Chaoyangopteridae
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Chaoyangopteridae (or chaoyangopterids) is a family of pterosaurs within the larger group
Azhdarchoidea Azhdarchoidea (or azhdarchoids) is a group of pterosaurs within the suborder Pterodactyloidea, more specifically within the group Ornithocheiroidea. Pterosaurs belonging to this group lived throughout the Early Cretaceous, Early and Late Cretaceo ...
. Chaoyangopterids lived mostly during the
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
period, though possible members, ''
Microtuban ''Microtuban'' is an extinct genus of azhdarchoid pterosaur known from the Late Cretaceous of northern Lebanon. Discovery ''Microtuban'' is known only from a partially preserved skeleton lacking the skull, holotype SMNK PAL 6595. It was acquire ...
'', '' Xericeps'' and ''
Argentinadraco ''Argentinadraco'' (meaning "Argentina dragon") 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 Jo ...
'', may extend the fossil range to the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
.


History

The
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
Chaoyangopteridae was first defined in 2008 by
Lü Junchang Lü Junchang (; 1965 – 9 October 2018) was a Chinese palaeontologist and professor at the Institute of Geology, Chinese Academy of Geological Sciences. An expert on Mesozoic reptiles, he described and named dozens of dinosaur and pterosaur taxa ...
and
David Unwin David (; , "beloved one") (traditional spelling), , ''Dāwūd''; grc-koi, Δαυΐδ, Dauíd; la, Davidus, David; gez , ዳዊት, ''Dawit''; xcl, Դաւիթ, ''Dawitʿ''; cu, Давíдъ, ''Davidŭ''; possibly meaning "beloved one". w ...
as: "'' Chaoyangopterus'', '' Shenzhoupterus'', their
most recent common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...
and all taxa more closely related to this clade than to '' Tapejara'', ''
Tupuxuara ''Tupuxuara'' is a genus of large, crested, and toothless Pterodactyloidea, pterodactyloid pterosaur from the Early Cretaceous period (Albian stage) of what is now the Romualdo Formation of the Santana Group, Brazil, about 125 to 112 million ye ...
'' or ''
Quetzalcoatlus ''Quetzalcoatlus'' is a genus of pterosaur known from the Late Cretaceous period of North America (Maastrichtian stage); its members were among the largest known flying animals of all time. ''Quetzalcoatlus'' is a member of the Azhdarchidae, ...
''". Based on neck and limb proportions, it has been suggested they occupied a similar ecological niche to that of
azhdarchid Azhdarchidae (from the Persian word , , a dragon-like creature in Persian mythology) 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 Cret ...
pterosaurs, though it is possible they were more specialized as several genera occur in
Liaoning Liaoning () is a coastal province in Northeast China that is the smallest, southernmost, and most populous province in the region. With its capital at Shenyang, it is located on the northern shore of the Yellow Sea, and is the northernmost ...
, while azhdarchids usually occur by one genus in a specific location.


Description

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 An antorbital fenestra (plural: fenestrae) is an opening in the skull that is in front of the eye sockets. This skull character is largely associated with archosauriforms, first appearing during the Triassic Period. Among extant archosaurs, birds ...
. 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.


Classification

Below is a
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
showing the
phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
conducted by paleontologist
Brian Andres Brian (sometimes spelled Bryan in English) is a male given name of Irish and Breton origin, as well as a surname of Occitan origin. It is common in the English-speaking world. It is possible that the name is derived from an Old Celtic word me ...
and colleagues in 2014. In the analysis, Chaoyangopteridae was found to consist of the genus ''
Eoazhdarcho ''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 ...
'' 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 Tapejaridae (from a Tupi word meaning "the old being") are a family of pterodactyloid pterosaurs from the Cretaceous period. Members are currently known from Brazil, England, Hungary, Morocco, Spain, the United States, and China. The most primiti ...
, 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''. A 2021 study focused on ''
Aerotitan ''Aerotitan'' is a genus of large azhdarchid pterosaur known from the Late Cretaceous period (Maastrichtian stage) of what is now the Allen Formation of the Neuquén Basin in northern Patagonia, Argentina. Discovery and naming The type species ...
'' recovers '' Xericeps'' and ''
Argentinadraco ''Argentinadraco'' (meaning "Argentina dragon") 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 Jo ...
'' as sister taxa within Chaoyangopteridae.


Paleobiology


Diet

Like their azhdarchid relatives, pterosaurs that belong to the Chaoyangopteridae were terrestrial
predators Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
.


Paleoecology

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 ''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 Discovery a ...
''. ''
Microtuban ''Microtuban'' is an extinct genus of azhdarchoid pterosaur known from the Late Cretaceous of northern Lebanon. Discovery ''Microtuban'' is known only from a partially preserved skeleton lacking the skull, holotype SMNK PAL 6595. It was acquire ...
'' may extend the clade's existence into the early
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
.Wilton, Mark P. (2013). Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press. .


References

{{Taxonbar, from=Q2951218 Early Cretaceous pterosaurs Early Cretaceous first appearances Early Cretaceous extinctions Chaoyang, Liaoning Paleontology in Liaoning Prehistoric reptile families