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Azhdarchoidea (or azhdarchoids) is a group of pterosaurs within the suborder
Pterodactyloidea Pterodactyloidea (derived from the Greek words ''πτερόν'' (''pterón'', for usual ''ptéryx'') "wing", and ''δάκτυλος'' (''dáktylos'') "finger" meaning "winged finger", "wing-finger" or "finger-wing") is one of the two traditional ...
, more specifically within the group
Ornithocheiroidea Ornithocheiroidea (or ornithocheiroids) is a group of pterosaurs within the extinct suborder Pterodactyloidea. They were typically large pterosaurs that lived from the Early to Late Cretaceous periods (Valanginian to Maastrichtian stages), with fo ...
. Pterosaurs belonging to this group lived throughout the Early and
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'', ...
periods, with one tentative member, ''
Tendaguripterus ''Tendaguripterus'' was a genus of pterodactyloid pterosaur from the Kimmeridgian to Tithonian-age Upper Jurassic Middle Saurian Beds (Tendaguru Formation) of Tendaguru, Lindi Region, Tanzania. Discovery and naming During the German paleontolog ...
'', that lived in the
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
period. The largest azhdarchoids include members of the family Azhdarchidae, examples of these are '' Quetzalcoatlus'', '' Hatzegopteryx'', and ''
Arambourgiania ''Arambourgiania'' is an extinct genus of azhdarchid pterosaur from the Late Cretaceous period (Maastrichtian stage) of Jordan, and possibly the United States.Harrell, T. Lynn Jr.; Gibson, Michael A.; Langston, Wann Jr. (2016). "A cervical ver ...
''. The Azhdarchoidea has been recovered as either closely related to the
Ctenochasmatoidea Ctenochasmatoidea is a group of early pterosaurs within the suborder Pterodactyloidea. Their remains are usually found in what were once coastal or lake environments. They generally had long wings, long necks, and highly specialized teeth. Evol ...
, as the sister taxon of the
Pteranodontoidea Pteranodontoidea (or pteranodontoids, from Greek meaning "toothless wings") is an extinct clade of ornithocheiroid pterosaurs from the Early to Late Cretaceous (early Valanginian to late Maastrichtian stages) of Asia, Africa, Europe, North Americ ...
within the Ornithocheiroidea, or within the
Tapejaroidea Tapejaroidea (or tapejaroids) 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 Ju ...
, which in turn was also within the Ornithocheiroidea.


Classification

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.Unwin, D. M., (2003). "On the phylogeny and evolutionary history of pterosaurs." Pp. 139-190. in Buffetaut, E. & Mazin, J.-M., (eds.) (2003). ''Evolution and Palaeobiology of Pterosaurs''. Geological Society of London, Special Publications 217, London, 1-347. There have been several competing views of azhdarchoid relationships. The first, presented by Felipe Pinheiro and colleagues in 2011, considered the
tapejarid 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 primit ...
s to be a monophyletic clade including the thalassodromines and chaoyangopterines.Pinheiro, F.L., Fortier, D.C., Schultz, C.L., De Andrade, J.A.F.G. and Bantim, R.A.M. (in press). "New information on ''Tupandactylus imperator'', with comments on the relationships of Tapejaridae (Pterosauria)." ''Acta Palaeontologica Polonica'', in press, available online 03 Jan 2011. 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
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 Cre ...
s. All azhdarchoids which are part of 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 Alexander Wilhelm Armin Kellner (born September 26, 1961) is a Brazilian geologist and paleontologist who is a leading expert in the field of studying pterosaurs. His research has focused mainly on fossil reptiles from the Cretaceous Period, i ...
created a different clade called
Tapejaroidea Tapejaroidea (or tapejaroids) 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 Ju ...
, which he defined as the most recent common ancestor and all descendants of ''Tapejara'', ''Quetzalcoatlus'', and ''
Dsungaripterus ''Dsungaripterus'' is a genus of dsungaripterid pterosaur with an average wingspan of . ''Dsungaripterus'' lived during the Early Cretaceous in what is now China, and its first fossil was found in the Junggar Basin. Description ''Dsungaripte ...
''. Kellner created this clade to include both Azhdarchoidea and the family
Dsungaripteridae Dsungaripteridae is a group of pterosaurs within the suborder Pterodactyloidea. They were robust pterosaurs with good terrestrial abilities and flight honed for inland settings. Classification In 1964 Young created a family to place the recently ...
, but as separate groups. A lot of recent studies have followed this concept.Borja Holgado, Rodrigo V. Pêgas, José Ignacio Canudo, Josep Fortuny, Taissa Rodrigues, Julio Company & Alexander W.A. Kellner, 2019, "On a new crested pterodactyloid from the Early Cretaceous of the Iberian Peninsula and the radiation of the clade Anhangueria", ''Scientific Reports'' 9: 4940 There are competing theories of azhdarchoid phylogeny; it is either recovered as closely related to the clade
Ctenochasmatoidea Ctenochasmatoidea is a group of early pterosaurs within the suborder Pterodactyloidea. Their remains are usually found in what were once coastal or lake environments. They generally had long wings, long necks, and highly specialized teeth. Evol ...
, or within the group
Ornithocheiroidea Ornithocheiroidea (or ornithocheiroids) is a group of pterosaurs within the extinct suborder Pterodactyloidea. They were typically large pterosaurs that lived from the Early to Late Cretaceous periods (Valanginian to Maastrichtian stages), with fo ...
, either as the sister taxon of the
Pteranodontoidea Pteranodontoidea (or pteranodontoids, from Greek meaning "toothless wings") is an extinct clade of ornithocheiroid pterosaurs from the Early to Late Cretaceous (early Valanginian to late Maastrichtian stages) of Asia, Africa, Europe, North Americ ...
or within the clade Tapejaroidea. The latter two of which are more widely accepted. 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 ...
showing the results of a
phylogenetic 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 ...
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. The result of 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. In 2022, Pêgas ''et al.'' named and officially registered two new clades: Azhdarchomorpha, the most inclusive clade containing ''Azhdarcho'' but not ''Tapejara'' or ''Thalassodromeus'', and Alanqidae, containing ''Alanqa'' but not ''Chaoyangopterus'' or ''Azhdarcho''. Their phylogeny is shown below:


References

{{Taxonbar, from=Q137513 Tithonian first appearances Maastrichtian extinctions