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Azhdarchoidea

From Wikipedia, the free encyclopedia
Superfamily of pterodactyloid pterosaurs

Azhdarchoids
Temporal range:
Early -Late Cretaceous,143–66 MaPossibleLate Jurassic record[1]
Four azhdarchoids (clockwise from top left):Quetzalcoatlus,Tapejara,Tupuxuara, andMeilifeilong
Scientific classificationEdit this classification
Kingdom:Animalia
Phylum:Chordata
Class:Reptilia
Order:Pterosauria
Suborder:Pterodactyloidea
Superfamily:Ornithocheiroidea
Clade:Azhdarchoidea
Unwin, 1995
Subgroups

Azhdarchoidea (/æʒdɑːrˈkɔɪdɪːə/, meaning "azhdarchid-like forms") is a group ofpterosaurs within the suborderPterodactyloidea. Pterosaurs belonging to this group lived throughout theEarly andLate Cretaceous periods, with one tentative member,Tendaguripterus, that lived in theLate Jurassic period. Remains of this group have been found in the Americas, Africa, and Eurasia, suggesting that they probably had a global distribution.

Azhdarchoids are generally distinguished from other pterodactyloids by their relatively low arm-to-leg-length ratio, suggesting that they were more proficient in moving on the ground than pterosaurs likePteranodon orAnhanguera (which had very long arms relative to the length of their legs). This has led some researchers to suggest that many azhdarchoids, such as the azhdarchids and dsungaripterids, may have been primarily terrestrial, while retaining the ability to fly when necessary.[2]

Classification

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Azhdarchoidea was named by paleontologist David Unwin in 1995.[3] He later gave the clade a phylogenetic definition in 2003. He defined the group as the most recent common ancestor ofQuetzalcoatlus andTapejara, and all its descendants.[4]

Relationships

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Azhdarchoidea is generally considered as one of three major clades of pterodactyloids. The other two major groups areArchaeopterodactyloidea andPteranodontoidea. In some older classification schemes, azhdarchoids were considered to be closely related toctenochasmatoids likePterodactylus andCtenochasma. However, most researchers no longer consider this arrangement to be very likely. Instead, azhdarchoids are grouped as close relatives of pteranodontoids under the cladeOrnithocheiroidea, which would comprise the bulk of pterodactyloid diversity.

A simplified cladogram illustrating the relationships between azhdarchoids and other pterosaurs based on the analysis of Brian Andres,James Clark, andXu Xing in 2014 is shown below.[5]

Monofenestrata

Taxonomy

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Kariridraco, a member of the disputed familyThalassodromidae

Azhdarchoidea is generally believed to be composed of several principlefamilies:Azhdarchidae,Chaoyangopteridae,Tapejaridae,Thalassodromidae, and potentiallyDsungaripteridae, although Thalassodromidae of these is considered to be a lineage tapejarids by some researchers. However, the interrelationships of these groups is not confidently understood. The azhdarchids and chaoyangopterids are generally considered to be one another's closest relatives, but the affinities of tapejarids, thalassodromids, and dsungaripterids are controversial.[6] Some researchers also consider an additional grouping,Alanqidae, composed of taxa formerly considered azhdarchids.[7]

There are several competing views of azhdarchoid relationships. An early study presented by paleontologist Felipe Pinheiro and colleagues in 2011 considered thetapejarids to be amonophyletic clade including the thalassodromines and the chaoyangopterids (therein called "chaoyangopterines").[8] Other earlier studies, such as that of paleontologistsDarren Naish &David Martill in 2006, and that ofLü Junchang and colleagues in 2008, considered the traditional "tapejarids" to be aparaphyletic grade of primitive azhdarchoids, with true tapejarids more basal, and the thalassodromines (alternatively called thalassodromids) and chaoyangopterids being successively more closely related toazhdarchids.[9][10]

Life reconstructions ofHatzegopteryx (top) andArambourgiania (bottom), giant members of the familyAzhdarchidae

