Article

A new crested ornithocheirid from the Lower Cretaceous of northeastern Brazil and the unusual death of an unusual pterosaur

Authors:
To read the full-text of this research, you can request a copy directly from the authors.

Abstract

An exceptionally well-preserved cranium and mandible of a new species of pterodactyloid pterosaur from the Nova Olinda Member of the Crato Formation (Aptian, Early Cretaceious) of the Araripe Basin, northeastern Brazil, is described. The new taxon is characterized by the presence of a caudally directed parietal crest similar to that seen in pteranodontids, but is referred to the Ornithocheiridae of the Ornithocheiroidea. The specimen is referred to a new genus within the Ornithocheiridae, as it lacks the diagnostic rostral crest and instead possesses this parietal crest oriented. A lanceolate leaf with frayed distal end wedged between the mandibular rami suggests the cause of death for the specimen.

No full-text available

Request Full-text Paper PDF

To read the full-text of this research,
you can request a copy directly from the authors.

... Between taxa there appear to be minor differences in tooth size and the degree of ornamentation. For example, the anterior-most teeth of Linlongopterus are smooth whereas the equivalent teeth in Ludodactylus Frey et al., 2003 are strongly ornamented with crenulations (Fig. 3). ...
... Conversely, teeth in more posterior parts of the jaws are straighter, or even straight and have oval cross-sections, and bear carinae on the anterior and posterior margins of the crown. The teeth become progressively smaller and wider spaced posteriorly along both jaws (Frey et al., 2003). The teeth from Malyy Prolom display all of these morphologies, and thus can satisfactorily be referred to Ornithocheiridae sensu Unwin (2003). ...
... Dentition in ornithocheirid pterosaurs. A, upper jaw dentition of Ludodactylus sibbicki SMNK PAL 3828Frey et al. (2003). Notice the size heterodonty between the front and the rear, as well as subtle shape changes, especially in the degree of curvature. ...
Article
Remains of Late Cretaceous toothed pterosaurs such as ornithocheirids and lonchodectids are almost unknown for post Cenomanian strata. Here we describe teeth referrable to ornithocheirid pterosaurs from the Upper Cretaceous (Santonian) Dmitrov Formation of Ryazan region, European Russia. The teeth are somewhat worn, but retain sufficient features to satisfactorily refer them to Ornithocheiroidea, including their slender curved shape, presence of faint reticulated ribbing and a curved margin of the enamel - dentine contact. Although occurring in a Santonian deposit, the teeth are most likely derived from the underlying Cenomanian Yakhroma Formation. Although ornithocheirid teeth have been reported previously from European Russia, these teeth are the first from this region and the first evidence for Pterosauria in the Dmitrov Formation, which is better known for its elasmobranch and marine reptile assemblage.
... The craniomandibular joint levels with the anterior margin of the orbit, similarly to both specimens of N. ignaciobritoi (see Fig. 3; both specimens, LPM 00023 and IVPP V-13288; see Wang et al., 2005Wang et al., , 2006Lü, Xu & Ji, 2008), Anhanguera spp. (see Kellner & Tomida, 2000) and Linlongopterus jennyae (see Rodrigues et al., 2015), but unlike Istiodactylus spp., in which the joint is located anterior to the orbit (see Andres & Ji, 2006;Witton, 2012), and Ikrandraco avatar (see Wang et al., 2015), Hamipterus tianshanensis (see Wang et al., 2014) and Ludodactylus sibbicki (Frey, Martill & Buchy, 2003), in which the joint is located under the middle of the orbit. The orbit is piriform, with the narrowest part being ventral, and without a suborbital vacuity. ...
... The nasal and lacrimal form the anterodorsal margin of the orbit and the posterodorsal margin of the nasoantorbital fenestra. The anterior end of the nasolacrimal coincides with the highest point of the nasoantorbital fenestra, as in both specimens of N. ignaciobritoi and also Ikrandraco avatar (Wang et al., 2005(Wang et al., , 2006Andres & Ji, 2006;Lü, Xu & Ji, 2008), but unlike Istiodactylus latidens and most anhanguerians (e.g., Anhanguera, Tropeognathus, and Hamipterus), except for Ludodactylus sibbicki, in which the highest point is posterior to the anterior end of the nasolacrimal (Campos & Kellner, 1985;Wellnhofer, 1987;Kellner & Tomida, 2000;Wang et al., 2014;Frey, Martill & Buchy, 2003). A nasal descending process cannot be seen in BPMC-0204, possibly because it is still covered by rock. ...
... Of the pterygoid, only the medial process can be seen. It is large and plate-like as that of Hongshanopterus lacustris (see Wang et al., 2008a) and, to a lesser extent, the anhanguerids, in which the process is also broad but less medially expanded (Campos & Kellner, 1985;Frey, Martill & Buchy, 2003). This differs from the slender medial processes of the pterygoid of azhdarchoids (Pinheiro & Schultz, 2012;Kellner, 2013;Pêgas, Costa & Kellner, 2018) or those of the referred specimen of N. ignaciobritoi (see Wang et al., 2006;Lü, Xu & Ji, 2008) and Ikrandraco avatar (see Wang et al., 2015). ...
Article
Full-text available
A new istiodactylid pterosaur, Nurhachius luei sp. nov., is here reported based on a complete skull with mandible and some cervical vertebrae from the lower part of the Jiufotang Formation of western Liaoning (China). This is the second species of Nurhachius , the type-species being N. ignaciobritoi from the upper part of the Jiufotang Formation. A revised diagnosis of the genus Nurhachius is provided, being this taxon characterized by the presence of a slight dorsal deflection of the palatal anterior tip, which is homoplastic with the Anhangueria and Cimoliopterus. N. luei sp. nov. shows an unusual pattern of tooth replacement, with respect to other pterodactyloid species. The relationships within the Istiodactylidae and with their closest taxa are investigated through a phylogenetic analysis by parsimony.
... As a rule, individual anhanguerid species were described mostly based in subtle morphological discrepancies in their premaxillary crests and overall skull proportions (see Pinheiro and Rodrigues, 2017). However, albeit still not being a consensus, it has been suggested by many authors (Bennett, 1992(Bennett, , 1994Carpenter et al., 2003;Frey et al., 2003;Unwin, 2005;Lu et al., 2011;Wang et al., 2014;Pinheiro and Rodrigues, 2017) that the premaxillary crest in pterosaurs is a sexually selected, dimorphic feature, being subject to ontogenetic variation and not suitable as a diagnostic feature. ...
... It has already been discussed (e.g., Bennett, 1992Bennett, , 1994Bennett, , 2006Carpenter et al., 2003;Frey et al., 2003;Unwin, 2005) that pterosaur premaxillary crests are potentially sexually dimorphic and subject to ontogenetic variation, being of limited use as diagnostic features. In 2014, two unexpected discoveries added to this discussion, shedding light over interesting aspects of pterosaur biology. ...
Article
Full-text available
The Anhangueridae are a clade of toothed pterodactyloid pterosaurs, known from their characteristicanteriorly expanded premaxillae and conspicuous rostral sagittal premaxillary and dentary crests. Most knownanhanguerids come from the Lower Cretaceous Romualdo Formation within the Araripe Basin of northeast Brazil. Withfour currently valid genera and several specimens referred to the clade, anhanguerids are the most abundant and diversetetrapod group in the Romualdo Formation. However, some studies suggest this diversity may be overestimated, as manytaxa have been diagnosed based on subtle differences in their premaxillary crests, a structure argued to be eitherontogenetically variable or sexually dimorphic. Here we describe an anterior fragment of a gracile pterosaur rostrum thatpossesses the single diagnostic feature of Anhanguera (fifth and sixth pairs of dental alveoli smaller than the fourth andseventh), but lacks a sagittal crest. The affinities of the new fossil among other toothed pterosaurs were tested throughboth cladistic and geometric morphometric approaches, which allow referral of the new specimen to Anhanguera. Theabsence of a crest in the new specimen of Anhanguera suggests that this structure varies in terms of ontogeny and/or sex, and that perhaps it was influenced by sexual selection.
... Currently, the law in force is The Federal Law of Archaeological, Artistic and Historic Monuments and Zones, published in 1972 [46]. The Instituto Nacional de Antropología e Historia, [26], ( publication retracted by publisher), (b) SMNK PAL 3828, holotype of the pterosaur Ludodactylus sibbicki [28], (c) SMNK 2344 PAL holotype of the pterosaur Tupandactylus navigans [95], (d ) SMNS 58022 holotype of the dinosaur Irritator challengeri [30] (e) SMNK PAL 3804, holotype of the crocodyliform Susisuchus anatoceps [96], ( f ) private collection BMMS BK 2-2, holotype of the putative legged-snake Tetrapodophis amplectus [27], currently interpreted as an aquatic lizard [33], photograph by Michael Caldwell. Abbreviations: SMNK, State Museum of Natural History Karlsruhe, Germany; SMNS, State Museum of Natural History Stuttgart. ...
... This fact, together with the absence of reported exportation permits, leads us to consider that these fossils may have been purchased (figure 5). Some publications state that the fossils were 'obtained from a quarry workman' [92] or 'from a fossil digger' [93] and eight publications [28,30,[94][95][96][97][98][99] directly acknowledge that the specimen was purchased. ...
Article
Full-text available
Scientific practices stemming from colonialism, whereby middle- and low-income countries supply data for high-income countries and the contributions of local expertise are devalued, are still prevalent today in the field of palaeontology. In response to these unjust practices, countries such as Mexico and Brazil adopted protective laws and regulations during the twentieth century to preserve their palaeontological heritage. However, scientific colonialism is still reflected in many publications describing fossil specimens recovered from these countries. Here, we present examples of ‘palaeontological colonialism’ from publications on Jurassic–Cretaceous fossils from NE Mexico and NE Brazil spanning the last three decades. Common issues that we identified in these publications are the absence of both fieldwork and export permit declarations and the lack of local experts among authorships. In Mexico, access to many fossil specimens is restricted on account of these specimens being housed in private collections, whereas a high number of studies on Brazilian fossils are based on specimens illegally reposited in foreign collections, particularly in Germany and Japan. Finally, we outline and discuss the wider academic and social impacts of these research practices, and propose exhaustive recommendations to scientists, journals, museums, research institutions and government and funding agencies in order to overcome these practices.
... Almost all of the alveoli are empty, except for partial teeth in the positions 1, 2, and 3 on the right side, and 1 and 7 on the left side. The partial teeth in the first pair and on the left side show that the lingual surface exhibits a distinct pattern of longitudinal striation, with striae that are relatively broad, shallow and sparsely placed; differently from the deeply excavated and numerous striae seen in anhanguerids such as Anhanguera sp (SMNK 2302)., Ludodactylus sibbicki and Guidraco venator (see Fastnacht 2001;Frey et al. 2003;Wang et al. 2012), while the labial surface exhibits a very fine striation pattern with very subtle, slender striae. The teeth are elliptical in cross-section, with the long axis forming an angle of 15°-22°with the sagittal line of the jaw. ...
... These authors regarded ZIN.PNT-S50-1 to be similar to Anhanguera on the basis of a slight upwards curvature of the symphysis (Unwin and Bakhurina 2000), but we note here that this feature is widespread among anhanguerians (e.g. Kellner and Tomida 2000;Frey et al. 2003;Wang et al. 2012Wang et al. , 2014Holgado et al. 2019) and targaryendraconians (Myers 2010;Kellner et al. 2011;Elgin and Frey 2011). ZIN.PNT-S50-1 exhibits strongly scalloped alveolar margins; subparallel lateral margins of the dentary symphysis in occlusal view; and total symphyseal width about three times alveolar width. ...
