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Record claims of pterosaur wingspans and equivalent standing heights compared to ( a ) a 3 m span Andean condor ( Vultur gryphus ) and ( b ) a 3 m span wandering albatross ( Diomedea exulans ). ( c ) Marsh’s (1876 a ) 7.6 m span Pteranodon longiceps . ( d ) Stoyanow’s (16 November 1936, Time Magazine ) apocryphal 10 m span Jurassic pterosaur. ( e ) Harksen’s (1966 ) 9.1 m span Pteranodon sternbergi . ( f ) Lawson’s (1975) 11 m span Quetzalcoatlus northropi . ( g ) The Buffetaut et al. (2002) 12 m span Hatzegopteryx thambema . ( h ) The erroneously reported BA Festival of Science 20 m span pterosaur. Humans used for scale are 1.75 m tall.
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The immense size of many pterosaurs is now well known to academics and laymen alike, but truly enormous forms with wingspans more than twice those of the largest modern birds were not discovered until 83 years after the first pterosaur fossils were found. These remains were discovered in an expedition to the Cretaceous chalk deposits of Kansas led...
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... with wingspans of 7.6 m ( Fig. 2c) and reallocate a previously named species of Pterano- don, P. gracilis, to a new genus of Niobrara ptero- saur, Nyctosaurus (Marsh 1876b; note that Marsh (1881) renamed this genus Nyctodactylus following presumption that his first name was preoccupied; this was shown to be erroneous by Williston 1903). His description of this 'eight to ...
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... this flexion into his wingspan estimates, making his accuracy against other Ptera- nodon size estimates difficult to fathom. Larger pterosaurs were reported in 1966 when an almost complete skull of a new Pteranodon species, Ptera- nodon sternbergi, was described and suggested to belong to an individual spanning 30 ft (9.1 m) across the wings (Fig. 2e) (Harksen ...
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... pterodactyloids (e.g. Hooley 1913;Gilmore 1928;Swinton 1948;Young 1964;Miller 1971). A potential record of a giant pterosaur was mentioned in a 1936 Time article (entitled 'Diggers' published 16 November) in which T. A. Stoyanow was reported to have discovered an enor- mous pterosaur in Jurassic deposits of Arizona. With a reported 10 m wingspan (Fig. 2d), this find would have been significant in not only being larger than Pteranodon but also in being three being times larger than any Jurassic pterosaur known, even today (see Carpenter et al. 2003). The find, however, was never documented beyond the Time article and was never followed up by other pterosaur workers. This lull in ...
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... than 15 or 20 m span estimates. Later discoveries of complete skeletons from smaller but closely related forms such as Zhejiangopterus (Cai & Wei 1994) add further confidence to the lower wingspan estimate of Quetzalcoatlus northropi. These estimates suggest that Quetzalcoatlus northropi had a wingspan almost 40% larger than that of Pteranodon (Fig. 2f), and it remains one of the largest known flying ...
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... If Hatzegopteryx has a skull like those of other azhdarchids, the estimated 2.5 m length would represent the jaws alone, granting it a larger gape than even the biggest theropod dinosaurs (see Dal Sasso et al. 2005). The Hatzegopteryx humerus (Fig. 3c) is also more robust than that of Quetzalcoatlus, suggesting it had a minimum wing- span of 12 m (Fig. 2g) and, when standing, a shoulder height of 3 m (Fig. ...
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... 10 m span Quetzalcoa- tlus, evidence of pterosaurs of equal or larger pro- portions have been reported in relatively quick succession (e.g. Padian 1984;Martill et al. 1996;Buffetaut et al. 1997Buffetaut et al. , 2002Company et al. 2001;Hwang et al. 2002). Even these giants, however, were dwarfed by the claim of a 20 m span pterosaur made in 2005 (Fig. 2h). Tales of enormous pterosaur footprints in Mexico and a huge wing bone from Israel were revealed in a press conference at the 2005 British Association Festival of Science prior to any formal publication of either find, and an excited media quickly widely reported this announcement in news- papers, magazines and numerous websites around ...
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... Arambourigiania and Hatzegopteryx were not represented as they have WS similar to Quetzalcoatlus (sensu Frey and Martill, 1996;Buffetaut et al., 2002;Witton and Habib, 2010;Naish and Witton, 2017). Redrawn silhouettes from Witton, 2010;Witton and Habib, 2010. Human silhouette and scale bar 180 cm. and Ornithocheiroidea) and, in the less inclusive clades within: Tapejaroidea (i.e. ...
