Chaoyangopterids | |
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Holotype specimen of Jidapterus | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | † Pterosauria |
Suborder: | † Pterodactyloidea |
Clade: | † Neoazhdarchia |
Family: | † Chaoyangopteridae Lü et al., 2008 |
Type species | |
† Chaoyangopterus zhangi Wang & Zhou, 2003 | |
Genera | |
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Chaoyangopteridae (or chaoyangopterids) is a family of pterosaurs within the larger group Azhdarchoidea. Chaoyangopterids lived mostly during the Early Cretaceous period, though possible members, Microtuban , Xericeps and Argentinadraco , may extend the fossil range to the Late Cretaceous. [1] [7]
The clade Chaoyangopteridae was first defined in 2008 by Lü Junchang and David Unwin as: " Chaoyangopterus , Shenzhoupterus , their most recent common ancestor and all taxa more closely related to this clade than to Tapejara , Tupuxuara or Quetzalcoatlus ". [8] Based on neck and limb proportions, it has been suggested they occupied a similar ecological niche to that of azhdarchid pterosaurs, [9] though it is possible they were more specialized as several genera occur in Liaoning, while azhdarchids usually occur by one genus in a specific location. [10]
Chaoyangopterids are distinguished from other pterosaurs by several traits of the nasoantorbital fenestra, a large hole on the side of the snout formed by the assimilation of the nares (nostril holes) into the antorbital fenestra. In members of this family, the nasoantorbital fenestra is massive, with the rear edge extending as far back as the braincase and jaw joint. The front edge is formed by a rod of bone known as the premaxillary bar, which is unusually slender in members of this family. [8]
Below is a cladogram showing the phylogenetic analysis conducted by paleontologist Brian Andres and colleagues in 2014. In the analysis, Chaoyangopteridae was found to consist of the genus Eoazhdarcho and the subfamily Chaoyangopterinae. [11] The subfamily Chaoyangopterinae was initially used by paleontologist Felipe Pinheiro and colleagues in 2011, which they assigned to the family Tapejaridae, [12] however, Andres and colleagues redefined the subfamily as the least inclusive clade containing Chaoyangopterus zhangi and Shenzhoupterus chaoyangensis. In the analysis by Andres and colleagues, Chaoyangopterinae consisted of the pterosaurs Chaoyangopterus, Shenzhoupterus, and Jidapterus. [11]
Neoazhdarchia | |
A 2021 study focused on Aerotitan recovers Xericeps and Argentinadraco as sister taxa within Chaoyangopteridae. [13]
Like their azhdarchid relatives, pterosaurs that belong to the Chaoyangopteridae were terrestrial predators. [14] [15]
The members of the family Chaoyangopteridae are mostly known from Asia, though the possible member Lacusovagus occurs in South America [10] and there are possible fossil remains from Africa, including the possible member Apatorhamphus . [16] Microtuban may extend the clade's existence into the early Late Cretaceous. [14]
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.
Tropeognathus is a genus of large pterosaurs from the late Early Cretaceous of South America. This genus is considered to be a member of the family Anhangueridae, however, several studies have also recovered it within another family called Ornithocheiridae. Both of these families are diverse groups of pterosaurs known for their keel-tipped snouts and large size. Tropeognathus is regarded as the largest pterosaur found in the Southern Hemisphere, only rivaled by the huge azhdarchids. The type and only species is Tropeognathus mesembrinus. Fossil remains of Tropeognathus have been recovered from the Romualdo Formation, which is a Lagerstätte located in the Santana Group of the Araripe Basin in northeastern Brazil.
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. A second species, E. yixianensis, was named in 2023, but its validity has been questioned.
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.
Sinopterus is a genus of tapejarid pterodactyloid pterosaur from the Aptian-age Lower Cretaceous Jiufotang Formation of Chaoyang, Liaoning, China. It was first described and named by Wang Xiaolin and Zhou Zhonghe. Historically, there were multiple species attributed to the genus although only one is considered to be valid. Sinopterus is known for its proportionally large skull, which has a birdlike pointed beak, a long bony crest that starts with a tall premaxilla and goes back along the middle of the skull to form a point overhanging the rear of the skull, and its lack of teeth.
Lonchodectidae or Lonchodraconidae is a group of pterosaurs within the clade Pterodactyloidea. It has variously been considered to be within Ctenochasmatoidea, Azhdarchoidea and Pteranodontoidea. They are notable for their high, conical tooth sockets and raised alveolar margins.
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.
Shenzhoupterus is a genus of chaoyangopterid pterosaur from the Jiufotang Formation of modern-day Liaoning, China. Fossil remains of Shenzhoupterus date back to the Early Cretaceous period, approximately 120 million years ago.
The Cambridge Greensand is a geological unit in England whose strata are earliest Cenomanian in age. It lies above the erosive contact between the Gault Formation and the Chalk Group in the vicinity of Cambridgeshire, and technically forms the lowest member bed of the West Melbury Marly Chalk Formation. It is a remanié deposit, containing reworked fossils of late Albian age, including those of dinosaurs and pterosaurs.
Alanqa is a genus of pterodactyloid pterosaur from the Late Cretaceous period of what is now the Kem Kem Beds of southeastern Morocco. The name Alanqa comes from the Arabic word العنقاءal-‘Anqā’, for a mythical bird of Arabian culture.
Thalassodrominae or Thalassodromidae is a group of azhdarchoid pterosaurs from the Cretaceous period. Its traditional members come from Brazil, however, other possible members also come from other places, including Morocco and Argentina. Thalassodrominae is considered either to be a subfamily within the pterosaur family Tapejaridae, or as a distinct family, Thalassodromidae, within the clade Neoazhdarchia, closely related to dsungaripterids or azhdarchids.
Aerotitan is a genus of large azhdarchid pterosaur known from the Late Cretaceous period of what is now the Allen Formation of the Neuquén Basin in northern Patagonia, Argentina.
Banguela is a genus of azhdarchoid pterosaur from the Early Cretaceous period of what is now Brazil. Only one species is known, Banguela oberlii.
Tapejaroidea is a group of pterosaurs belonging to the clade Ornithocheiroidea. Tapejaroids lived from the Early to Late Cretaceous periods, with one possible member, Tendaguripterus, extending the fossil range to the Late Jurassic period. Tapejaroidea contains two groups, the Dsungaripteridae and the Azhdarchoidea, which in turn includes the azhdarchids, the group that contains some of the largest flying animals. The group was named by Brazilian paleontologist Alexander Wilhelm Armin Kellner in 1996.
Xericeps is a genus of pterosaur from the Cenomanian stage of the Late Cretaceous. It was discovered from the Kem Kem Beds of southeastern Morocco.
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.
Keresdrakon is an extinct genus of azhdarchoid pterosaur from the Goio-Erê Formation of Brazil, which dates back to the Early Cretaceous period, 125 to 100.5 million years ago. Keresdrakon contains a single species, Keresdrakon vilsoni.
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)
Nicorhynchus is a genus of anhanguerid pterosaur from the Cretaceous period. It contains two species, the type species, N. capito, from the Cambridge Greensand of England, and N. fluviferox from the Kem Kem Group of Morocco. These species were previously assigned to Coloborhynchus.
Huaxiadraco is a genus of tapejarid pterodactyloid pterosaur from the Aptian-age Lower Cretaceous Jiufotang Formation of Chaoyang, Liaoning, China. It is the third valid genus of tapejarid from the Jehol Biota, after Sinopterus and Eopteranodon. It contains one species, Huaxiadraco corollatus, originally assigned to the defunct genus Huaxiapterus.