Article
Full-text available
Ornithocheirus wiedenrothi, from the Hauterivian (Early Cretaceous of Germany), is a taxon represented by three-dimensional remains of the lower jaw and wing elements. Its phylogenetic affinities have for long been elusive, though several works had already pointed out that it probably did not belong within the wastebasket genus Ornithocheirus. In the present contribution, we redescribe this species, assigning it to the new genus Targaryendraco and offering updated morphological comparisons. Subsequently, we present a phylogenetic analysis in which we recover a clade formed by Targaryendraco, Aussiedraco, Barbosania, Aetodactylus, Camposipterus and Cimoliopterus. This newly recognised clade is interesting in being quite cosmopolitan and spanning from the Hauterivian to the Cenomanian, like its sister-group, the Anhangueria. The recognition of this clade helps fill the temporal gap between the Anhangueria and Cimoliopterus, and also demonstrates that the diversity of Cretaceous toothed pterosaurs was higher than previously thought.
... However, as with all pterosaurs, the record of Ornithocheiridae is extremely patchy. The group was originally based on fragmentary remains from the Early Cretaceous of southern and eastern England (Seeley, 1870;Owen, 1874;Unwin, 2001), but discoveries in Lagerstӓtten such as the Santana and Crato formations of Brazil and the Yixian Formation of China have increased our knowledge of the Cretaceous members of that clade considerably in the last 30 years (Campos and Kellner, 1985;Fastnacht, 2001;Frey et al., 2003;Martill and Frey, 1998;Unwin and Martill, 2007;Veldmeijer, 2003aVeldmeijer, , 2006Wang et al., 2002Wang et al., , 2012Zhou, 2003, 2004;Wellnhofer, 1985Wellnhofer, , 1987 and show that they were more widely distributed. ...
... Subsequent authors (Unwin, 2001;Fastnacht, 2001;Frey et al., 2003;Ibrahim et al., 2010;Martill and Unwin, 2012) considered Siroccopteryx moroccensis to be a species of Coloborhynchus. These authors considered that the deltoid facet with an anteriorly directed first pair of teeth was a character of the genus Coloborhynchus. ...
... 1), Tropeognathus mesembrinus (Wellnhofer, 1987, fig. 2) and Ludodactylus sibbicki (Frey et al., 2003, fig. 1). ...
... The posterior portion of the left maxilla, ventral to the nasoantorbital fenestra, preserves the contact between the maxilla and maxillary process of the jugal (Fig. 4), as described by Molnar and Thulborn (2007). The anterior margin of the nasoantorbital fenestra in Mythunga is similar to that of Ludodactylus (Frey et al., 2003) and Maaradactylus (Bantim et al., 2014), and appears more rounded in comparison to NHMUK PV 39410 , SNSB-BSPG 1982I 89 (Wellnhofer, 1985, Anhanguera piscator (Kellner and Tomida, 2000), AMNH 22555 (Wellnhofer, 1991b) and Guidraco (Wang et al., 2012). ...
Article
Mythunga camara, the most complete pterosaur described from Australia, derives from the Lower Cretaceous (Albian) Toolebuc Formation of central Queensland, Australia. Although it was originally described as an archaeopterodactyloid, more recently Mythunga has been designated as an indeterminate anhanguerid or ornithocheiroid. The holotype specimen is a partial skull, comprising a fragment of the premaxilla, incomplete maxillae and dentaries, and an isolated cranial element preserved in matrix within the nasoantorbital fenestra. Reassessment of the holotype specimen has revealed several new features, which enable a re-evaluation of the phylogenetic position of this taxon. Primary among these observations is that the anteriormost preserved teeth are, in fact, the root of a maxillary tooth and its replacement, rather than a pair of contralateral teeth; thus, they do not represent the anteriormost tooth pair as previously proposed. Redescription of the holotype specimen has also resulted in the identification of several previously unrecognised features, including several nutrient foramina on the maxilla and mandible and shallow channels on the maxilla. Furthermore, the bone lodged within the nasoantorbital fenestra is identified herein as the right splenial. Following the redescription and reinterpretation of the holotype specimen, the phylogenetic position of Mythunga camara was assessed. Although this analysis placed Mythunga within the Anhangueria, most of this clade was reduced to a polytomy; only the clades representing Anhanguera and Boreopterinae + Guidraco were resolved. The classification of Mythunga within Anhangueria indicates that this clade was widespread, and perhaps achieved a global distribution by the late Early Cretaceous.
... The Crato Formation comprises mostly micritic laminated limestone rocks, interpreted as deposits from the shallow waters of a coastal lagoon with both marine and fluvial influences [23]. Although its fossils are generally compressed to some level, these beds are famous for yielding exceptionally preserved remains [23, 26], including a pterosaur patagium [27] , soft tissue crests [20, 21, 28], rhamphothecae [20, 21, 28], and possible pycnofibers [28]. Furthermore, possible muscle fibers of a turtle [29] and several feathers [30][31][32][33]are preserved, some of them having even revealed melanosomes [34]. ...
... The Crato Formation comprises mostly micritic laminated limestone rocks, interpreted as deposits from the shallow waters of a coastal lagoon with both marine and fluvial influences [23]. Although its fossils are generally compressed to some level, these beds are famous for yielding exceptionally preserved remains [23, 26], including a pterosaur patagium [27] , soft tissue crests [20, 21, 28], rhamphothecae [20, 21, 28], and possible pycnofibers [28]. Furthermore, possible muscle fibers of a turtle [29] and several feathers [30][31][32][33]are preserved, some of them having even revealed melanosomes [34]. ...
Article
Full-text available
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.
... Entre os tetrápodes, os pterossauros são os mais diversos e abundantes da sucessão Lagerstätte Crato com sete espécies formalmente descritas, Arthurdactylus conandoylei, Aymberedactylus cearensis, Brasileodactylus araripensis, Lacusovagus magnificens, Ludodactylus sibbicki, Tupandactylus imperator e Tupandactylus navigans, distribuídas em três famílias, Tapejaridae, Thalassodromidae e Ornithocheiridae (Kellner, 1984;Frey & Martill, 1994;Campos & Kellner, 1997;Frey et al., 2003a;2003b;Beccari et al., 2021). A espécie T. navigans constitui um importante registro para os tapejarídeos devido à presença de um esqueleto articulado e praticamente completo com preservação das feições morfológicas pós-cranianas (Beccari et al., 2021). ...
Chapter
O rifteamento do Gondwana resultou em mudanças importantes na composição química e distribuição de nutrientes dos oceanos, além de ter contribuído para modificações na circulação de massas de água, o que acarretou na abertura de novos espaços ecológicos. Com a reativação de sistemas de falhas do embasamento durante a ruptura, tem-se a formação de lagos de riftes perenes e temporários, por vezes alimentados por incursões fluviais. Além disso, os pulsos de incursões marinhas durante os eventos de transgressão no final do Cretáceo Inicial também geraram importantes modificações na biota da Bacia do Araripe. A gênese desses novos ecoespaços resultou em grandes transformações bióticas, criando novas possibilidades para o surgimento, a diversificação e a extinção de espécies em um curto espaço temporal. As rochas da Bacia do Araripe registram este momento em que ficam evidentes as relações intrínsecas entre as alterações dos espaços ecológicos e as consequentes biotas registradas em ambientes continentais e marinhos. A integração entre os dados sedimentológicos, estratigráficos e paleontológicos das rochas da Bacia do Araripe permite categorizar três biotas, Missão Velha, Crato e Romualdo, produtos das dinâmicas tectônicas globais associadas à ruptura do Gondwana e suas decorrentes variações climáticas e oceanográficas. Essas biotas apresentam registros paleobiológicos singulares, com animais e vegetais abundantes, diversificados e muitas vezes de caráter endêmico. Muitos dos fósseis identificados são espécies com preservação excepcional, que possibilitam o entendimento de aspectos-chave para a evolução biológica e a relação com as espécies viventes.
... Based on what is preserved of the premaxilla, it is clear that the premaxillary crest is confined to the anterior third of the skull. The presence of a large, blade-like premaxillary crest differentiates Ferrodraco from other anhanguerians which lack crests, such as Liaoningopterus gui (Wang and Zhou, 2003), Ludodactylus sibbicki (Frey et al., 2003), Guidraco venator (Wang et al., 2012) and Cearadactylus atrox (Vila Nova et al., 2014). The premaxillary crest of Ferrodraco is confluent with the anterior margin of the skull, and thus differs from Anhanguera blittersdorffi (Wellnhofer, 1987), Anhanguera sp. ...
Article
Full-text available
Ferrodraco lentoni, an anhanguerid from the Upper Cretaceous Winton Formation of northeast Australia, is the most complete Australian pterosaur described to date, represented by a partial cranium, incomplete cervical series and wing elements. Herein we present a comprehensive osteological description of Ferrodraco, as well as an emended diagnosis for this taxon. In addition, we compare Ferrodraco with other isolated pterosaur remains from Australian Cretaceous deposits. Subtle, yet salient, differences indicate that at least three of these specimens, all derived from the upper Albian Toolebuc Formation, are distinct from Ferrodraco. However, we are uncertain whether these specimens are attributable to Mythunga camara, Aussiedraco molnari, Thapunngaka shawi, or an as yet un-named taxon. Detailed description of the postcranial material of Ferrodraco also provides an opportunity to reassess its phylogenetic position. In one analysis, Ferrodraco and Mythunga are resolved as sister taxa within Tropeognathinae, whereas in another, Ferrodraco, Mythunga, and Tropeognathus form a polytomy within Coloborhynchinae. Either way, these slight differences notwithstanding, a close relationship between Ferrodraco and Mythunga is evident, supporting the interpretation that they form a clade. By contrast, Aussiedraco molnari is resolved as a member of Targaryendraconia, a clade with a cosmopolitan distribution. The presence of several anhanguerian taxa or lineages in the late Early and early Late Cretaceous of northeast Australia is suggestive of even greater diversity in the Australian pterosaur fauna.
... Among these are the world famous Konservat-Lagerstätten (sense Seilacher, 1970;Seilacher et al., 1985;Itano, 2019) recorded in the Aptian Crato and Romualdo formations of the Santana Group, Araripe Basin (Maisey, 1991;Martill et al., 2007;Varejão et al., 2019a;Ribeiro et al., 2021) and the Upper Jurassic-Lower Cretaceous Muzinho Shale, Patos Bons Formation, Parnaíba Basin (Cardoso et al., 2020). Thanks to their exceptional preservation, many of the fossils have significantly contributed to our understanding of the paleobiology of various groups, including pterosaurs (Kellner, 1984;Frey and Martill, 1994;Campos and Kellner, 1997;Frey et al., 2003aFrey et al., , 2003bWitton, Freitas et al. (2017). C. Detail of the sedimentary succession of the Amargosa Bed in the Tucano Basin, according to Freitas et al. (2017) and Varejão et al. (2019b). ...