The largest pterosaur discovered in South America and one of the largest flying vertebrates in the world is described. Thanatosdrakon amaru gen. et sp. nov. is a new azhdarchid found in the upper-most levels of the Plottier Formation (upper Coniacian–lower Santonian, Neuquén Basin), Mendoza, Argentina. Two specimens were identified, the holotype (UNCUYO-LD 307) and the paratype, with an estimated wingspan of ∼ 7 m and ∼ 9 m, respectively. The cladistic analysis, based on 216 characters and 97 taxa, recovered Thanatosdrakon deeply nested within Azhdarchidae. Relationships within this clade are well resolved and the clade Quetzalcoatlinae presents two well-defined sister-groups: (Arambourgiania, Mistralazhdarcho, Aerotitan, Hatzegopteryx, Albadraco) + (Cryodrakon, Thanatosdrakon, Quetzalcoatlus ssp.). Thanatosdrakon is the oldest taxon of the clade Quetzalcoatlinae so far. It is represented by several well-preserved axial and appendicular bones in three dimensions. Some of these elements have never been described in giant azhdarchids (e.g. complete norarium, dorsosacral vertebrae and caudal vertebra) and this allows to expand the knowledge about the anatomy of this diverse group of pterosaurs. Finally, from a paleoecological point of view, Thanatosdrakon was found in floodplain deposits of ephemeral meandering systems indicating that this large flying species inhabited continental environments.
... Pteranodon is a large pterodactyloid pterosaur from the Late Cretaceous (Coniacian-Campanian) of North America with an estimated maximum wingspan of 7.25 m (Bennett, 2001). The genus was among the first pterosaurs reported from North America (Marsh, 1876)-see (Bennett, 2001;Witton, 2010 for context of its discovery) and has become one of the best known flying reptiles thanks to a representation of over 1,100 specimens-the highest sample size for any pterosaur genus. Although most specimens are incomplete and crushed, every component of its osteology is known and has been described in detail (Eaton, 1910;Bennett, 1991;Bennett, 1994;Bennett, 2001;Bennett, 2018;Bennett & Penkalski, 2018). ...
A cervical vertebra of the large, pelagic pterodactyloid pterosaur Pteranodon sp. from the Late Cretaceous Niobrara Formation of Kansas, USA is significant for its association with a tooth from the large lamniform shark, Cretoxyrhina mantelli . Though the tooth does not pierce the vertebral periosteum, the intimate association of the fossils—in which the tooth is wedged below the left prezygapophysis—suggests their preservation together was not mere chance, and the specimen is evidence of Cretoxyrhina biting Pteranodon . It is not possible to infer whether the bite reflects predatory or scavenging behaviour from the preserved material. There are several records of Pteranodon having been consumed by other fish, including other sharks (specifically, the anacoracid Squalicorax kaupi ), and multiple records of Cretoxyrhina biting other vertebrates of the Western Interior Seaway, but until now interactions between Cretoxyrhina and Pteranodon have remained elusive. The specimen increases the known interactions between large, pelagic, vertebrate carnivores of the Western Interior Seaway of North America during the Late Cretaceous, in addition to bolstering the relatively small fossil record representing pterosaurian interactions with other species.
... Perhaps surprisingly, based on the limited information provided by the fossil evidence, a number of elaborate theories have been proposed regarding the morphology and ecology of these giant pterosaurs, including tentative reconstructions of mass, height, posture, body plan, wingspan, terrestrial locomotion, flight capability (including take-off and landing), dietary habits and ecological role (for different approaches, interpretations and results, see, e.g. Chatterjee & Templin 2004;Witton 2007Witton , 2008Witton , 2010Witton , 2013aWitton & Naish 2008, 2015Sato et al. 2009;Habib 2010Habib , 2013Henderson 2010;Witton & Habib 2010;Martin & Palmer 2014;Naish & Witton 2017). Such circumstantial evidence, especially in reconstructing giants such as Arambourgiania philadelphiae (Arambourg 1959), Q. northropi, or Hatzegopteryx thambema Buffetaut, Grigorescu, Csiki 2002, was inferred from more complete fossils of smaller pterosaur taxa (e.g. the azhdarchids Zhejiangopterus linhaiensis Cai, Wei 1994emend. ...
