Bennettazhia | |
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Skeletal diagram of Bennettazhia oregonensis | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | † Pterosauria |
Suborder: | † Pterodactyloidea |
Clade: | † Tapejaromorpha |
Genus: | † Bennettazhia Nesov, 1991 |
Type species | |
†Pteranodon oregonensis Gilmore, 1928 | |
Species | |
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Synonyms | |
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Bennettazhia 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 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, in the state of Oregon, United States, to which the specific epithet refers. Gilmore noted similarities to Nyctosaurus though the specimens were larger. [2]
In 1989, S. Christopher Bennett concluded that the remains might be those of a member of the Azhdarchidae instead of a pteranodontid. [3] 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. [4] Bennett himself in 1994 changed his opinion and stated that it belonged to the Dsungaripteridae. [5] 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. [6] [7]
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 the lower mandible (F127960). [1]
Bennettazhia was a medium-sized pterosaur with an estimated wingspan of 4 metres (13 ft). [1] 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 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 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. [8]
The cladogram below follows a 2014 phylogenetic analysis by Brian Andres and colleagues. In the analysis, they placed Bennettazhia within the clade Tapejaromorpha, as the basalmost member. [9]
Azhdarchoidea |
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Azhdarchidae is a family of pterosaurs known primarily from the Late Cretaceous Period, though an isolated vertebra apparently from an azhdarchid is known from the Early Cretaceous as well. Azhdarchids are mainly known for including some of the largest flying animals discovered, but smaller cat-size members have also been found. Originally considered a sub-family of Pteranodontidae, Nesov (1984) named the Azhdarchinae to include the pterosaurs Azhdarcho, Quetzalcoatlus, and Titanopteryx. They were among the last known surviving members of the pterosaurs, and were a rather successful group with a worldwide distribution. Previously it was thought that by the end of the Cretaceous, most pterosaur families except for the Azhdarchidae disappeared from the fossil record, but recent studies indicate a wealth of pterosaurian fauna, including pteranodontids, nyctosaurids, tapejarids and several indeterminate forms. In several analyses, some taxa such as Navajodactylus, Bakonydraco and Montanazhdarcho were moved from Azhdarchidae to other clades.
Tupuxuara is a genus of large, crested, and toothless pterodactyloid pterosaur from the Early Cretaceous period of what is now the Romualdo Formation of the Santana Group, Brazil, about 125 to 112 million years ago. Tupuxuara is a close relative of Thalassodromeus, and both form a group that is either called Thalassodrominae or Thalassodromidae.
Montanazhdarcho is a genus of azhdarchoid pterosaur from the Late Cretaceous period of what is now the state of Montana, United States. Montanazhdarcho is known from only one species, M. minor.
Pterodactyloidea is one of the two traditional suborders of pterosaurs, and contains the most derived members of this group of flying reptiles. They appeared during the middle Jurassic Period, and differ from the basal rhamphorhynchoids by their short tails and long wing metacarpals. The most advanced forms also lack teeth, and by 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.
Ornithostoma is a genus of pterodactyloid pterosaur that lived during the Early Cretaceous period of Europe, around 110 million years ago. Ornithostoma was once thought to have been a senior synonym of the pteranodontid Pteranodon due to its toothless anatomy and prior naming.
Eopteranodon is a genus of tapejarid pterosaur from the Aptian-age Lower Cretaceous Yixian Formation of Beipiao City, Liaoning, China. The genus was named in 2005 by paleontologists Lü Junchang and Zhang Xingliao. The type species is Eopteranodon lii.
Chaoyangopterus is a genus of chaoyangopterid pterosaur known from a partial skeleton found in Liaoning, China. Chaoyangopterus was found in rocks dating back to the Aptian-age Lower Cretaceous Jiufotang Formation of Dapingfang, Chaoyang.
Eoazhdarcho is a genus of azhdarchoid pterodactyloid pterosaur named in 2005 by Chinese paleontologists Lü Junchang and Ji Qiang. The type and only known species is Eoazhdarcho liaoxiensis. The fossil was found in the Aptian-age Lower Cretaceous Jiufotang Formation of Chaoyang, Liaoning, China.
Jidapterus is a genus of chaoyangopterid pterosaur from the Aptian-age Lower Cretaceous Jiufotang Formation of Chaoyang, Liaoning, China. The genus was in 2003 named by Dong Zhiming, Sun Yue-Wu and Wu Shao-Yuan. The type species is Jidapterus edentus. The genus name is derived from Jílín Dàxué or "Jilin University" and a Latinized Greek pteron, "wing". The specific name means "toothless" in Latin.