Generally speaking, in the modern scientific literature, there are two major competing hypotheses regarding the classification of azhdarchoids. The first of these is based on the work of Brian Andres and colleagues, who have published multiple papers on the taxaKryptodrakon,Elanodactylus, andQuetzalcoatlus among others, in which they examined the phylogenetic relationships of these groups.[5][11] Andres and colleagues have generally suggested thatDsungaripteridae andThalassodromidae are closely related and that they form a clade (Dsungaripteromorpha) which is more closely related toAzhdarchidae thanTapejaridae.[5] This result has been corroborated by some other authors.[12] A simplified version of Andres and colleagues' phylogeny is shown below.[5]

Ornithocheiroidea
Tapejarids, one of the prominent azhdarchoid families.Sinopterus (left) andHuaxiadraco (right) are pictured

The second hypothesis of azhdarchoid relationships is that Dsungaripteridae is thesister taxon of Azhdarchoidea within the larger clade Tapejaroidea. This hypothesis has been published on by R. Pêgas and colleagues in their extensive studies of pterosaurs likeSinopterus,Thalassodromeus, andAerotitan.[7][13][14][15] This hypothesis has also received support from other authors.[16][17] A simplified version of their phylogeny is shown below.[7]

Ornithocheiroidea

In 2025, Henry N. Thomas and Skye N. McDavid performed an extensive revision of pterosaur phylogenetics with a focus on Azhdarchomorpha. Their results align with a previous analysis by Pêgas et al., (2024)[15] findingTapejaroidea to be monophyletic containingDsungaripteromorpha and Azhdarchoidea, as well asTapejaromorpha andAzhdarchomorpha being sister taxa within Azhdarchoidea.[18]A simplified version of their results is shown below.

Ornithocheiroidea

Subclades

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A wide variety of subclades have been defined within Azhdarchoidea in the last 15 years. Many of these have contradictory or overlapping definitions, and so whether or not these clades represent truemonophyletic groups remains hotly debated. Some of the uncertainty regards the position ofThalassodromidae, which may be closely related to eitherTapejaridae orDsungaripteridae. Another source of uncertainty is the affinities of the dsungaripterids themselves. This problematic group has been found to be closely related toOrnithocheiromorpha,[19] just outside of Azhdarchoidea,[20][16][17][21] or within Azhdarchoidea.[5][22]

NameNamed byDefinitionNotes
AzhdarchiformesAndres, 2021[23]Most-inclusive clade containingQuetzalcoatlus, but notChaoyangopterus
AzhdarchomorphaPêgas et al., 2021[7]Most-inclusive clade containingAzhdarcho, but notTapejara orThalassodromeusMay be synonymous with Neopterodactyloidea ifThalassodromeus is a member ofTapejaridae
ConcilazhiaThomas & McDavid, 2025[18]Least-inclusive clade containingAzhdarcho,Chaoyangopterus, andAlanqaSimilar definition to 'Neopterodactyloidea' but usesAzhdarcho as an internal specifier rather thanQuetzalcoatlus
DsungaripteromorphaAndres et al., 2014[5]Most-inclusive clade containingDsungaripterus but notQuetzalcoatlusMay be synonymous withDsungaripteridae if Dsungaripteridae is outside of Azhdarchoidea
NeoazhdarchiaUnwin, 2003[4]Least-inclusive clade containing bothAzhdarcho andTupuxuaraMay be synonymous with Azhdarchoidea ifTupuxuara belongs toTapejaridae; has alternately been used interchangeably with Azhdarchomorpha[6]
NeopterodactyloideaAndres et al., 2014[5]Least inclusive clade containing bothAzhdarcho andChaoyangopterusMay be used interchangeably with Azhdarchomorpha ifChaoyangopterus is its most basal member; however, under thePhyloCode, this clade is improperly named, as it is derived fromPterodactylus despite not including the genus in the clade
TapejariformesPêgas et al., 2024[15]The clade characterized by a downturned rostrum synapomorphic with that ofTapejaraMay be synonymous withTapejaridae ifCaupedactylus is a tapejarid
TapejaroideaKellner, 1996[24]Least-inclusive clade containingQuetzalcoatlus,Tapejara, andDsungaripterusMay be synonymous with Azhdarchoidea ifDsungaripterus is more closely related toQuetzalcoatlus than either are toTapejara; alternatively used for a clade containingTapejaridae andThalassodromidae[25]
TapejaromorphaAndres et al., 2014[5]Most inclusive clade containingTapejara but notAzhdarcho