... The above estimates suggest that the V alioara mandible belonged to an animal somewhat smaller than the type specimen of H. thambema from the same general area and stratigraphic level, with an estimated wingspan of 12 m (Buffetaut et al. 2002). Indeed, originally, H. thambema was thought to possess an extremely elongated, robust skull, up to 2.5 m in length (Buffetaut et al. 2002(Buffetaut et al. , 2003Witton & Naish 2008;Witton 2010Witton , 2013a. More realistic size estimates based on the holotype skull remains indicate a total skull length of not more than 1.6 m, with a proportionally short and robust neck. ...
We describe and interpret a posterior mandibular symphysis of a very large azhdarchid pterosaur. The specimen LPB (FGGUB) R.2347 exhibits a series of morphological characters present in both azhdarchid and tapejarid pterosaurs, suggesting a more basal position within the clade Azhdarchidae. This fossil was collected from Maastrichtian continental deposits near Vălioara in the Hațeg Basin, Romania, but cannot be confidently referred to the contemporaneous giant Hatzegopteryx thambema, also from Vălioara, due to the absence of overlapping skeletal elements. Remarkably, this mandibular symphysis shares a number of features the smaller azhdarchoid Bakonydraco galaczi from the Santonian of Hungary. Additional comparisons with previously described large‐sized azhdarchid mandibles indicate a certain degree of morphological and probably ecological disparity within the group. This specimen represents the largest pterosaur mandible ever found and provides insights into the anatomy of the enigmatic giant pterosaurs.
... Though it was considered a rather enigmatic vertebrate ichnotaxon for decades (Delair, 1963;Walkden and Oppé, 1969;MacFayden, 1970;Haubold, 1971;Delair and Lander, 1973;Sarjeant, 1974;Ensom, 1984Ensom, , 1986, Purbeckopus was finally identified as pterosaur track by M. G. Lockley in Wright et al. (1997;see also Ensom, 2002a;Lockley et al., 2008;Witton, 2010). Billon-Bruyat and Mazin (2003) questioned the affiliation of the tracks with pterosaurs, arguing that there is no clear manus-pes association identifiable in the available material. ...
A hypichnium of a manus imprint (preserved as plaster cast) indicates for the first time the presence of the large pterosaur ichnotaxon Purbeckopus cf. pentadactylus Delair, 1963 in the late Berriasian of northwest Germany. It is only the second record of Purbeckopus globally and the first pterosaur track from Germany.
It provides evidence of a very large pterosaur (wingspan c. 6 m) in this area and from this time period not yet represented by skeletal remains. When compared with the English type material, the specimen exhibits some differences that are related mostly to different properties of the substrate on which both were left. These include, in the German track, an impression of the metacarpophalangeal joint of the wing finger, normally not present in pterosaur tracks. Also interesting is the rather blunt termination of the deeply impressed digits I–III, indicating rather short and blunt claws, which seem more suitable for walking than for grasping or climbing. The specimens of Purbeckopus in England and Germany occur in different environments: the English locality was situated close to a brackish lagoon, while the German site belongs to a limnic-deltaic system at the margin of a large, freshwater lake.
... The first well known toothless pterosaur was Pteranodon Marsh, 1876 from the Late Cretaceous of North America. The first discovered specimen of this pterosaur was a wing metacarpal bone described as Pterodactylus oweni Marsh, 1871 and its toothless nature was not apparent until discovery of a more complete cranial material, upon which a new genus, "pterodactyl lacking teeth" (meaning of the word Pteranodon) was established in 1876 (see Witton 2010 for the history of discoveries). Seven years earlier, Seeley in the catalogue of fossils from the Sedgwick Museum in Cambridge mentioned three jaw fragments from the phosphorite mines within the Albian Cambridge Greensand near Cambridge, England, numbers 1-3 from cabinet J, tablet 16 (Seeley 1869: xvi). ...