Article
We report the Amargosa Biota from the middle part of the Lower Cretaceous Marizal Formation (Central Tucano Sub-Basin, NE Brazil), as a new Konservat-Lagerstätte. Exceptionally preserved fossils are confined to the lower part of an up to 15-m-thick, mud-dominated succession, named Amargosa Bed. Seven bedding planes (L0-L6) with distinct sedimentological and taphonomic attributes were identified in the type section (Amargosa Village, Euclides da Cunha County, Bahia State), distributed in an ~1-m-thick succession of well-laminated claystone, mudstone, siltstone, and very fine-grained sandstone. These contain ostracods, spinicaudatan carapaces, palaemonid shrimps, fish, and comminuted plant remains. Fossils occur in high concentration on at least four bedding planes (i.e., L2, L3, L5, and L6), forming polytypical assemblages that are dominated by one of the fossil groups. Assemblages are formed mainly by autochthonous to parautochthonous elements, representing variable, but limited, temporal mixing. A key attribute of some fossil-rich strata (L3, L5, and L6) is the preservation of poorly biomineralized organisms and/or of complete soft-bodied parts, which are typically prone to destruction due to rapid decay or bioturbation. The polytypical nature of these fossil assemblages, interbedded with non-fossiliferous intervals, suggests mass mortality events, probably caused by abrupt changes in water parameters (anoxia, salinity, pH, among others). The dark greenish gray color (yellowish when weathered), and the finely laminated nature of the claystone, siltstone, and mudstone containing members of the Amargosa Biota indicates that the benthic infaunal life was absent or, at least, very scarce in a locally, relatively deep, oxygen-poor lake bottom. Anoxia and high salinity, linked with local semi-arid conditions during the Lower Cretaceous may have played key roles in the exceptional preservation of some fossils (shrimps, fish). Finally, our data provide a more comprehensive understanding of the temporal distribution of taxa and taphonomic processes associated with the complex genesis of the fossil-bearing interval of the Amargosa Bed in its type locality.
... Hence, this is termed the maxillojugal bar. The jugal maxillary ramus extends along the dorsal margin of the bar and terminates anteriorly with a sharp tip posterior to the nasoantorbital fenestra anterior margin, unlike the Pteranodontia and the clade containing the Ornithocheirae and Ludodactylus sibbicki Frey et al., 2003a(Andres, 2021. A thickened ventral margin of the maxillojugal bar anterior to the left jugal main body in TMM 41961-1.1 is identified as the posterior extent of the maxilla, and this is confirmed in crosssection. ...
Article
Full-text available
Quetzalcoatlus is the largest flying organism ever known and one of the most familiar pterosaurs to the public. Despite a half century of interest, it remains very incompletely described. This shortfall is addressed here through a full morphological description of Quetzalcoatlus and the other pterosaur material of Big Bend National Park, Texas. The first reported material was described and named Quetzalcoatlus northropi by Douglas Lawson in 1975, but in two separate publications. A ruling by the International Commission of Zoological Nomenclature was required for the name to be made available. Review of the pterosaur fauna of the Park recovers three valid species of azhdarchid pterosaurs in the latest Cretaceous Period Javelina and Black Peaks formations. The size and occurrence of these species are correlated with depositional environment. The holotype of the giant Quetzalcoatlus northropi and six other giant specimens referred to it occur in stream-channel deposits, including the youngest reported pterosaur. The vast majority of specimens (200+) are from large pterosaurs found in the abandoned channel-lake deposits at Pterodactyl Ridge; they form a diagnosable natural group erected as the new species Quetzalcoatlus lawsoni. A moderate-sized partial skull and cervical series also found in the abandoned channel-lake deposits at Pterodactyl Ridge, but lower in the section, is distinct from both species and is erected as Wellnhopterus brevirostris, gen. et sp. nov. Overbank flood-plain facies preserve another eleven specimens of extreme size variation, including small azhdarchids. The Big Bend pterosaur fauna provides the greatest known sample of azhdarchid pterosaurs and three-dimensional pterosaur morphology.
... The Araripe Basin in northeastern Brazil -a region of low socioeconomic status -where 'Ubirajara' was found has suffered from illicit fossil trafficking for decades 4,5 . In fact, some foreign researchers openly admit to having obtained Araripe fossils from commercial sources or fossil dealers (for example, the dinosaur Irritator challengeri, the crocodilian Susisuchus anatoceps and the pterosaurs Ludodactylus sibbicki, Lacusovagus magnificens, Unwindia trigonus and Tupandactylus navigans [6][7][8][9][10][11] ). In one notorious case, researchers even conceded they were "deceived by the fossil dealers" 9 . ...
Article
To the Editor — The dinosaur fossil ‘Ubirajara jubatus’ was allegedly exported from Brazil in 1995 and is currently held by State Museum of Natural History Karlsruhe (SMNK) in Germany. It has attracted the attention of the scientific community not only because of its intrinsic interest but also for the controversy surrounding its export from Brazil, which has put into question the legal and ethical context of this work and led to the removal of the publication. Note that ‘Ubirajara jubatus’ is no longer a valid scientific name due to this retraction, hence its appearance in quotation marks here. ‘Ubirajara’ is far from a unique case of an illegally acquired and exported fossil residing in a foreign museum. The Araripe Basin in northeastern Brazil — a region of low socioeconomic status — where ‘Ubirajara’ was found has suffered from illicit fossil trafficking for decades. In fact, some foreign researchers openly admit to having obtained Araripe fossils from commercial sources or fossil dealers (for example, the dinosaur Irritator challengeri, the crocodilian Susisuchus anatoceps and the pterosaurs Ludodactylus sibbicki, Lacusovagus magnificens, Unwindia trigonus and Tupandactylus navigans). In one notorious case, researchers even conceded they were “deceived by the fossil dealers”. Many of the illicit fossils, some of which are holotypes (specimens used to designate new species), are stored in German museums. Like many other Latin American countries, Brazil adopted rigid laws...
... Pterosaurs (Archosauria: Avemetatarsalia: Pterosauromorpha) are the most diverse and abundant group of tetrapods in the CKL fossil record. To date, the following six species are reported: Arthurdactylus conandoylei (Pterodactyloidea: Ornithocheiridae) (Frey and Martill, 1994); Aymberedactylus cearensis (Pterodactyloidea: Tapejaridae) (Pêgas et al., 2016); Brasileodactylus araripensis (Pterodactyloidea: Ornithocheiridae) (Kellner, 1984); Lacusovagus magnificens (Pterodactyloidea) (Witton, 2008); Ludodactylus sibbicki (Pterodactyloidea: Ornithocheiridae) (Frey et al., 2003a); Tupandactylus imperator (Pterodactyloidea: Tapejaridae) (Campos and Kellner, 1997); and Tupandactylus navigans (Pterodactyloidea: Tapejaridae) (Frey et al., 2003b). Pterosaurs are completely extinct lineages; hence, their biology and lifestyles are mostly unknown and the focus of much debate. ...
Article
An alternative hypothesis concerning the paleoecological and paleoenvironmental depositional conditions of the Crato Konservat-Lagerstätte (CKL), Crato Formation, Aptian, NE Brazil, one of the most extraordinary Gondwana fossil sites, is proposed. Following an actualistic approach, the ecology of extant relatives of the most abundant and diverse fossil groups recorded in the CKL (i.e., vascular plants, arthropods, fishes, and tetrapods) is considered. Data is based on an extensive literature review followed by a re-examination of recently collected fossils. This approach allowed a detailed appraisal of the stratigraphic/ecological distribution of the main fossil groups preserved in the CKL. Plant and animal groups are recorded in three main stratigraphic intervals, named Intervals I–III in ascending order. Most fossils are to be considered autochthonous to parautochthonous and have been preserved in distinct stages of base-level fluctuations within a shallow lacustrine depositional system, subject to periodic flooding in large, depressed areas. Exceptional preservation in such environments was mediated by microbially-induced processes (i.e., microbial mat entombment), mostly in the coastal areas of the alkaline lake. Based on the distinct sedimentary facies and autecological attributes of dominant paleo-bioindicators, a new paleoenvironmental model for the CKL is proposed, encompassing a seasonal, semi-arid, shallow lacustrine wetland. Faunal and floral content were ecologically arranged in long-lasting aquatic zones, surrounded by periodically flooded mesophytic ecotones and outer xeric habitats, as in the modern alkaline lake Chad in Africa. Our data show the relevance of multiproxy analyses (i.e., paleontological, sedimentological, geochemical, and stratigraphic) of exceptional fossil sites for assessing paleoenvironmental conditions in depositional settings subject to continuous base-level changes, such as those existing in complex, present-day wetland ecosystems. The recognition of key parameters in ancient wetlands is of great importance concerning the formation of non-marine Konservat- Lagerstätten in the geological record.
... Fastnacht (2001) modified the diagnosis after Lee (1994) add ing the following characters: upper jaw laterally expanded in a spoonshape in dorsal view from the second to the fourth pair of alveoli; first pair of teeth projecting anteriorly from the blunt anterior margin of the upper jaw at a significant eleva tion above the palate relative to subsequent teeth; second and third pair of alveoli of the upper and lower jaw enlarged rela tive to other alveoli; lower jaw with medial crest rising from its anterior end; and lower jaw laterally expanded in a spoon shape from the first to the third pair of alveoli (Fastnacht 2001). This new diagnosis was considered in the subsequent literature by some authors (e.g., Unwin 2001Unwin , 2002Unwin , 2003Frey et al. 2003;Veldmeijer 2003Veldmeijer , 2006Lü et al. 2006;Martill and Naish 2006;Fastnacht 2008;Martill and Unwin 2012;Witton 2013;Martill 2015;MartinSilverstone et al. 2018). Unwin (2001), accepting the revalidation of Colo borhynchus, followed the works of Lee (1994) and Fastnacht (2001) and further added other species to this genus. ...
Article
Full-text available
Anhanguerids are a particular group of pterodactyloid pterosaurs, characterized mainly by their rostral sagittal crests, well laterally expanded jaw tips and enlarged anterior teeth. Due to the fragmentary nature of most known specimens, including holotypes, the taxonomy of the group has proved particularly difficult and controversial. Coloborhynchinae is a recently proposed clade within the Anhangueridae, and was defined as the most inclusive clade containing Coloborhynchus clavirostris but not Anhanguera or Ludodactylus. Coloborhynchinae was originally thought to include Coloborhynchus, Uktenadactylus, and Siroccopteryx. Here we present a reassessment of the taxonomy and phylogeny of all proposed members of the Coloborhynchinae and Coloborhynchus complex, with new anatomical comparisons and a novel phylogenetic analysis. Several features allow us to establish that coloborhynchines were much more diverse than previously thought, englobing four genera and seven species: Aerodraco sedgwickii gen. et comb. nov., Coloborhynchus clavirostris, Nicorhynchus capito gen. et comb. nov., Nicorhynchus fluviferox gen. et comb. nov., Uktenadactylus rodriguesae sp. nov., and Uktenadactylus wadleighi. Nicorhynchus and Uktenadactylus are considered sister taxa, being distinct on the basis of several rostral characters. Although with a homoplastic flat rostrum surface, Siroccopteryx was recovered out of the Coloborhynchinae, as sister taxon of Tropeognathus, due to similarities on the palatal ridge (which is broad and deep, and starting at the same level) and the relatively stout teeth compared to other anhanguerids. Tropeognathus and Siroccopteryx are further related to the Australian taxa Ferrodraco and Mythunga, which are all grouped in a new clade: the Tropeognathinae. Our analysis suggests that morphological evolution within anhanguerids was quite more complex than previously thought, with coloborhynchines representing the oldest recorded lineage of Anhangueridae, which achieved a worldwide distribution at least from the Aptian to the Cenomanian.
