21-519: Eupterodactyloidea (meaning "true Pterodactyloidea ") is an extinct group of pterodactyloid pterosaurs that existed from the latest Late Jurassic to the latest Late Cretaceous periods ( Tithonian to Maastrichtian stages). Eupterodactyloids have been found on all continents except Antarctica . Eupterodactyloidea was named by S. Christopher Bennett in 1994 as an infraorder of the suborder Pterodactyloidea. Bennett defined it as an apomorphy -based clade . However, in 2010, Brian Andres re-defined
42-1472: A branch-based definition for Eupterodactyloidea, making them very similar in content. Haopterus gracilis Piksi barbarulna Tethydraco regalis Pteranodon longiceps [REDACTED] Pteranodon sternbergi Alamodactylus byrdi Volgadraco bogolubovi Cretornis hlavaci Alcione elainus Simurghia robusta Muzquizopteryx coahuilensis Barbaridactylus grandis Nyctosaurus lamegoi Nyctosaurus nanus Nyctosaurus gracilis [REDACTED] Hongshanopterus lacustris Lonchodraco giganteus Lonchodectes compressirostris Boreopterus cuiae Zhenyuanopterus longirostris Nurhachius ignaciobritoi Liaoxipterus brachyognathus Istiodactylus sinensis Istiodactylus latidens Aetodactylus halli Cimoliopterus dunni Cimoliopterus cuvieri Guidraco Ludodactylus [REDACTED] Cearadactylus Brasileodactylus Anhangueridae [REDACTED] Ornithocheiridae Bennettazhia oregonensis Tapejara wellnhoferi Europejara olcadesorum Vectidraco daisymorrisae Caiuajara dobruskii Tupandactylus navigans Tupandactylus imperator [REDACTED] Bakonydraco galaczi [REDACTED] "Huaxiapterus" benxiensis "Huaxiapterus" corollatus Eopteranodon lii Huaxiapterus jii Sinopterus dongi Pterodactyloidea Pterodactyloidea (derived from
63-1247: A different type of analysis have found that this basic division into primitive (archaeopterodactyloid) and advanced (eupterodactyloid) species may not be correct. Beginning in 2014, Steven Vidovic and David Martill constructed an analysis in which several pterosaurs traditionally thought of as archaeopterodactyloids closely related to the ctenochasmatoids may have been more closely related to ornithocheiroids , or in some cases, fall outside both groups. The results of their updated 2017 analysis are shown below. Eosipterus yangi Pterodactylus antiquus [REDACTED] Diopecephalus kochi Aerodactylus scolopaciceps [REDACTED] Aurorazhdarchidae Gallodactylidae Ctenochasmatidae Altmuehlopterus ramphastinus [REDACTED] Germanodactylus cristatus Elanodactylus prolatus Lonchodectes compressirostris [REDACTED] Prejanopterus curvirostra Kepodactylus insperatus Dsungaripteridae [REDACTED] Ornithocheiromorpha [REDACTED] Hamipterus tianshanensis Ikrandraco avatar Istiodactylus Nyctosauridae [REDACTED] Pteranodon [REDACTED] Sinopterus dongi Bennettazhia oregonensis Bennettazhia
84-560: A much more exclusive group including only the branch of traditional ornithocheirid pterosaurs, though this use has since fallen out of favor by many researchers after years of competing definitions for the various pterodactyloid clades. The compromise definitions by Andres and others have since become more widely adopted. Below is a cladogram showing the results of a phylogenetic analysis first presented by Andres et al. in 2014, and updated by Longrich, Martill, and Andres in 2018. Andres and colleagues followed this definition, and also used
105-673: A new species of Pteranodon : P. oregonensis . A humerus ( holotype MPUC V.126713 ), two fused dorsal vertebrae and the broken-off end of some joint bone had been unearthed from the Lower Cretaceous ( Albian stage) beds of the Hudspeth Formation in Wheeler County , Oregon , United States, to which the specific epithet refers. Gilmore noted similarities to Nyctosaurus though the specimens were larger. In 1989, S. Christopher Bennett concluded that
126-471: A pterosaur. O'Sullivan and Martill (2018) described a partial synsacrum from the Stonesfield Slate identified as possibly pterodactyloid based on the number of incorporated sacrals although they commented that the morphology was perhaps closer to that of wukongopterids. If correctly identified, it would be the oldest pterodactyloid fossil known. Pterodactyloidea is traditionally considered to be
147-619: Is a genus of tapejaromorph pterosaur which lived during the Albian stage of the Early Cretaceous from what is now the Hudspeth Formation of the state of Oregon in the United States . Although originally identified as a species of the pteranodontoid pterosaur Pteranodon , Bennettazhia is now thought to have been a different animal. The type and only species is B. oregonensis . In 1928, Charles Gilmore named
168-561: Is about 5 million years older than the oldest previously known confirmed specimens. Previously, a fossil jaw recovered from the Middle Jurassic Stonesfield Slate formation in the United Kingdom, was considered the oldest known. This specimen supposedly represented a member of the family Ctenochasmatidae , though further examination suggested it belonged to a teleosaurid stem-crocodilian instead of
189-473: Is also a common term for pterodactyloid pterosaurs, though it can also be used to refer to Pterodactylus specifically. Well-known examples of pterodactyloids include Pterodactylus , Pteranodon , and Quetzalcoatlus . In 2014, fossils from the Shishugou Formation of China were classified as the most basal pterodactyloid yet found, Kryptodrakon . At a minimum age of about 161 my, it
