurus. The specimens suggest that the cranial crest was developed late in ontogeny, which is consistent with the interpretation of pterosaur cranial crests as intraspecific display structures. Zwei neue Exemplare des FlugsauriersNyctosaurus unterscheiden sich von anderen Exemplaren, weil sie einen hohen, abzweigenden Schädelkamm haben. Der Schädelkamm dehnt sich aufwärts und rückwärts von dem hinteren Schädeldach und ist dreimal so lang wie der Schädel. Trotz des Schädelkamms unterscheiden sich die neuen Exemplare in der wichtigen Morphologie nicht von anderen Exemplaren. Demnach sind sie keine neue Art. Wahrscheinlich entwickelte sich der Schädelkamm nur, als das Individuum ein fast ausgewachsenes Tier war. Das stimmt mit der Vermutung überein, dass der Schädelkamm ein intraspezifisches Zeigmerkmal ist.
Article
Two juvenile specimens of the pterodactyloid pterosaur Germanodactylus cristatus are identified from the Solnhofen Limestone of southern Germany. The specimens had been referred to the nominal species Pterodactylus kochi and P. micronyx. They have edentulous tips on the upper and lower jaws; their skulls are taller and less elongate and have fewer teeth than similarly sized specimens assigned to P. antiquus and P. micronyx; and they have tall neural spines on their cervical vertebrae. In addition, they exhibit a distinctive suite of skeletal proportions that distinguishes them from similarly sized specimens assigned to P. antiquus and P. micronyx. The specimens do not exhibit any trace of a cranial crest, which supports the interpretation that the cranial crest developed late in ontogeny, probably as the individual approached skeletal and/or sexual maturity. The taxonomy of Germanodactylus is reviewed and revised diagnoses are presented.
Article
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.
Article
The increasing numbers of pterosaur finds from Brazil obviously increased our knowledge, but through the years, taxonomy has become less clear as one would expect. Various reasons can be given for this. One is that important morphological features have not been incorporated in the discussion whereas others are, but have questionable taxonomic value. Also, sometimes unstated assumptions play an important role on which pterosaurs have been re-classified. It is understandable that one seeks a connection with the pterosaurs from the Cambridge Greensands in England, but this has had a negative impact on the systematics: the fossils from the Cambridge Greensands themselves have been subject to fierce debates and much confusion ever since their first discovery in the latter half of the 19th century. The present work presents a reappraisal of four closely related taxa with teeth (Anhanguera, Brasileodactylus, Coloborhynchus en Criorhynchus) by means of the study of unpublished and published material in the major collections all around the world, concentrating on cranial parts. In chapter 2, a largely complete skeleton of Coloborhynchus is described. The specimen has been classified as new species. Chapter 3 presents various post-cranial elements, on the basis of which it is concluded that interspecific differences in the post-cranial skeleton are so small that an evaluation is necessary. More important for the present work, however, is the description of the mandible of Criorhynchus, which confirms the strong deviating dentition pattern relative to the other taxa. Chapter 4 presents one of the most complete mandibles of Anhanguera. The study of this specimen resulted among others in the discovery of an important diagnostic morphological feature of the mandible. The next chapter (chapter 5) is a preliminary description of a largely unprepared skull and wing of a fossil in the collection of the AMNH, New York. The specimen is tentatively classified as Brasileodactylus mainly because of the crestless jaws (currently, the fossil is completely prepared and under study). In chapter 6, an almost complete mandible of Brasileodactylus is described. Furthermore, post-cranial material as well as a small piece of maxilla, belonging to the same individual of Brasileodactylus is presented. This is the first description of post-cranial material of this genus; as expected, there is a high degree of conformity with the other taxa. Chapter 7 summarises and shortly discusses all taxa, presents the main conclusions again, and furthermore, presents the measurements of the dentition, showing specific patterns for each taxon. The final remarks in chapter 8 offer a discussion on the problems with the Cambridge Greensands (Ornithocheiridae versus Anhangueridae). This study resulted in the refinement of various diagnoses and makes the distinction between the four taxa clearer: animals with a premaxillary and dentary sagittal crest (Coloborhynchus, Criorhynchus and Anhanguera) belong to Anhangueridae; the crestless Brasileodactylus however, cannot be classified to Anhangueridae, despite the close relationship due to the incompleteness of its remains. The diagnosis of Ornithocheiridae, as recently proposed in a re-evaluation, is not based on the type specimen and the used specimen does not show the described characters.