... Fossilized fish are dominated by Dastilbe crandalli, Cladocyclus, Lepidotes and Araripelepidotes (Davis and Martill, 1999), with Vinctifer, Cladocyclus, Rhacolepis, Notelops, Mawsonia and Axelrodichthys also being present in the Romualdo Formation. Further, well preserved wing membranes and wing fibres, claw sheaths, foot webs, and a heel pad of pterosaurs (Arthurdactylus, Ludodactylus, Ingridia, Santanadactylus, Araripesaurus, Cearadactylus, Brasileodactylus, Anhanguera, Lacusovagus) have been discovered from the Crato and Romualdo formations (Martill and Unwin, 1989;Martill and Frey, 1998;Frey et al., 2003;Unwin and Martill, 2007;Witton, 2007), with additional dinosaurs (Irritator, Angaturama, Santanaraptor and Mirischia) being known in the Romualdo Formation (Kellner, 1996(Kellner, , 1999Martill et al., 1996Martill et al., , 2000. ...
Article
The Ipubi Formation of the Santana Group, Araripe Basin, Brazil, is characterized by black shales and overlying evaporite deposits and is suggested to record the transition from lacustrine to marine depositional environments. To date, the age of the black shales, constrained only by microfossils, is poorly determined, with ages spanning ∼25 myrs from 125 to 100.5 Ma (Aptian-Albian). Here we present new Re–Os elemental and isotopic data to provide the first absolute age for those rocks of the Ipubi Formation and an improved understanding of the depositional paleoenvironment of the Araripe Basin. The Re–Os isotope data for Ipubi Formation black shales yield a depositional age of 123 ± 3.5 Ma, with a highly radiogenic initial ¹⁸⁷Os/¹⁸⁸Os composition (Osi) of 1.97 ± 0.02. The Re–Os age indicates that the deposition of the Ipubi Formation black shales occurred during the Late Barremian/Early Aptian, prior to the onset of OAE 1a, in a highly restricted marine/lacustrine setting.
... The total length of the preserved portion of the specimen, restored as in life, is estimated at 150 mm, although it is possible that the gaps between preserved fragments were larger than assumed here. Typically, in ornithocheirid pterosaurs, the length of the mandible is approximately fifteen times its depth just anterior to the articular region (based on seven examples; Frey et al., 2003;Veldmeijer, 2005). On this basis the mandible of UR CP 0002 is likely to have reached approximately 0.4 m in length, comparable in size to the mandible of Anhanguera santanae Wellnhofer, 1991 (AMNH 22555) which has an estimated wingspan of 4.0 m. ...
Article
The global fossil record of lowermost Cretaceous pterosaurs is meagre, and much of the material is fragmentary. Here we report three occurrences of pterosaurs from the Rosablanca Formation (Valanginian), the first records of this extinct group of flying reptiles from Colombia. Specimens from Zapatoca, Santander Department, consist of fragments of the left mandible and the proximal portion of a wing phalange. A third specimen, from Cundinamarca Department, is the proximal termination of a radius. Although fragmentary, these remains are clearly pterosaurian, on account of their remarkably thin bone walls and provide evidence of pterodactyloids, including a large non-pteranodontian ornithocheiroid, in the northernmost part of South America in the early Lower Cretaceous.
... Tooth morphology and size can vary quite considerably along the tooth row in ornithocheirids (e.g. Campos and Kellner 1985, Wellnhofer 1985, Kellner and Tomida 2000, Frey et al. 2003, Veldmeijer 2003, Wang et al. 2012. Moreover, this variation was likely compounded by allometric shape changes related to ontogeny, with hatchlings of 0.3-0.4 ...
Article
Full-text available
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
... In some specimens the apex is frayed (Fig. 2B). W. brasiliense has also been recorded in association with the pterosaur Ludodactylus; it was first thought that the rigid and sharp leaf pierced the gular of the pterosaur leading to its eventual death (Frey et al., 2003;Martill et al., 2007, Fig. 17.4). An alternative theory is that this is in fact an abiotic association, due to the hyoid apparatus being preserved on top of the Welwitschiophyllum leaf, rather than wrapping around either side of the leaf (Witton, 2017), which we deem less likely given the positioning of the leaf inside the mandibular apparatus. ...
Article
The Lower Cretaceous Crato Formation of north-east Brazil yields a diverse plant assemblage. It has yielded many macrofossils thought to be related to the enigmatic gymnosperm group Gnetales, including the long leaf Welwitschiophyllum brasiliense Dilcher et al., 2005. This fossil plant is considered to be related to the extant gnetalean Welwitschia mirabilis Hooker, 1863, despite lacking many gnetalean characteristics. Presently, this macrophyte fossil is known only from detached leaves and much anatomical information is currently unavailable. The reproductive structures assigned to the family Welwitschiaceae in the Crato Formation, as well as several key morphological features of the leaves are currently thought to relate fossil Welwitschiophyllum to extant Welwitschia. These leaf characters include: isobilateral leaf form, triangular elongated leaf shape with a wide base, longitudinal splitting from a frayed leaf apex, numerous parallel veins and possible thickening of the epidermis. However, many of these leaf characteristics also occur in many other macrophytes, perhaps as a result of convergence. Anatomical and morphological data described here from fossil Welwitschiophyllum leaves is compared with extant Welwitschia. Our results show that Welwitschiophyllum can only be placed tentatively in Gnetales, as many of the features we report are not diagnostic, and may have resulted from convergent evolution (e.g., gum). Fossils with better anatomical preservation and the reconstruction of the whole plant are really needed to better understand the affinities of Welwitschiophyllum.
... The offset placement of alveoli and a ventral crista are present in both A. piscator and A. robustus, but is also seen in several other ornithocheirids, e.g. Ludodactylus Frey et al., 2003. The ventral crista, however, is deeper with a steeper posterior margin in A. robustus. ...
Article
Pterodactyloid pterosaurs underwent a diversification in the Late Jurassic and Early Cretaceous, followed by a major turnover event in the mid-Cretaceous, when ornithocheiroids and basal azhdarchoids were replaced by pteranodontids, nyctosaurids and azhdarchids in the latest Cretaceous. However, precise patterns of turnover are obscured by the incompleteness of the pterosaur fossil record. Fossils from the middle Cretaceous Kem Kem beds of Morocco (?Albian –Cenomanian) have helped shed light on the diversity of pterosaurs from this time and provide a window into the diversity of a continental pterosaur assemblage from this critical transitional period. Two toothed pterosaurs, the ornithocheirids Siroccopteryx moroccensis and Coloborhynchus fluviferox, have been reported from the Kem Kem beds. Here, we report a partial mandible and two premaxillae representing three additional taxa of toothed pterosaurs. The mandibular symphysis closely resembles that of Anhanguera piscator from the Romualdo Member of the Santana Formation of Brazil in the arrangement and spacing of the alveoli, the weak anterior upturn of the jaw, and the ventral crest. One premaxilla closely resembles that of the ornithocheirid Ornithocheirus simus from the Cambridge Greensand Formation of eastern England. A second premaxilla is referred to Coloborhynchus, bearing similarities to C. clavirostris from the Hastings Group of southern England, and C. fluviferox from the Kem Kem beds of Morocco. In total, the Kem Kem pterosaur fauna includes at least nine species, of which three are ornithocheirids. The Kem Kem assemblage supports the idea that toothed pterosaurs remained diverse during the mid Cretaceous before disappearing from post-Cenomanian strata.
... 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). ...
Article
Full-text available
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.
... Piscivory interpretations are based on content fossils, associations, ichnology, isotope analyses, comparative anatomy and functional morphology (Fig. 3). Most ornithocheirids are known from lagoonal, coastal and marine deposits (Frey, Martill & Buchy, 2003;Chatterjee & Templin, 2004;Unwin, 2006;Molnar & Thulborn, 2007;Unwin & Martill, 2007;Kear, Deacon & Siverson, 2010;Veldmeijer et al., 2012;Wretman & Kear, 2013). A coprolite interpreted to be from Guidraco venator contains fish bones (Wang et al., 2012). ...
Article
Full-text available
Pterosaurs are an extinct group of Mesozoic flying reptiles, whose fossil record extends from approximately 210 to 66 million years ago. They were integral components of continental and marginal marine ecosystems, yet their diets remain poorly constrained. Numerous dietary hypotheses have been proposed for different pterosaur groups, including insectivory, piscivory, carnivory, durophagy, herbivory/frugivory, filter‐feeding and generalism. These hypotheses, and subsequent interpretations of pterosaur diet, are supported by qualitative (content fossils, associations, ichnology, comparative anatomy) and/or quantitative (functional morphology, stable isotope analysis) evidence. Pterosaur dietary interpretations are scattered throughout the literature with little attention paid to the supporting evidence. Reaching a robustly supported consensus on pterosaur diets is important for understanding their dietary evolution, and their roles in Mesozoic ecosystems. A comprehensive examination of the pterosaur literature identified 314 dietary interpretations (dietary statement plus supporting evidence) from 126 published studies. Multiple alternative diets have been hypothesised for most principal taxonomic pterosaur groups. Some groups exhibit a high degree of consensus, supported by multiple lines of evidence, while others exhibit less consensus. Qualitative evidence supports 87.3% of dietary interpretations, with comparative anatomy most common (62.1% of total). More speciose groups of pterosaur tend to have a greater range of hypothesised diets. Consideration of dietary interpretations within alternative phylogenetic contexts reveals high levels of consensus between equivalent monofenestratan groups, and lower levels of consensus between equivalent non‐monofenestratan groups. Evaluating the possible non‐biological controls on apparent patterns of dietary diversity reveals that numbers of dietary interpretations through time exhibit no correlation with patterns of publication (number of peer‐reviewed publications through time). 73.8% of dietary interpretations were published in the 21st century. Overall, consensus interpretations of pterosaur diets are better accounted for by non‐biological signals, such as the impact of the respective quality of the fossil record of different pterosaur groups on research levels. That many interpretations are based on qualitative, often untestable lines of evidence adds significant noise to the data. More experiment‐led pterosaur dietary research, with greater consideration of pterosaurs as organisms with independent evolutionary histories, will lead to more robust conclusions drawn from repeatable results. This will allow greater understanding of pterosaur dietary diversity, disparity and evolution and facilitate reconstructions of Mesozoic ecosystems.
... Piscivory interpretations are based on content fossils, associations, ichnology, isotope analyses, comparative anatomy and functional morphology (Fig. 3). Most ornithocheirids are known from lagoonal, coastal and marine deposits (Frey, Martill & Buchy, 2003;Chatterjee & Templin, 2004;Unwin, 2006;Molnar & Thulborn, 2007;Unwin & Martill, 2007;Kear, Deacon & Siverson, 2010;Veldmeijer et al., 2012;Wretman & Kear, 2013). A coprolite interpreted to be from Guidraco venator contains fish bones (Wang et al., 2012). ...
Conference Paper
Full-text available
Pterosaurs were a successful group of Mesozoic flying reptiles. For 150 million years they were integral components of terrestrial and coastal ecosystems, yet their feeding ecology remains poorly constrained. Postulated pterosaur diets include insectivory, piscivory and/or carnivory, but many dietary hypotheses are little more than speculation based on scant evidence. We have developed a more robust approach based on quantitative analysis of the micron-scale 3D textures of worn pterosaur tooth surfaces – dental microwear texture analysis – never before applied to pterosaurs. Microwear is produced as scratches and chips generated by food items create characteristic surface textures on teeth that vary according to diet. We compared microwear from non-occlusal tooth surfaces of 11 species of pterosaur with data from extant organisms with known diets, (bats, monitor lizards and crocodilians, including insectivorous, piscivorous and carnivorous species). This allowed for robust testing of previous pterosaur dietary hypotheses. Microwear from Dimorphodon for example, previously hypothesised as a piscivore, indicates a diet of vertebrates and invertebrates. Microwear from basal monofenestratans, previously hypothesised as carnivores, provides evidence of piscivory in these pterosaurs. Dietary evidence from microwear therefore provides novel insights into the ecological roles of respective pterosaurs and pterosaur dietary evolution.