Ornithostoma sedgwicki Seeley, 1891 from the Lower Cretaceous (Albian) Cambridge Greensand of England is represented by edentulous jaw fragments, posterior skull fragment with the supraoccipital crest, and by several postcranial bones attributed previously to Lonchodectes. Ornithostoma is referred to the Azhdarchoidea based on a combination of derived characters (teeth absent, middle cervicals moderately elongated, pneumatic foramen on anterior side of humerus, large pneumatic foramen on posterior side of femur at greater trochanter) and plesiomorphic characters (deltopectoral crest not warped, femoral neck to shaft angle less than 145°). The structure of the supraoc-cipital crest and humerus resembles those in Tapejara, but Ornithostoma differs from that taxon by a strong median ridge on the occiput presumably associated with a more elongated rostrum. At least three taxa of basal azhdarchoids were present in the British Lower Cretaceous. РЕЗЮМЕ Ornithostoma sedgwicki Seeley, 1891 из нижнемелового (альб) кембриджского зеленого песчаника Англии представлен фрагментами беззубых челюстей, фрагментом задней части черепа с затылочным гребнем, и некоторыми костями посткраниального скелета, которые раньше относили к Lonchodectes. Ornithostoma от-носится к Azhdarchoidea на основании комбинации продвинутых признаков (отсутствие зубов, умеренно удлиненные среднешейные позвонки, отверстие пневматизации на передней стороне плечевой кости, круп-ное отверстие пневматизации на задней стороне бедренной кости у большого трохантера) и плезиоморфных признаков (дельтопекторальный гребень не загнут, угол между шейкой и телом бедренной кости меньше 145°). Строение затылочного гребня и плечевой кости сходно с таковыми у Tapejara, но Ornithostoma отлича-ется от последнего таксона мощным медиальным гребнем на затылке, предположительно связанным с более удлиненной ростральной частью черепа. В нижнем мелу Великобритании существовало, по крайней мере, три таксона базальных аждархоидов.
... Some of the longest pterosaur necks, in proportion to body size, are seen in the giant species with wingspans of over (possibly well over) 10 m (Whitton, 2010). Many of these large pterosaurs, such as Quetzalcoatlus, are more commonly found in terrestrial sediments rather than the marine sediments more typical for smaller pterosaurs (Whitton & Naish, 2008). ...
... The ecology of these animals has been reconstructed as variously, 'typical' fishing pterosaurs (albeit very large ones), carrion feeders or ground predators-with Whitton & Naish (2008) preferring ground-hunting predators. Recently Henderson (2010) argued that their reconstructed body mass was such that they were probably secondarily flightless, although most authors have assumed they could fly (reviewed by Whitton, 2010). One obvious speculation about their feeding ecology is that they could have been carrion feeders-effectively Mesozoic vultures-specializing in the bodies of large dinosaurs. ...
There has been recent discussion about the evolutionary pressures underlying the long necks of extant giraffes and extinct sauropod dinosaurs. Here we summarise these debates and place them in a wider taxonomic context. We consider the evolution of long necks across a wide range of (both living and extinct) taxa and ask whether there has been a common selective factor or whether each case has a separate explanation. We conclude that in most cases long necks can be explained in terms of foraging requirements, and that alternative explanations in terms of sexual selection, thermoregulation and predation pressure are not as well supported. Specifically, in giraffe, tortoises, and perhaps sauropods there is likely to have been selection for high browsing. It the last case there may also have been selection for reaching otherwise inaccessible aquatic plants or for increasing the energetic efficiency of low browsing. For camels, wading birds and ratites, original selection was likely for increased leg length, with correlated selection for a longer neck to allow feeding and drinking at or near substrate level. For fish-eating long-necked birds and plesiosaurs a small head at the end of a long neck allows fast acceleration of the mouth to allow capture of elusive prey. A swan's long neck allows access to benthic vegetation, for vultures the long neck allows reaching deep into a carcass. Geese may be an unusual case where anti-predator vigilance is important, but so may be energetically efficient low browsing. The one group for which we feel unable to draw firm conclusions are the pterosaurs, this is in keeping with the current uncertainty about the biology of this group. Despite foraging emerging as a dominant theme in selection for long necks, for almost every taxonomic group we have identified useful empirical work that would increase understanding of the selective costs and benefits of a long neck.
