21-794: 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 . 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 ". Based on neck and limb proportions, it has been suggested they occupied
42-1087: A more recent phylogenetic analysis, by Kellner and colleagues in 2019, had recovered Azhdarchoidea within the larger group Tapejaroidea. Unlike the analysis by Andres and colleagues, Kellner and colleagues had found Azhdarchoidea to only consist of three groups: Azhdarchidae , Chaoyangopteridae, and Tapejaromorpha . Their cladogram is shown below. Dsungaripterus weii Noripterus parvus Azhdarcho lancicollis Quetzalcoatlus northropi [REDACTED] Zhejiangopterus linhaiensis Chaoyangopterus zhangi Jidapterus edentus Shenzhoupterus chaoyangensis Keresdrakon vilsoni Thalassodromeus sethi Tupuxuara leonardii [REDACTED] Caupedactylus ybaka Aymberedactylus cearensis Eopteranodon lii "Huaxiapterus" benxiensis "Huaxiapterus" corollatus Sinopterus dongi Europejara olcadesorum Caiuajara dobruskii Tapejara wellnhoferi Tupandactylus imperator [REDACTED] In 2022, Pêgas et al. named and officially registered two new clades: Azhdarchomorpha ,
63-482: A similar ecological niche to that of azhdarchid pterosaurs, 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. 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
84-992: Is a cladogram showing the results of a phylogenetic analysis presented by Andres and colleagues in 2014. This study found the a grouping of tapejarids at the base of the clade, with thalassodromines more closely related to azhdarchids and chaoyangopterids, as well as dsungaripterids. Their cladogram is shown below. Bennettazhia oregonensis Eopteranodon lii "Sinopterus" gui Nemicolopterus crypticus Huaxiapterus jii Tapejaridae [REDACTED] Dsungaripterus weii Domeykodactylus ceciliae [REDACTED] Noripterus parvus Noripterus complicidens Thalassodromeus sethi Tupuxuara longicristatus [REDACTED] Tupuxuara leonardii Eoazhdarcho liaoxiensis Shenzhoupterus chaoyangensis Chaoyangopterus zhangi Jidapterus edentus Azhdarcho lancicollis Zhejiangopterus linhaiensis Arambourgiania philadelphiae Quetzalcoatlus northropi [REDACTED] The result of
105-642: 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
126-792: The Ctenochasmatoidea , as the sister taxon of the Pteranodontoidea within the Ornithocheiroidea, or within the Tapejaroidea , which in turn was also within the Ornithocheiroidea. Azhdarchoidea was given a phylogenetic definition by David Unwin in 2003. Unwin defined the group as the most recent common ancestor of Quetzalcoatlus and Tapejara , and all its descendants. There have been several competing views of azhdarchoid relationships. The first, presented by Felipe Pinheiro and colleagues in 2011, considered
147-424: 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. Below is a cladogram showing the phylogenetic analysis conducted by paleontologist Brian Andres and colleagues in 2014. In
168-488: The tapejarids to be a monophyletic clade including the thalassodromines and chaoyangopterines . The second, found by Naish & Martill (2006), as well as Lü et al. (2008), considered the traditional "tapejarids" to be a paraphyletic grade of primitive azhdarchoids. with true tapejarids most basal, and the thalassodromines (alternatively called thalassodromids) and chaoyangopterids being successively more closely related to azhdarchids . All azhdarchoids which are part of
189-626: The Chaoyangopteridae were terrestrial predators . The members of the family Chaoyangopteridae are mostly known from Asia, though the possible member Lacusovagus occurs in South America and there are possible fossil remains from Africa, including the possible member Apatorhamphus . Microtuban may extend the clade's existence into the early Late Cretaceous . [REDACTED] [REDACTED] [REDACTED] Azhdarchoidea Azhdarchoidea (or azhdarchoids )
210-615: The Late Cretaceous of northern Lebanon . Microtuban is known only from a partially preserved skeleton lacking the skull, holotype SMNK PAL 6595 . It was acquired from a local fossil dealer by the Staatliches Museum für Naturkunde Karlsruhe and was, as can be deduced from the qualities of the stone matrix, collected in the locality of Hjoûla in a marine layer of the Sannine Formation , dating to
231-754: The analysis by Andres and colleagues, Chaoyangopterinae consisted of the pterosaurs Chaoyangopterus , Shenzhoupterus , and Jidapterus . Chaoyangopterus zhangi Jidapterus edentus Shenzhoupterus chaoyangensis Eoazhdarcho liaoxiensis Azhdarcho lancicollis Zhejiangopterus linhaiensis Arambourgiania philadelphiae Quetzalcoatlus northropi A 2021 study focused on Aerotitan recovers Xericeps and Argentinadraco as sister taxa within Chaoyangopteridae. Shenzhoupterus chaoyangensis Xericeps curvirostris Argentinadraco barrealensis Chaoyangopterus zhangi Jidapterus edentus Lacusovagus magnificens Like their azhdarchid relatives, pterosaurs that belong to