... Although I do refer the reader to the discussion presented previously (Kellner 2010), just for the sake of argumentation, I will highlight two points regarding the diagnosis employed by them in the taxonomic discussion of UALVP 24238: the presence of a cranial crest made by the frontals directed up and back, and the presence of proximal caudal vertebrae with duplex centra. Cranial crests made by the frontals have since been reported in other pterosaurs such as Guidraco and Ludodactylus (Frey et al. 2003, Wang et al. 2012 and proximal caudals with duplex centra were reported in Anhanguera (Kellner & Tomida 2000). That said, regarding the cranial crest, a highly controversial topic these days (see Cheng et al. 2017), I hardly believe that Guidraco or Ludodactylus should be classified into the genus Pteranodon, but just would like to stress that diagnoses can change with the advent of new specimens or more information. ...
... Some atoposaurid crocodylomorphs of the Jurassic and Cretaceous have extreme lateral compression, but not with backwardly deflected and inwardly curved crown apices, and always with strong eocristae and accesory ridges (Salisbury 2002;Oreska et al. 2013). Lateral compression and recurved apices occur among dromaeosaurid and ceratosaurid theropod dinosaurs, though never to the extreme seen in HMNS/BB 5032; these dinosaurs always have strong, serrated posterior Frey et al. (2003)), Dorygnathus (data from Wellnhofer (1991)) and Archaeopteryx (outline from Wellnhofer (2009)). Outer (buccal) view of premaxillary tooth edited from Feduccia (2012). ...
Article
Pterosaur fossils are rare in the Morrison Formation, and most are poorly preserved. The Breakfast Bench Facies (BBF) at Como Bluff produces incomplete but uncrushed limbs. One proximal and two distal femora match a complete femur (BYU 17214) referred to Mesadactylus. Unexpectedly, both of the BBF distal femora possess a large intercondylar pneumatopore. BYU 17214 also possesses an intercondylar pneumatopore, but it is smaller than in the BBF femora. Distal femoral pnuematicity is previously recognized only in Cretaceous azhdarchoids and pteranodontids. A peculiar BBF jaw fragment shows strongly labiolingually compressed, incurved crowns with their upper half bent backwards; associated are anterior fangs. We suspect this specimen is a previously undiagnosed pterosaur. Additional BBF material documents a diverse pterosaur fauna including a humerus with a greatly expanded ectepicondyle possibly from a non-pterodactyloid monofenestratan.
... However, more recently described specimens challenge some of these features and show that they are more widespread among dsungaripteroid pterosaurs. Characters (1) and (2) are present on Ludodactylus sibbicki from the Crato Formation (Frey, Martill & Buchy, 2003), and characters (4), (5) and (6) are also found in Brasileodactylus sp. (SNSB-BSPG 1991 I 27;Veldmeijer, Meijer & Signore, 2009) and in Istiodactylus (Hooley, 1913;Andres & Ji, 2006). ...
Article
Full-text available
Background Anhanguerids comprise an important clade of pterosaurs, mostly known from dozens of three-dimensionally preserved specimens recovered from the Lower Cretaceous Romualdo Formation (northeastern Brazil). They are remarkably diverse in this sedimentary unit, with eight named species, six of them belonging to the genus Anhanguera. However, such diversity is likely overestimated, as these species have been historically diagnosed based on subtle differences, mainly based on the shape and position of the cranial crest. In spite of that, recently discovered pterosaur taxa represented by large numbers of individuals, including juveniles and adults, as well as presumed males and females, have crests of sizes and shapes that are either ontogenetically variable or sexually dimorphic. Methods We describe in detail the skull of one of the most complete specimens referred to Anhanguera, AMNH 22555, and use it as a case study to review the diversity of anhanguerids from the Romualdo Formation. In order to accomplish that, a geometric morphometric analysis was performed to assess size-dependent characters with respect to the premaxillary crest in the 12 most complete skulls bearing crests that are referred in, or related to, this clade, almost all of them analyzed first hand. Results Geometric morphometric regression of shape on centroid size was highly statistically significant (p = 0.0091) and showed that allometry accounts for 25.7% of total shape variation between skulls of different centroid sizes. Premaxillary crests are both taller and anteroposteriorly longer in larger skulls, a feature consistent with ontogenetic growth. A new diagnosis is proposed for Anhanguera, including traits that are nowadays known to be widespread within the genus, as well as ontogenetic changes. AMNH 22555 cannot be referred to “Anhanguera santanae” and, in fact, “Anhanguera santanae”, “Anhanguera araripensis”, and “Anhanguera robustus” are here considered nomina dubia. Discussion Historically, minor differences in crest morphology have been used in the definition of new anhanguerid species. Nowadays, this practice resulted in a considerable difficulty in referring well-preserved skulls into known taxa. When several specimens are analyzed, morphologies previously believed to be disparate are, in fact, separated by a continuum, and are thus better explained as individual or temporal variations. Stratigraphically controlled excavations on the Romualdo Formation have showed evidence for faunal turnover regarding fish communities. It is thus possible that some of the pterosaurs from this unit were not coeval, and might even represent anagenetic morphotypes. Unfortunately, amateur collecting of Romualdo Formation fossils, aimed especially at commerce, resulted in the lack of stratigraphic data of virtually all its pterosaurs and precludes testing of these further hypotheses.
... [52,53]), amphibians (e.g. [54,55]), pterosaurs [56][57][58][59][60][61][62], crocodylomorphs [17,19] and feathers (e.g. [63]). ...
Article
Full-text available
Susisuchus anatoceps is a neosuchian crocodylomorph lying outside the clade Eusuchia, and associated with the transition between basal and advanced neosuchians and the rise of early eusuchians. The specimen MPSC R1136 comprises a partially articulated postcranial skeleton and is only the third fossil assigned to this relevant taxon. Thin sections of a right rib and right ulna of this specimen have been cut for histological studies and provide the first paleohistological information of an advanced non-eusuchian neosuchian from South America. The cross-section of the ulna shows a thick cortex with 17 lines of arrested growth (LAGs), a few scattered vascular canals, and primary and secondary osteons. This bone has a free medullary cavity and a spongiosa is completely absent. Thin sections of the rib show that remodeling process was active when the animal died, with a thin cortex and a well-developed spongiosa. In the latter, few secondary osteons and 4 LAGs were identified. According to the observed data, Susisuchus anatoceps had a slow-growing histological microstructure pattern, which is common in crocodylomorphs. The high number of ulnar LAGs and the active remodeling process are indicative that this animal was at least a late subadult, at or past the age of sexual maturity. This contradicts previous studies that interpreted this and other Susisuchus anatoceps specimens as juveniles, and suggests that full-grown adults of this species were relatively small-bodied, comparable in size to modern dwarf crocodiles.
... Various authors have suggested Brasileodactylus to be synonymous with Anhanguera (Unwin 2001) or Coloborhynchus (Frey et al. 2003). As remarked by Veldmeijer & Signore (2004) "The explanation of the supposed difference as the result of ontogeny, sexual dimorphism or variation cannot be proven, mainly due to the scanty fossil record (most of the species are represented by only one (published) specimen, often consisting only of parts of the skull); the fossils should therefore be treated as a different species unless proven (and not just suggested) otherwise." ...
... Comparison to more complete material from the Santana Group (Araripe Basin, northeastern Brazil) showed that they closely resemble anhanguerid teeth, such as the long and slender teeth of Anhanguera blittersdorffi Campos & Kellner, 1985 (Pz-DBAV-UERJ 40;Kellner & Tomida, 2000: fig. 66b) from the Romualdo Formation (Aptian/Albian; Beurlen, 1971) and the possibly related Ludodactylus sibbicki Frey, Martill & Buchy, 2003 (SMNK PAL 3828;Frey et al., 2003) (Figure 5) from the Crato Formation (Aptian; Beurlen, 1971), where the first teeth also show longitudinal lines, such as BMNH R 8662, and similar shape and curvature of the crown. It is noteworthy that this morphology is present in several pterosaurs from the Romualdo and Crato formations of the Araripe Basin. ...
Article
The reported occurrences of pterosaur specimens from the Lower Cretaceous Recôncavo Basin (Bahia), northeastern Brazil, is reviewed herein. All material was described by the British paleontologist A. S. Woodward and is housed at the Natural History Museum in London. The review confirms that all isolated and incomplete quadrates first regarded by Woodward as pterosaurian were later correctly referred to an osteichthyan coelacanthid species, and possibly represent Mawsonia gigas. Two isolated teeth were also found, one of which (BMNH R 8662) is likely the one briefly mentioned by Woodward in 1907 as pterosaurian. Those specimens, described and figured here for the first time, belong to a pterodactyloid pterosaur with affinities to the Anhangueridae. Despite not presenting new morphological data, the pterosaur tooth BMNH R 8662 is of historical importance since it is the first pterosaur from South America to be recorded in the literature. Furthermore, this material shows the presence of anhanguerid-like pterosaurs in the Lower Cretaceous of Bahia, extending the geographical record of this group in Brazil.
... On the posterior side of this foramen position, an apparently unique process directed posteriorly in the orbit is present. A lacrimal spine in Ludodactylus, which is more slender than the process of this specimen, was the only similarity the process in B. giganticus (Frey et al., 2003a). The condition of an extensively fenestrated lacrimal has been reported only in some tapejarids, such as Tapejara wellnhoferi (Kellner, 1989), Sinopterus dongi (Wang and Zhou, 2003a), and Tupandactylus navigans (Frey et al., 2003b). ...
Article
Full-text available
A new species of boreopterid pterosaur from the new fossil locality, Heichengzi, Beipiao, western Liaoning, China allows a reassessment of the Boreopteridae. In this new analysis, three species, Boreopterus cuiae, Boreopterus giganticus n. sp., and Zhenyuanopterus longirostris, are included within the Boreopteridae united by the autopomorphic occurrence of two main tooth morphologies, an equal length of the tibia and femur, and weak feet. Other taxa previously placed within the Boreopteridae are not in a monophyletic group with the former three species. Boreopterus has fewer teeth and a shorter tooth row than that in Zhenyuanopterus. This new Boreopterus species has a large size, a piriform orbit, an extensively fenestrated lacrimal, and a posteriorly directed lacrimal process, that differs from Boreopterus cuiae.
... Executive producer Steven Spielberg evidently agreed that the teeth were indeed essential selling points. The genus name of the new species Ludodactylus sibbicki Frey et al., 2003 is a combination of the Greek ludo (game, plaything) and Latin dactyl (finger), to honor the fact that this creature was reconstructed by plastic model artists long before paleontologists ever discovered its fossils. Professional paleontologists used to complain about the hybrid models as "highly inaccurate," but have been forced to swallow their criticisms in the face of Ludodactylus. ...
Book
Full-text available
This book represents Matt Lafreniere's student internship project for Holyoke Catholic High School, undertaken with Professor Mark McMenamin at Mount Holyoke College in summer 2012. The subject matter lends itself to both K-12 education and to scientifically fruitful consideration of a wide variety of ancient organisms. It may truly be said that a good dinosaur model represents a summary of what we think we know about any particular ancient creature at the time the model was made. Each realistically sculpted and painted model furthermore provides an artist's interpretation of the contours and coloration of the animal, and as such the model represents a potentially important hypothesis concerning what the animal actually looked like. And as we will see in the case of Ludodactylus, such models can on occasion anticipate future results of ongoing field research, excavation and fossil discovery.