Santanadactylus was a genus of pterodactyloid pterosaur from the Albian-age Romualdo Member of the Upper Cretaceous Santana Formation, of Barra do Jardim, Araripe Plateau, Ceará State, Brazil. Four species have been named, but today are no considered congeneric with each other. It was a rather large pterosaur.
Azhdarchoidea is a group of pterosaurs within the suborder Pterodactyloidea, more specifically within the group Ornithocheiroidea. Pterosaurs belonging to this group lived throughout the Early and Late Cretaceous periods, with one tentative member, Tendaguripterus, that lived in the Late Jurassic period. The largest azhdarchoids include members of the family Azhdarchidae, examples of these are Quetzalcoatlus, Hatzegopteryx, and Arambourgiania. The Azhdarchoidea has been recovered as either closely related to the Ctenochasmatoidea, as the sister taxon of the Pteranodontoidea within the Ornithocheiroidea, or within the Tapejaroidea, which in turn was also within the Ornithocheiroidea.
Lacusovagus is a genus of azhdarchoid pterodactyloid pterosaur from the Lower Cretaceous of Brazil. It is based on SMNK PAL 4325, a partial upper jaw comprising sections of the skull in front of the eyes. This specimen was found in rocks of the Early Cretaceous-age Nova Olinda Member of the Crato Formation. The skull was long, and unusually wide. The section in front of the combined nasal-antorbital fenestra was relatively short. Also unusual was the combination of its toothless jaws and no bony head crest. Lacusovagus was described in 2008 by Mark Witton. The type species is L. magnificens, meaning "grand lake wanderer", in reference to its large size—it is currently the largest pterosaur known from the Crato Formation with an estimated wingspan of approximately 4 metres (13 ft) and a body mass of 30 kg (66 lb).
Caupedactylus is an extinct genus of tapejarid azhdarchoid pterosaur known from the Early Cretaceous period of what is now the Romualdo Formation of the Araripe Basin in northeastern Brazil. The type and only species of Caupedactylus is Caupedactylus ybaka.
Radiodactylus is an extinct genus of non-azhdarchid azhdarchoid pterosaur known from the Early Cretaceous period of what is now Texas, southern United States. It contains a single species, Radiodactylus langstoni.
This timeline of pterosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, and taxonomic revisions of pterosaurs, the famed flying reptiles of the Mesozoic era. Although pterosaurs went extinct millions of years before humans evolved, humans have coexisted with pterosaur fossils for millennia. Before the development of paleontology as a formal science, these remains would have been interpreted through a mythological lens. Myths about thunderbirds told by the Native Americans of the modern Western United States may have been influenced by observations of Pteranodon fossils. These thunderbirds were said to have warred with water monsters, which agrees well with the co-occurrence of Pteranodon and the ancient marine reptiles of the seaway over which it flew.
Argentinadraco is an extinct genus of azhdarchoid pterosaur from the Late Cretaceous Portezuelo Formation of Argentina. It contains a single species, A. barrealensis, named in 2017 by Alexander Kellner and Jorge Calvo. Argentinadraco is unusual for bearing a bottom jaw with a concave bottom edge, as well as a pair of ridges and depressions on the top surface. These features distinguish it from all other azhdarchoid groups, complicating its assignment, but it may belong to the Chaoyangopteridae. The ridges on the lower jaw may have been used to feed on small invertebrates in loose sediment within the system of lakes and rivers that it resided in.
Apatorhamphus is an extinct genus of azhdarchoid pterosaur from the Kem Kem Group of Morocco. It might have been part of the Chaoyangopteridae. It is only known from a few snout fragments and it likely had a wingspan of between 3–7 metres (9.8–23.0 ft)
Leptostomia is a genus of long-beaked pterosaur from the mid-Cretaceous (Cenomanian) of Morocco, North Africa. The type species is L. begaaensis, which was named and described in 2021 from sediments of the Kem Kem Group in Morocco. It was a small animal with a long, slender bill which is thought to have been used to probe sediments for worms and other invertebrates, similar to kiwi birds and curlews. Leptostomia is likely a member of the Azhdarchoidea.
Wellnhopterus is an azhdarchid pterosaur recovered from the Late Cretaceous Javelina Formation in Texas that was previously identified as a thalassodromine. It consists of a set of upper and lower jaws, as well as some cervical vertebrae and a fragmentary long bone. In July 2021, the jaws were given the genus name "Javelinadactylus", with the type and only species as "J. sagebieli"; however, this article has now been retracted. In a paper published in December 2021, the complete holotype was independently named Wellnhopterus, with the only species being W. brevirostris. As of 2022, this is the formal name of this pterosaur.