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#1732801697325252-439: The analysis, Chaoyangopteridae was found to consist of the genus Eoazhdarcho and the subfamily Chaoyangopterinae . The subfamily Chaoyangopterinae was initially used by paleontologist Felipe Pinheiro and colleagues in 2011, which they assigned to the family Tapejaridae , however, Andres and colleagues redefined the subfamily as the least inclusive clade containing Chaoyangopterus zhangi and Shenzhoupterus chaoyangensis . In
273-416: The clade formed by Quetzalcoatlus and Tupuxuara are included in the group Neoazhdarchia ("new azhdarchids") as defined by Unwin in 2003. In 1996, Alexander Kellner created a different clade called Tapejaroidea , which he defined as the most recent common ancestor and all descendants of Tapejara , Quetzalcoatlus , and Dsungaripterus . Kellner created this clade to include both Azhdarchoidea and
294-581: The early Cenomanian stage of the Late Cretaceous , about 99.6–96 million years ago . The fossil dealer however, indicated it came from a quarry at nearby Hâqel . Regardless, it is only the second pterosaur fossil found in Lebanon and a rare example of a pterosaur from the African continental plate, to which the area in Cretaceous times belonged. In this period the site was located hundreds of miles from
315-435: The family Dsungaripteridae , but as separate groups. A lot of recent studies have followed this concept. There are competing theories of azhdarchoid phylogeny; it is either recovered as closely related to the clade Ctenochasmatoidea , or within the group Ornithocheiroidea , either as the sister taxon of the Pteranodontoidea or within the clade Tapejaroidea. The latter two of which are more widely accepted. Below
336-655: The four known azhdarchoid groups they excluded the Tapejaridae and Azhdarchidae , leaving the possibility that Microtuban was member of either the Thalassodromidae or the Chaoyangopteridae . As the two latter groups are mainly distinguished on the basis of traits in the skull, the very body part the Microtuban holotype lacks, it proved impossible to decide whether it were a thalassodromid or
357-401: The mainland coastline. The specimen consists of the last cervical and first dorsal vertebrae, a shoulder girdle, the left wing and fragments of the hindlimbs. It represents an immature individual. Many bones are broken but the whole is still articulated. This was explained as being caused by a collision or a bite by a predator, just before or after death, after which the carcass quickly sank to
378-400: The most inclusive clade containing Azhdarcho but not Tapejara or Thalassodromeus , and Alanqidae , containing Alanqa but not Chaoyangopterus or Azhdarcho . Their original concept of Alanqidae was later refuted and updated by Pêgas. The updated phylogeny of Pêgas (2024) is shown below: Microtuban Microtuban is an extinct genus of azhdarchoid pterosaur known from
399-433: The sea bottom. Microtuban was named by Ross A. Elgin and Eberhard Frey in 2011 and the type species is Microtuban altivolans . The generic name is derived from Greek μικρός, mikros , "little", and Arabic ثعبان, tuban , "dragon" or basilisk , also referring to Thuban , the ancient Polar Star in the constellation Draco . The specific name is derived from Latin altus , "high", and volare , "to fly". As
420-482: The second phalanx of the wing-finger is long, with 85% of the length of the first phalanx; the fourth phalanx is extremely reduced measuring but 1.1% of wing-finger length. Possible juvenile traits included the lack of a notarium , being a fusion of the front dorsal vertebrae, and an unfused scapula and coracoid . The describers assigned Microtuban to the Azhdarchoidea , using the comparative method. Of
441-501: The specimen is that of a not yet fully grown animal, it is difficult to estimate the adult size. The wingspan of the holotype can be deduced from the length of the wing elements. The upper arm, damaged in the fossil, measured about nine centimetres, the lower about seven. The fourth metacarpal is 122 millimetres long. The four phalanges of the, fourth, wing-finger measure 135, 114.5, 63.5 and 3.5 millimetres respectively. The authors indicated two unique derived traits, or autapomorphies :
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