... Throat tissues are preserved in the ctenochasmatoid Pterodactylus antiquus and the rhamphorhynchid Rhamphorhynchus muensteri (e.g. Frey et al. 2003a), and are inferred for the ornithocheiroids Ludodactylus sibbicki and Pteranodon sp. from the preservation of ingested material between their mandibular rami (Frey et al. 2003b;Bennett 2001). The exact nature of these tissues (e.g. ...
Article
Full-text available
The lifestyles of all pterosaurs are contentious, but those of the pterodactyloid clade Azhdarchidae are particularly debated. A 2008 review of the functional morphology of azhdarchid pterosaurs concluded that they were probably terrestrial foragers, as evidenced by their long limbs, generalised skull construction, the arthrological limitations of their cervical series, trackway data indicating terrestrial proficiency, a strong continental skew in the depositional context of their fossils, and several additional lines of corroborating evidence. This hypothesis was recently challenged on three counts: 1) azhdarchid fossils routinely occur in aquatic deposits; 2) terrestrially-foraging pterosaurs were highly vulnerable to predation and 3), aerial ‘water trawling’, where the mandible is pulled though water to catch food in a distended throat pouch, is a more likely foraging strategy. Pelican-like jaw mechanics were suggested for azhdarchids because of the asymmetrical jaw joints in these pterosaurs, which permit lateral deflection of the mandibular rami during jaw extension. We evaluate these three claims and conclude that all are flawed. The frequent occurrence of azhdarchid fossils in aquatic sedimentary systems is not significant with regard to ecology or behaviour, since these provide the overwhelming mechanism for the preservation of all fossil terrestrial animals. Likely pterosaur takeoff abilities and the ubiquitous nature of modern, terrestrially-foraging birds indicate that predation risks on ground-foraging pterosaurs are probably overstated. The kinematics of pterosaur jaws are entirely different to those of pelicans, which are highly specialised compared to those of all other tetrapods, and there are no indications from azhdarchid jaw anatomy that azhdarchids indulged in pelican-like foraging behaviour. The estimated amount of jaw expansion present in azhdarchids was minimal compared to that of pelicans, even when the asymmetrical jaw joints of azhdarchids are taken into account. Moreover, the widespread occurrence of asymmetrical jaw joints in other reptiles demonstrates that they are not related to any specific feeding habits. We conclude that terrestrial foraging remains the most parsimonious habit for azhdarchid
Article
Full-text available
In the palaeogeographical context of the opening of the South Atlantic Ocean at the end of the Early Cretaceous, we document here the first microvertebrate fauna recorded from the Aptian deposits of Gadoufaoua, Niger. A systematic study of the fauna has resulted in a significant expansion of the existing faunal list and increase our knowledge of the palaeobiodiversity of the Gadoufaoua fossil site. Some taxa were previously recorded (lungfishes, crocodilians, chelonians, etc.), but several new taxa are described here for the first time, including the first occurrence of a stem-boreosphenidan mammal in Africa for this time-interval. In addition, chondrichthyans, pterosaurs and lissamphibians are documented. The analysis of taxonomic diversity and preservation of the fossils confirms a floodplain depositional environment (more precisely channel-type), in a higher velocity flow regime than previously thought based on studies of the macrofauna. To confirm the depositional environment inferred by the state of preservation of the fossils, a preliminary comparison with the fauna of similar age from the Santana Formation (Brazil) has been undertaken. The hypothesis of a communication between Africa and South America during the Aptian, as had already been demonstrated based on studies of the macrofauna, is strengthened.
Article
Full-text available
Diagnostic characters from 227 pterosaur species were listed, separated into cranial or post-cranial elements and counted. From 21 post-cranial and 23 cranial elements, most diagnostic characters were related to phalanges (15%) and rostrum. Post-cranial characters comprise 44.23%, and cranial characters 55.77% of all characters used in pterosaur diagnoses. The highest correlation between diagnostic features occurs between the coracoid and the scapula. 25.11% and 28.63% of sampled taxa were diagnosed with 3-4 and 5-6 characters, respectively. The mean number of 6.79 characters was found in specimens with both cranial and post-cranial elements, and 4.86 and 4.17 in those with just cranial or post-cranial elements, respectively. 31 from 227 species (13.7%) were erected based on single elements, which are most frequently complete or partial mandibles (n=18). We estimate that 23.4% of the total pterosaur genera are currently known, with 90% of this diversity to be unveiled up to 2145. As the requirements of broad and cautious revision of genus/group must be undertaken, and some deposits will provide mostly fragmented and incomplete material, the assignment of fairly incomplete specimens to the most inclusive taxonomic level is feasible. Tracing this scenario can guide future works on the description of new pterosaur taxa.
Thesis
Full-text available
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.
Article
The pterosaurs were the first vertebrates to evolve powered flight and they staged a major radiation in the Cretaceous. Cretaceous pterosaurs occupied many of the niches occupied today by birds, including aerial insect hawkers, piscivores, and filter feeders. The diversity of this radiation remains poorly known due to the uneven stratigraphic and geographic distribution of pterosaur fossils. Much of what is known about pterosaur diversity comes from a handful of Lagerstätten, representing primarily lacustrine, lagoonal, and marine palaeoenvironments, primarily in Laurasia. These sites may fail to capture pterosaur diversity in other habitats and regions, such as Gondwana. Here, we describe a unique small, long-beaked pterosaur from fluvial mid-Cretaceous (?Albian–Cenomanian) strata of Morocco, North Africa, with adaptations for sediment probing. The upper and lower jaws form a hyperelongate dorsoventrally flattened beak, with thickened bony walls. The morphology most closely resembles that of probing birds such as kiwis, ibises, and curlews that probe in mud or earth for invertebrates. The affinities of the new pterosaur are unclear. It likely represents an azhdarchoid, but does not clearly fit within any known azhdarchoid clade. The new pterosaur adds to the remarkable diversity of pterosaurs known from the mid-Cretaceous, and suggests that pterosaur diversity remains under sampled.
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.
Article
As more new fossils are discovered, a great number of extraordinary specimens have greatly enhanced our understanding of bone features in dinosaurs. The fracture patterns of dinosaur bones are distinctive and easily identifiable, and thus have received increasing attention. Some pioneering modern medical studies have categorised two types of bone fractures, including traumatic (a broken bone caused by an injury) and pathologic (a broken bone caused by a disease) fractures. Here, we investigated two fossil specimens collected from Yangchuanosaurus hepingensis and Gigantospinosaurus sichuanensis, respectively, which were both uncovered in the Sichuan Basin, southwestern China. These specimens were subjected to osteological examination, CT scanning and spectral data analysis. Thus, multiple lines of evidence indicate that the bone fracture in the two dinosaurs is attributed to different factors.
Article
Full-text available
The pterosaur is the first known vertebrate clade to achieve powered flight. Its hyoid apparatus shows a simplification similar to that of birds, although samples of the apparatus are rare, limiting the ability to make an accurate determination. In this study we reveal a new pterosaur specimen, including the first definite basihyal. Through the comparison of pterosaur hyoids, a trend has been discovered for the shortened hyoid relative to the length of the skull, indicating a diminished role of lingual retraction during the evolution of the pterosaur. The new material, possibly from a gallodactylid Gladocephaloideus , represents one of the least effective lingual retractions in all pterosaurs. Based on the structure of an elongated ceratobranchial and retroarticular process on mandibles, the function of the Y-shaped istiodactylid tongue bone is similar to those of scavenger crows rather than chameleons, which is consistent with the interpretation of the scavenging behavior of this taxon. More fossil samples are needed for further study on the function of other pterosaur hyoids.
Article
Full-text available
The pterosaur record from the Iberian Peninsula is mostly scarce and undefined, but in the last few years some new taxa have been described from different Lower Cretaceous sites of Spain. Here we describe a new genus and species of toothed pterodactyloid pterosaur from the Barremian of the Iberian Peninsula, Iberodactylus andreui gen. et sp. nov., that shows a close and rather unexpected relationship with Hamipterus tianshanensis from China. A review of the phylogenetic relationships of the Anhangueria reveals a new family of pterodactyloid pterosaurs, the Hamipteridae fam. nov. being recovered as sister-group of the Anhangueridae. This latter clade can be in turn divided into the new clades Anhanguerinae and Coloborhynchinae. The close relationships of Iberodactylus and Hamipterus shows an interesting palaeobiogeographical correlation between the Chinese and Iberian pterosaur faunas during the Barremian (Lower Cretaceous). The discovery of Iberodactylus strongly suggests that the clade Anhangueria has clear ancestral ties in eastern Laurasia.
Article
Full-text available
Background In the Early Cretaceous Jehol Biota, the toothless pterosaurs flourished with the chaoyangopterids and tapejarids playing a key role in understanding the early diversity and evolution of the Azhdarchoidea. Unlike the more diverse tapejarids, the rarer chaoyangopterids are characterized by a long and low rostrum, supporting a close relationship with the huge azhdarchids. Unfortunately, our knowledge is still limited in the osteology, paleoecology, and taxonomy of the Chaoyangopteridae. As one of the best preserved skeletons, the type and only specimen of Jidapterus edentus provides an opportunity to understand the morphology and paleoecology of the chaoyangopterids. Results Our study of the osteology of Jidapterus edentus reveals valuable information about the morphology of the Chaoyangopteridae such as a rostrum with a curved dorsal profile, high Rostral Index (RI), larger angle between the dorsal and postorbital processes of the jugal, sequentially shorter fourth to seventh cervical vertebrae, sternum with a plate wider than long, contact of the metacarpal I with the distal syncarpal, pneumatic foramen on first wing phalanx, hatchet-like postacetabular process with unconstricted neck and small dorsal process, distinctly concave anterior margin of pubis, subrectangular pubic plate with nearly parallel anterior and posterior margins, longer proximal phalanges of pedal digits III and IV, as well as reduced and less curved pedal unguals. These features further support the validity of Jidapterus edentus as a distinct species and the close relationship of the chaoyangopterids with the azhdarchids. Paleoecologically, the chaoyangopterids are probably like the azhdarchids, more terrestrial than the contemporaneous and putatively arboreal tapejarids, which may have been limited to the forest-dominated ecosystem of the Jehol Biota. Discussion The osteology of Jidapterus edentus further supports the close relationship of the Chaoyangopteridae with the Azhdarchidae in sharing a high RI value and reduced and mildly-curved pedal unguals, and it also implies a possible paleoecological similarity in their terrestrial capability. Combined with the putatively arboreal and herbivorous tapejarids, this distinct lifestyle of the chaoyangopterids provides new insights into the diversity of pterosaurs in the ecosystem of the Jehol Biota.