210-536: Is unwarped. Both dsungaripterids and azhdarchoids show this feature, but only the latter group is typified by such a very thin outer bone wall. Habib concluded that Bennettazhia was a member of the Azhdarchoidea , a more encompassing group than the Azhdarchidae. The cladogram below follows a 2014 phylogenetic analysis by Brian Andres and colleagues. In the analysis, they placed Bennettazhia within
231-855: The Dsungaripteridae . Since then, some authors including Bennett himself have classified this genus as a sister group to the Tapejaridae or possibly a member of the Thalassodrominae . In 2023, the discovery of guano and unusual fragmentation of ammonites from the Hudspeth Formation have been interpreted as evidence that this pterosaur ate molluscs and formed large colonies on nearby cliffs similar to modern seagull rookeries. New remains of this pterosaur are also reported, including two isolated teeth (F127985A and F127910B) and an edentulous (teeth-lacking) section of
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#1732801274776252-759: The Eupterodactyloidea . Anurognathidae [REDACTED] Kryptodrakon [REDACTED] "Painten pro-pterodactyloid" ( Propterodactylus ) Germanodactylidae [REDACTED] Pterodactylus [REDACTED] Gallodactylidae Ctenochasmatidae [REDACTED] Haopterus Serradraco Aussiedraco Lonchodectidae [REDACTED] Lonchodraconidae Istiodactylidae Ornithocheiriformes [REDACTED] Pteranodontia [REDACTED] Tapejaridae [REDACTED] Thalassodromidae [REDACTED] Dsungaripteridae [REDACTED] Chaoyangopteridae Radiodactylus Azhdarchidae [REDACTED] Some studies based on
273-534: The Greek words πτερόν ( pterón , for usual ptéryx ) "wing", and δάκτυλος ( dáktylos ) "finger") is one of the two traditional suborders of pterosaurs ("wing lizards"), and contains the most derived members of this group of flying reptiles. They appeared during the middle Jurassic Period, and differ from the basal (though paraphyletic ) rhamphorhynchoids by their short tails and long wing metacarpals (hand bones). The most advanced forms also lack teeth, and by
294-453: The bone structure. Apart from the thin bone wall, the humerus was filled with a spongy tissue consisting of trabeculae , very thin bone layers and struts, forming a light yet strong construction. Habib inferred that such strength would have allowed even very large pterosaurs to launch themselves from the ground using their forelimbs. The same investigation made a better classification possible. The humerus has an elongated deltopectoral crest that
315-499: The group as a stem-based taxon in his dissertation, and then formalized the definition in 2014 as all pterosaurs more closely related to Pteranodon longiceps than to Pterodactylus antiquus . The slightly more exclusive group Ornithocheiroidea was re-defined in 2003 by Alexander Kellner. He defined it as the least inclusive clade containing Anhanguera blittersdorffi , Pteranodon longiceps , Dsungaripterus weii , and Quetzalcoatlus northropi . Ornithocheiroidea has often been used for
336-468: The group of short-tailed pterosaurs with long wrists (metacarpus), compared with the relatively long tails and short wrist bones of basal pterosaurs ("rhamphorhynchoids"). In 2004, Kevin Padian formally defined Pterodactyloidea as an apomorphy -based clade containing those species possessing a metacarpal at least 80% of the length of the humerus , homologous with that of Pterodactylus . This definition
357-490: The late Cretaceous, all known pterodactyloids were toothless. Many species had well-developed crests on the skull, a form of display taken to extremes in giant-crested forms like Nyctosaurus and Tupandactylus . Pterodactyloids were the last surviving pterosaurs when the order became extinct at the end of the Cretaceous Period, together with the non-avian dinosaurs and most marine reptiles. " Pterodactyl "
378-421: The lower mandible (F127960). Bennettazhia was a medium-sized pterosaur with an estimated wingspan of 4 metres (13 ft). In 2007, American biologist Michael Habib revealed the result of a study by CAT-scan of the type specimen of Bennettazhia . The humerus, 183 millimeters (7.2 in) long, is uncrushed, which is uncommon for a pterosaur fossil and therefore offered a rare opportunity to investigate
399-484: The remains might be those of a member of the Azhdarchidae instead of a pteranodontid . In 1991, Russian paleontologist Lev Nesov therefore named a new azhdarchid genus: Bennettazhia . The genus name honors Bennett and combines his name with Persian azhdarha , "dragon", a reference to Azhdarcho , the type genus of the Azhdarchidae. Bennett himself in 1994 changed his opinion and stated that it belonged to
420-543: Was adopted by the PhyloCode in 2020. A subgroup of pterodactyloids, called the Lophocratia , was named by David Unwin in 2003. Unwin defined the group as the most recent common ancestor of Pterodaustro guinazui and Quetzalcoatlus northropi , and all its descendants. This group was named for the presence of a head crest in most known species, though this feature has since been found in more primitive pterosaurs and
441-400: Was probably an ancestral feature for all pterodactyloids. There are competing theories of pterodactyloid phylogeny. Below is cladogram following a topology recovered by Brian Andres, using the most recent iteration of his data set (Andres, 2021). This study found the two traditional groupings of ctenochasmatoids and kin as an early branching group, with all other pterodactyloids grouped into
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