... The unearthing of Pteranodon eclipsed previous pterosaur discoveries from the English Chalk and Cambridge Greensand but there is little need to recount its early history, which has been adequately examined by Bennett (1994Bennett ( , 2001 and Everhart (2005). Pteranodon was the first of the giant pterosaurs to be known from more than just fragments (see Witton 2010) and it became iconic as one of the central 'prehistoric characters' in film versions of Arthur Conan Doyle's (1922) Lost World and, latterly, Michael Crichton's Jurassic Park: The Lost World (Cearadactylus in the novel, Pteranodon in the film). The discovery of this incredible animal, with a wingspan now reliably known to be around 6-7 m, projected Othniel C. Marsh into the limelight and drew attention away from Britain: the Great American dinosaur rush had begun. ...
The first pterosaur fossil was described by Cosimo Alessandro Collini in 1784, but the epithet ptero dactyle was not applied until Georges Cuvier recognized the fossil as that of a volant animal in 1801. In eighteenth-century Britain, pterosaur bones had been discovered in Jurassic strata at Stonesfield, Oxfordshire but were considered to be bird bones, and largely went unnoticed. Bones of pterosaurs considerably larger than those of the first pterosaurs were discovered in the early nineteenth century by Gideon Mantell, but because of their comparatively large size were considered by Cuvier to also be the bones of birds. This perception by early nineteenth-century palaeontologists, including William Buckland and Gideon Mantell, that pterosaurs were relatively small animals was probably the reason their remains went unrecognized in British Jurassic and Cretaceous strata for several decades. Furthermore, the eighteenth-and early nineteenth-century dogmatic acceptance that fossil birds were present in the Jurassic Stonesfield 'slate' of Oxfordshire delayed the identification of medium-sized pterosaurs until the late 1820s, when Dean William Buckland described the Liassic Pterodactylus (= Dimorphodon) macronyx in 1829. Even after that date many fragmentary, but large, pterosaur bones were misidentified as avian, despite there being no convincing evidence for Mesozoic birds until the discovery of Archaeopteryx in the 1860s. Truly gigantic pterosaurs were first discovered in Great Britain some 20 years before Pteranodon was found in the Late Cretaceous of Kansas. However, the British material was so fragmentary that it was easily eclipsed by the spectacular, near-complete skeletons of Pteranodon found by O. C. Marsh and others from the 1870s onwards.
The Campanian Beloe Ozero locality within the Rybushka Formation in Saratov Province, Russia, is one of the richest and most diverse Upper Cretaceous pterosaur localities in Europe. It produces identifiable remains of Pteranodontidae indet. and Azhdarchidae indet., as well as bones which can be attributed to either of these groups. The pteranodontid specimens from the Beloe Ozero locality described in this paper include a cervical III, distal scapula, hume-rus deltopectoral crest, proximal syncarpal, preaxial carpal and complete femur. Based on the femur and proximal syncarpal, the wingspan estimate for the Beloe Ozero pteranodontid varies from 5.2 to 6.5 m. Volgadraco bogolubovi, known from the neighbouring Shyrokii Karamysh locality of the same formation and attributed previously to the Azhdarchidae, is more likely pteranodontid than azhdarchid. The other putative records of the Pteranodontidae in the Late Cretaceous of North America, Europe and Asia are discussed. Pteranodontid pterosaurs had a much wider distribution on the northern continents in the Late Cretaceous than previously thought.
Pterosaurs have fascinated scientists and nonscientists alike for over 200 years, as one of the three known clades of vertebrates to have evolved flapping flight. The smallest pterosaurs were comparable in size to the smallest extant birds and bats, but the largest pterosaurs were vastly larger than any extant flier. This immense size range, coupled with poor preservation and adaptations for flight unknown in extant vertebrates, have made interpretations of pterosaur flight problematic and often contentious. Here we review the anatomical, evolutionary, and phylogenetic history of pterosaurs, as well as the views, perspectives, and biases regarding their interpretation. In recent years, three areas of pterosaur biology have faced challenges and made advances: structure of the wing membrane, function of the pteroid, body size and mass estimates, as well as flight mechanics and aerodynamics. Comparative anatomical and fossil study, simulated bone loading, and aerodynamic modeling have all proved successful in furthering our understanding of pterosaur flight. Weagree with previous authors that pterosaurs should be studied as pterosaurs, a diverse but phylogenetically, anatomically, and mechanically constrained clade that can offer new insights into the diversity of vertebrate flight. © 2015, National Research Council of Canada, All Rights Reserved.