Article
Full-text available
The fossil record of Australian pterosaurs is sparse, consisting of only a small number of isolated and fragmentary remains from the Cretaceous of Queensland, Western Australia and Victoria. Here, we describe two isolated pterosaur teeth from the Lower Cretaceous (middle Albian) Griman Creek Formation at Lightning Ridge (New South Wales) and identify them as indeterminate members of the pterodactyloid clade Anhangueria. This represents the first formal description of pterosaur material from New South Wales. The presence of one or more anhanguerian pterosaurs at Lightning Ridge correlates with the presence of ‘ornithocheirid’ and Anhanguera-like pterosaurs from the contemporaneous Toolebuc Formation of central Queensland and the global distribution attained by ornithocheiroids during the Early Cretaceous. The morphology of the teeth and their presence in the estuarine- and lacustrine-influenced Griman Creek Formation is likely indicative of similar life habits of the tooth bearer to other members of
Article
Understanding the ecological roles of pterosaurs is a challenging pursuit, but one aided by a growing body of fossil evidence for their dietary preferences and roles as food sources for other species. Pterosaur foraging behaviour is represented by preserved gut content, stomach regurgitates, coprolites and feeding traces. Pterosaurs being eaten by other species are recorded by tooth marks and teeth embedded in their fossil bones, consumer gut content and regurgitate, and their preservation entangled with predatory animals. This palaeoecological record has improved in recent years, but remains highly selective. The Jurassic rhamphorhynchid Rhamphorhynchus, Cretaceous ornithocheiroid Pteranodon and azhdarchid pterosaurs currently have the most substantial palaeoecological records. The food species and consumers of these taxa conform to lifestyle predictions for these groups. Rhamphorhynchus and Pteranodon ate and were eaten by aquatic species, matching expectations of these animals as sea-going, perhaps partly aquatic species. Possible azhdarchid pterosaur foraging traces alongside pterosaur tracks, and evidence that these animals were eaten by dinosaurs and Crocodyliformes, are consistent with hypotheses that azhdarchids foraged and lived in terrestrial settings. Fossil evidence of pterosaur palaeoecology remains rare: researchers are strongly encouraged to put specimens showing details of dietary preferences, foraging strategies or interactions with other animals on record.
Article
A pterosaur from the Lower Cretaceous (Lower Aptian) Leza Formation of La Rioja (Spain) is assigned to a new genus and species of Pterodactyloidea, Prejanopterus curvirostra n. gen. n. sp., based on the holotype FA 112 (rostrum) and associated craneal and postcraneal elements. The rostrum is elongated, sharp and curved to the left and has a pronounced midline palatal groove. The 1-5 and 10-11 pairs of dental alveoli are significantly smaller than pairs 6-9 and 12-20. The teeth of the maxilla are short and labiolingually compressed. The mandible is long and thin, and not curved. The mandibular symphysis is long, occounting for 35% of the length of the lower jaw. It has no crest either on the bill or the mandible. The new taxon represents the first genus and species of pterosaur from the Lower Cretaceous in Spain.
Article
Full-text available
Pterosaurs have been found in eight provinces in China, and their geological ages are from the Middle Jurassic to the Late Cretaceous. We have analyzed the U-Pb zircon ages of two pterosaur localities, Linglongta in Liaoning and Tangshang Formation in Zhejiang, whose ages were disputable. The youngest age of detrital zircons is 150 Ma, which suggested that the age of the Linglongta Pterosaur Fauna was younger than 150 Ma. Two different ages of tuffs from Tangshang Formation in different basins in Zhejiang were obtained: one from Tiantai basin, underlying the dinosaur-egg bearing strata, is 113 Ma and one from Shangpan locality of Linhai is 90 Ma; the pterosaurs were found in the latter basin. These results suggested that Zhejiangopterus was in the Late Cretaceous, and the Tangshang Formation in Tiantai and Xiaoxiong basins were deposited in different times. On the basis of these new ages of two localities, we preliminarily summarized the geochronology and stratigraphic sequence of pterosaur fossil-bearing beds in China, and the pterosaurs from the Early Cretaceous were in the main position.
Article
The feeding behavior of pterosaurs was varied in different groups. Judging by the tooth morphologies and fossilized stomach contents of pterosaurs, the feeding habits included fish-eating, insect-eating, filtering small aquatic organisms, eating shelled crabs and snails, and fruit-eating. Because of the need for survival, they occupied different ecological niches, which determined their different food sources. Herein described is an almost complete, well-preserved hyoid apparatus of Liaoxipterus brachyognathus in comparison with the hyoid apparatus of the modern lizard Chameleon. The long processus lingualis (processus entoglossus) is similar to that of the modern lizard Chameleon, which captures prey by tongue, implying that Liaoxipterus might share a similar lingual feeding behavior. This phenomenon, plus its special tooth morphology, further suggests that it was an insect-eating rather than fish-eating pterosaur.
Article
Pterosaurs first appeared in the Late Triassic and persisted until the terminal Cretaceous: they achieved a global distribution during the Mesozoic. Here, we attempt to provide the first comprehensive summary of pterosaur distribution through time and space, including information on the taxonomie composition of pterosaur faunas and the lithostratigraphic units in which they occur. We hope that this compilation will be used as a primary research tool, permitting more detailed and rigorous analyses of pterosaur diversity and palaeobiogeography than have been possible to date.
Chapter
Full-text available
Paleoart is a widely used term in the paleontological world. This word can be found in different media, used in relation to different kinds of artistic manifestations of paleontological theme. Paleoart has a long history and has helped paleontology to became one of the most popular sciences. The production of this artwork means that paleoartists must have high skills in both disciplines, arts and paleontology. Due to its scientist, artistic and cultural significance, paleoart and paleoartists must be recognized and valued in both paleontology and fine arts. In this situation paleoart needs a clear definition that distinguishes it from other artistic representations. Taking this into account, we have consulted to a series of paleontologists, and this survey has allowed us to outline such a definition. All the original artistic manifestations that pretend to reconstruct o depict prehistoric life according to the current knowledge and scientific evidence at the moment of creating the artwork can be considered paleoart.
Article
Full-text available
The Crato Formation paleoflora is one of the few equatorial floras of the Early Cretaceous. It is diverse, with many angiosperms, especially representatives of the clades magnoliids, monocotyledons and eudicots, which confirms the assumption that angiosperm diversity during the last part of the Early Cretaceous was reasonably high. The morphology of a new fossil monocot is studied and compared to all other Smilacaceae genus, especially in the venation. Cratosmilax jacksoni gen. et sp. nov. can be related to the Smilacaceae family, becoming the oldest record of the family so far. Cratosmilax jacksoni is a single mesophilic leaf with entire margins, ovate shape, with acute apex and base, four venation orders and main acrodromous veins. It is the first terrestrial monocot described for the Crato Formation, monocots were previously described for the same formation, and are considered aquatics. Cratosmilax jacksoni is the first fossil record of Smilacaceae in Brazil, and the oldest record of this family.
Article
Full-text available
The Cambridge Greensand, a remanié deposit that crops out in Cambridgeshire, eastern England, has yielded numerous, though fragmentary, late Early Cretaceous (Albian) vertebrate fossils including more than 2000 isolated pterosaur bones. So far, 32 species of pterosaur have been proposed in connection with the Cambridge Greensand material, but there has been and continues to be considerable confusion concerning the validity of these taxa, their relationships to each other and to other pterosaurs, and the material upon which they were established. A comprehensive systematic revision identified eleven valid species distributed among three families: the Ornithocheiridae (Ornithocheirus simus and possibly a second, as yet unnamed species of Ornithocheirus, Coloborhynchus capito, Coloborhynchus sedgwickii, Anhanguera cuvieri, and Anhanguera fittoni); the Lonchodectidae (Lonchodectes compressirostris, Lonchodectes machaerorhynchus, Lonchodectes microdon and Lonchodectes platystomus); and a species of edentulous pterosaur (Ornithostoma sedgwicki) that may represent the earliest record for the Pteranodontidae. It is possible that some of the taxa currently recognised represent sexual dimorphs (Coloborhynchus capito and Coloborhynchus sedgwickii, Lonchodectes compressirostris and Lonchodectes machaerorhynchus), or disjunct populations of a single species (Ornithocheirus simus and Ornithocheirus sp., Lonchodectes compressirostris and Lonchodectes microdon) and that there may be as few as seven valid species, but the Cambridge Greensand pterosaurs are too poorly known to demonstrate this at present. The Cambridge Greensand pterosaur assemblage is similar to a slightly younger, but much smaller assemblage from the Lower Chalk of England and shares some elements, such as ornithocheirids, in common with many other late Early and early Late Cretaceous assemblages. It is distinguished by the absence of tapejarids and the presence of Lonchodectes which, so far, is only known from the Cretaceous of England. The disparity in taxonomic composition is possibly related to ecological differentiation, and might also reflect some provincialism in late Early and early Late Cretaceous pterosaur faunas. Der Cambridge Greensand, eine in Ostengland aufgeschlossene Remanié-Ablagerung, hat zahlreiche Wirbeltiere aus der oberen Unterkreide (Alb) geliefert. Darunter fanden sich mehr als 2000 isolierte Pterosaurierknochen. Insgesamt wurden aus dem Greensand bis zu 32 Flugsauriertaxa beschrieben, was zu einer beträchtlichen taxonomischen und nomenklatorischen Verwirrung geführt hat, die bis heute andauert. Eine vollständige Revision erkennt 11 Arten aus drei Familien an: (1) die Ornithocheiridae (Ornithocheirus simus und vielleicht eine zweite, bislang unbenannte Art von Ornithocheirus, sowie Coloborhynchus capito, Coloborhynchus sedgwickii, Anhanguera cuvieri und Anhanguera fittoni); (2) die Lonchodectidae (Lonchodectes compressirostris, Lonchodectes machaerorhynchus, Lonchodectes microdon und Lonchodectes platystomus); und schließlich einen zahnlosen Flugsaurier (Ornithostoma sedgwicki). der zu keiner der vorgenannten Familien gehört und sich als stratigraphisch ältester Nachweis der Pteranodontidae erweisen könnte. Es ist nicht auszuschließen, dass einige der gegenwärtig erkannten Taxa eher einen ausgeprägten Sexualdimorphismus illustrieren denn taxonomisch distinkte Arten darstellen (Coloborhynchus capito und Coloborhynchus sedgwickii, Lonchodectes compressirostris und Lonchodectes machaerorhynchus) oder sogar lediglich Endpunkte einer intraspezifisch variablen Population (Ornithocheirus simus und Ornithocheirus sp., Lonchodectes compressirostris und Lonchodectes microdon). In dieser strengeren Fassung bestünden nur sieben gültige Arten, doch leider sind die Flugsaurier des Cambridge Greensand zu schlecht bekannt, um diese Fragen zu beantworten. Die Flugsaurierfauna des Cambridge Greensand ähnelt jüngeren kreidezeitlichen Faunen aus dem Lower Chalk von England. Weiter-hin enthält sie Faunenelemente, wie etwa Ornithocheiriden. die auch für zahlreiche andere Faunen der hohen Unterkreide und tiefen Oberkreide charakteristisch sind. Das Fehlen von Tapejariden und das Auftreten des anscheinend endemischen Lonchodectes sind weitere Kennzeichen des Cambridge Greensand. Die Zusammensetzung der Pterosaurierfaunen folgte offenbar ökologischen Differenzierungen und illustriert einen gewissen Provinzialismus an der Grenze Unter-Oberkreide. doi:10.1002/mmng.20010040112
Article
Full-text available
An almost symmetrical, isolated feather from the Crato Formation is associated with numerous spherical, hollow structures approximately 75 microns diameter which are interpreted as the eggs of an avian ectoparasite, probably a mite. The feather shape resembles the caudal feathers of modern birds and Archaeopteryx, but it appears to have only simple barbules. This specimen may represent the oldest occurrence of ectoparasitism on feathered maniraptorans.