Article
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'.
Article
The Harderian gland is an orbital feature found in most terrestrial vertebrates. Although there have been several reports on the structure of the squamate Harderian gland, there has been little recent discussion as to its potential function. This article reviews both the recent morphological observations and their implications on the potential functions of the squamate Harderian gland. Literature on the gross structure, histochemistry, and ultrastructure of the squamate Harderian gland and associated structures was reviewed. These observations were then used to assess morphologically the likelihood of the proposed functions. A high level of morphological variation was found in the squamate Harderian gland. Three functional hypotheses, including roles in orbital lubrication, digestion, and vomerolfaction, were considered. Both morphology of the squamate Harderian gland and the presence of alternate secretory sources suggest that it is unlikely to function in orbital lubrication. There is little evidence to suggest a function in digestion. Both the presence of the connecting lacrimal apparatus and the reduced intrinsic secretory capacity of the vomeronasal organ suggest that the Harderian gland may function in vomerolfaction. The most likely role of the squamate Harderian gland seems to be in vomerolfaction. Morphological variations observed in the Harderian gland may mirror the different degrees and mechanisms of vomerolfaction. Further studies, including comparative morphological, experimental, and microchemical analyses, are required to test this hypothesis.
Article
Toothless pterosaurs played a key role in broadening the taxonomic, morphological and ecological diversity of Cretaceous pterosaurs. Here we report a complete, articulated skeleton of a 1.4-m-wingspan pterosaur from the Lower Cretaceous Jiufotang Formation of Liaoning Province, China which is identified as a new genus and species, Shenzhoupterus chaoyangensis gen. et sp. nov. The new taxon is edentulous, with a relatively large skull and a remarkably large, tall nasoantorbital fenestra that extends well above the main part of the braincase. Phylogenetic analysis shows that Shenzhoupterus gen. nov. belongs in a distinct clade of azhdarchoid pterosaurs, formally recognised here as a new family, Chaoyangopteridae, that also includes Chaoyangopterus, Jidapterus and Eoazhdarcho from the Jiufotang Formation and Eopteranodon from the Yixian Formation. These new data clarify recent confusion surrounding the systematics of these Lower Cretaceous taxa and provide new insights into the evolutionary history of pterosaurs.
Window into an Ancient World: The Crato Fossil Beds of Brazil
  • D M Unwin
  • D M Martill
Unwin, D.M., Martill, D.M., 2007. Pterosaurs from the Crato Formation. In: Martill, D.M., Bechly, G., Loveridge, R.F. (Eds.), Window into an Ancient World: The Crato Fossil Beds of Brazil. Cambridge University Press, Cambridge, pp. 475-491.
Gigantskiye lyetayushchiye yashchyeryi semyeistva Azhdarchidae. I. Morfologiya, sistematika. Vestnik Leningradskogo Gosudarstvennogo Universiteta
  • L A Nesov
Nesov, L.A., 1991. Gigantskiye lyetayushchiye yashchyeryi semyeistva Azhdarchidae. I. Morfologiya, sistematika. Vestnik Leningradskogo Gosudarstvennogo Universiteta, Seriya 7 (2), 14-23 (in Russian).
Toothed pterosaurs of the Santana Formation (Cretaceous
  • A J Veldmeijer
Veldmeijer, A.J., 2006. Toothed pterosaurs of the Santana Formation (Cretaceous;
The palaeoecology and diversity of pterosaurs
  • M P Witton
Witton, M.P., 2008. The palaeoecology and diversity of pterosaurs. Unpublished Ph.D. Thesis, University of Portsmouth, 307 pp.