Article
Full-text available
A lizard from the Early Cretaceous Crato Formation of Brazil is the first from this Formation and one of the earliest recorded lizards from South America. The specimen represents a very young individual, probably recently hatched. The lower jaws are characterised by an elongated dorsal ramus of the dentary which replaces the surangular along much of the posterodorsal margin ofthe mandible. This unusual feature distinguishes the lizard from all those previously described. It is here designated Olindalacerta brasiliensis n. g. n. sp. Zusammenfassung: Ein Eidechsen-Skelett aus dem Araripe-Becken ist der erste a us der Crato-Formation und einer der friihesten Belege fur Eidechsen in Siidamerika. Das Exemplar ist ein Jungtier, vermutlich irn Stadium kurz nach dem Ausschliipfen. Die Kiefer tragen einen verlangerten dorsalen Ast des Dentale, der auf eine weite Strecke des posterodorsalen Unterkiefer-Randes das Supraangulare ersetzt. Mit diesem unterscheidenden Merkmal wird das neue Taxon Olindalacerta brasiliensis n. g. n. sp. begriindet.
Article
Full-text available
A partial rostrum of a new species of scaphognathine pterosaur, distinguished by a thin median crest along its dorsal margin and a deep embayment of the dental margin, is the first identifiable cranial fragment of a pterosaur from the Upper Jurassic Morrison Formation of western North America. By contrast with pterodactyloids, cranial crests are rare in "rhamphorhynchoids" and this is the first record of such a structure. The new material provides fresh insights into the taxonomic diversity of Late Jurassic North American pterosaurs. Based on the ratio of the skull and skeleton of Scpahognathus, the fragment represents an individual with an estimated wing span of 2.5 m. Consequently, this is one of the largest "rhamphorhynchoids" found so far. A mandible fragment from the same quarry has closely spaced alveoli, therefore cannot be referred to the rostrum. Its large size indicates another large "rhamphorhynchoid" in the Morrison Formation.
Article
Full-text available
The partially articulated, distal portion of a left wing finger of a pterosaur from the Crato Formation of northeast Brazil has a T-shaped cross-section to the second and third phalanges. This cross-sectional shape is one of several characters diagnostic of the pterodactyloid pterosaur family Azhdarchidae (Unwin & Lu 1997). Until now, this family of pterosaurs was known exclusively from the Late Cretaceous. The specimen described here may be the first recorded azhdarchid from the southern hemisphere and the earliest recorded member of the family.
Article
A semi-plume feather from the Nova Olinda Member of the Crato Formation (Aptian, Lower Cretaceous) of north-east Brazil is only the second feather reported from this remarkable lagerstatte. The new specimen is presumed to be preserved as an organic film in organic-rich, laminated limestones, and is from an unknown bird probably in the size range 150 to 300 mm. -Authors
Article
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.
Article
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
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
The first Mesozoic japygid (Hexapoda: Diplura), Ferrojapyx vivax gen. et sp. nov., is described from the Lower Cretaceous (Aptian) Crato Formation of north-east Brazil. There are only two previously described occurrences of japygids in the fossil record: in Miocene or Pliocene onyx marble from Arizona and from the Carboniferous (Westphalian D) Francis Creek Shale of Mazon Creek, Illinois.
Article
We examine the preservation, autecology and morphological variation for several characters of the Cretaceous gonorynchiform fish Dastilbe from the Lower Cretaceous of Brazil and Africa. More than 83 specimens were examined. We test species validity using characters of the caudal endoskeleton and meristic counts of fin-rays vs length. Evidence provided by fossilized soft tissues and slabs containing large individuals ‘freeze framed’ in the process of swallowing smaller prey meals, show that Dastilbe was predatory, at least as adults, as well as cannibalistic. Dastilbe was probably an anadromous fish tolerant of hypersalinity and in Araripe was subjected to frequent mass mortality events. Observations of the otic region indicate that the lagenar statolith is consistently larger than the saccular statolith, hence revealing a primitive actinopterygian condition. For the first time, a lagenar statolith from Dastilbe has been cleaved to expose putative annuli-like ridges. Our results clearly show that there is a wide degree of morphological plasticity of the endoskeleton coupled with wide meristic variation, and as such, overall length, fin-ray count and even absence or presence of caudal diastema are not suitable criteria for species recognition in Dastilbe. New specimens from the Crato Formation (Aptian) and statistical tests suggest rejection of all species of Dastilbe erected subsequent to Jordan (1910). All Brazilian specimens of Lower Cretaceous Dastilbe can be assigned to the single species D. crandalli Jordan. The African D. batai Gayet is also placed within D. crandalli
Article
The anterior tips of associated upper and lower jaws of a pterosaur from the Lower Cretaceous of Brazil are described and assigned to the taxonColoborhynchus in the family Ornithocheiridae. It is characterized by the shape and position of the sagittal crest on the upper and lower jaw, the arrangement and length of the teeth and the spoon-like lateral expansion of the anterior parts of the jaws. It closely resemblesColoborhynchus wadleighi from North America andColoborhynchus clavirostris from England. Diagnostic anatomical characteristics permit a revision of the genusTropeognathus, which is shown here to be a junior synonym of other described taxa.Tropeognathus mesembrinus is referred toCriorhynchus andT. robustus toColoboryhnchus. Consistent anatomical features enable the new jaw fragment to be assigned toColoborhynchus robustus. Das anteriore Ober- und Unterkieferfragment eines Pterosauriers aus der Unteren Kreide von Brasilien wird beschrieben und zur GattungColoborhynchus in die Familie Ornithocheiridae gestellt. Die charakteristischen antomischen Eigenschaften sind der sagitale Kamm am Ober- und Unterkiefer, die Anordnung und Länge der Zähne und die löffeiförmige laterale Verbreiterung der anterioren Kieferenden. Das Stück ähnelt starkColoborhynchus wadleighi aus Nordamerika undColoborhynchus clavirostris aus England. Die diagnostischen morphologischen Eigenschaften des beschriebenen Stückes ermöglichen eine Revision der GattungTropeognathus, die synonym mit anderen Gattungen ist.Tropeognathus mesembrinus wird in die GattungCriorhynchus, T. robustus in die GattungColoborhynchus gestellt. Aufgrund der starken anatomischen Übereinstimmung kann das beschriebene Fragment alsColoborhynchus robustus identifiziert werden.
Taxonomy and systematics of the Late Cretaceous pterosaur Pteranodon (Pterosauria, Pterodactyloidea) Natural History Museum
  • S C Benneat
BENNEaT, S. C. 1994. Taxonomy and systematics of the Late Cretaceous pterosaur Pteranodon (Pterosauria, Pterodactyloidea). Natural History Museum, University of Kansas Occasional Papers, 169, 1-70.
The osteology and functional morphology of the Late Cretaceous pterosaur Pteranodon. Part 1. General description of the osteol-ogy
  • S C Bennetr
BENNETr, S. C. 2001. The osteology and functional morphology of the Late Cretaceous pterosaur Pteranodon. Part 1. General description of the osteol-ogy. Palaeontographica A, 260, 1-112.
Un novo exam-plar de Anhanguera blittersdorffi (Reptilia, Pterosauria) da formac~o Santana, Cret~iceo Inferior do Nordeste do Brasil
  • D A Cameos
  • A W A K~llner
CAMEOS, D. A. & K~LLNER, A. W. A. 1985. Un novo exam-plar de Anhanguera blittersdorffi (Reptilia, Pterosauria) da formac~o Santana, Cret~iceo Inferior do Nordeste do Brasil. Boletim de Resumos, 9th Congresso Brasileiro de Paleontologia, 13. CAMeoS, D. A. & KELLrCER, A. W. A. 1997. Short note on the first occurrence of Tapejaridae in the Crato Member (Aptian), Santana formation, Araripe Basin, northeast Brazil. Anais de Academia Brasileira de Ci~ncias, 69, 83-87.
Undetermined Santana frog Santana Fossils, An Illustrated Atlas. Tropical Fish Hobbyist Preservation of fish in the Cretaceous of Brazil
  • J G Maisey
  • D M Martill
MAISEY, J. G. 1991. Undetermined Santana frog. In: MAISEY, J. G. (ed.) Santana Fossils, An Illustrated Atlas. Tropical Fish Hobbyist, Neptune City, New Jersey, 459 pp. MARTILL, D. M. 1988. Preservation of fish in the Cretaceous of Brazil. Palaeontology, 31, 1-18.
Fossils of the Santana and Crato Formations, Brazil. Field Guides to Fossils, Palaeontological Association A feather with pos-sible ectoparasite eggs from the Crato Formation
  • D M Martill
  • Pp
  • D M Martill
  • P G Davis
MARTILL, D. M. 1993. Fossils of the Santana and Crato Formations, Brazil. Field Guides to Fossils, Palaeontological Association, vol. 5,159 pp. MARTILL, D. M. & DAVIS, P. G. 2001. A feather with pos-sible ectoparasite eggs from the Crato Formation (Lower Cretaceous, Aptian) of Brazil. Neues Jahrbuch fiir Geologie und Paliiontologie, Abhandlungen, 219, 241-259.
Stratigraphy Fossils of the Santana and Crato Formations, Brazil. Field Guides to Fossils
  • D M Martill
  • P Wilby
MARTILL, D. M. & WILBY, P. 1993. Stratigraphy. In: MARTILL, D. M. (ed.) Fossils of the Santana and Crato Formations, Brazil. Field Guides to Fossils, Palaeontological Association, vol. 5,159 pp. PONS, D., BERTHOU, P.-Y. & CAMPOS, D. A. 1990. Quelques
Undetermined Santana frog
  • J G Maisey
Un novo examplar de Anhanguera blittersdorffi (Reptilia, Pterosauria) da formacâo Santana, Cretáceo Inferior do Nordeste do Brasil Boletim de Resumos 1985 13 9th Congresso Brasileiro de Paleontologia
  • D A Campos
  • A W A Kellner
  • Campos D. A.
Fossils of the Santana and Crato Formations, Brazil, Field Guides to Fossils
  • D M Martill
Congresso Brasileiro de Geologia (Anais), Ocorrencia de uma mandibula de Pterosauria (Brasileodactylus araripensis nov. gen.; nov. sp.) na formaçáo Santana
  • A W A Kellner
The Early Cretaceous pterodactyloid pterosaur Coloborhynchus from North America
  • Y N Lee
  • Lee Y. N.
The Ornithosauria: An Elementary Study of the Bones of Pterodactyles, Made from Fossil Remains Found in the Cambridge Greensand, and Arranged in the Woodwardian Museum of the University of Cambridge 1870 Cambridge Deighton Bell & Co
  • H G Seeley
  • Seeley H. G.
Dragons of the Air: An Account of Extinct Flying Reptiles
  • H G Seeley
  • Seeley H. G.
Taxonomy and systematics of the Late Cretaceous pterosaur Pteranodon (Pterosauria, Pterodactyloidea) 1994 169 1 70 Natural History Museum
  • S C Bennett
  • Bennett S. C.
Crato’ et ‘Ipubi’ (Aptien supérieur à Albien inférieur-moyen, Bassin d’Araripe, NE du Brésil) Géologie de l’Afrique et de l’Atalantique Sud (Actes Colloques Angers
  • D Pons
  • P.-Y Berthou
  • D A Campos
  • S Jardine
  • L Klasz
  • De
  • J-P Debaney
  • Pons D.
Ocorrencia de uma mandibula de Pterosauria (Brasileodactylus araripensis nov. gen.; nov. sp
  • A W A Kellner
  • Kellner A. W. A.
Fossils of the Santana and Crato Formations, Brazil 1993 5 159 pp Field Guides to Fossils
  • D M Martill
  • Martill D. M.
A new feather from the Lower Cretaceous of Brazil
  • D M Martill
  • B J Filguiera
  • Martill D. M.