See also

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References

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  1. ^Unwin, David M.; Heinrich, Wolf-Dieter (1999)."On a pterosaur jaw from the Upper Jurassic of Tendaguru (Tanzania)".Mitteilungen aus dem Museum für Naturkunde in Berlin, Geowissenschaftliche Reihe.2 (1):121–134.Bibcode:1999FossR...2..121U.doi:10.1002/mmng.1999.4860020109.
  2. ^Smyth, Robert S.H.; Breithaupt, Brent H.; Butler, Richard J.; Falkingham, Peter L.; Unwin, David M. (2024)."Hand and foot morphology maps invasion of terrestrial environments by pterosaurs in the mid-Mesozoic".Current Biology.34 (21): 4894–4907.e3.Bibcode:2024CBio...34.4894S.doi:10.1016/j.cub.2024.09.014.PMID 39368469.
  3. ^Unwin, D. M. 1995. Preliminary results of a phylogenetic analysis of the Pterosauria (Diapsida: Archosauria); pp. 69–72 in A. Sun and Y. Wang (eds.), Sixth Symposium on Mesozoic Terrestrial Ecosystems and Biota, Short Papers. China Ocean Press, Beijing.
  4. ^abUnwin, 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.
  5. ^abcdefghAndres, B.; Clark, J.; Xu, X. (2014)."The Earliest Pterodactyloid and the Origin of the Group".Current Biology.24 (9):1011–6.Bibcode:2014CBio...24.1011A.doi:10.1016/j.cub.2014.03.030.PMID 24768054.
  6. ^abWitton, M. P. (2013).Pterosaurs: Natural History, Evolution, Anatomy (1st ed.). Princeton University Press.ISBN 978-0-691-15061-1.
  7. ^abcdPêgas, Rodrigo V.; Holgado, Borja; Ortiz David, Leonardo D.; Baiano, Mattia A.; Costa, Fabiana R. (2022). "On the pterosaur Aerotitan sudamericanus (Neuquén Basin, Upper Cretaceous of Argentina), with comments on azhdarchoid phylogeny and jaw anatomy".Cretaceous Research.129.Bibcode:2022CrRes.12904998P.doi:10.1016/j.cretres.2021.104998.
  8. ^Pinheiro, F.L., Fortier, D.C., Schultz, C.L., De Andrade, J.A.F.G. and Bantim, R.A.M. (in press). "New information onTupandactylus imperator, with comments on the relationships of Tapejaridae (Pterosauria)."Acta Palaeontologica Polonica, in press, available online 03 Jan 2011.doi:10.4202/app.2010.0057
  9. ^Martill, D. M.; Naish, D. (2006)."Cranial crest development in the Azhdarchoid pterosaurTupuxuara, with a review of the genus and tapejarid monophyly".Palaeontology.49 (4):925–941.Bibcode:2006Palgy..49..925M.doi:10.1111/j.1475-4983.2006.00575.x.S2CID 15609202.
  10. ^Lü, J.; Unwin, D.M.; Xu, L.; Zhang, X. (2008). "A new azhdarchoid pterosaur from the Lower Cretaceous of China and its implications for pterosaur phylogeny and evolution".Naturwissenschaften.95 (9):891–897.Bibcode:2008NW.....95..891L.doi:10.1007/s00114-008-0397-5.PMID 18509616.S2CID 13458087.
  11. ^Andres, Brian; Qiang, JI (2008)."A New Pterosaur from the Liaoning Province of China, the Phylogeny of the Pterodactyloidea, and Convergence in Their Cervical Vertebrae"(PDF).Palaeontology.51 (2):453–469.Bibcode:2008Palgy..51..453A.doi:10.1111/j.1475-4983.2008.00761.x.
  12. ^Longrich, Nicholas R.; Martill, David M.; Andres, Brian (2018)."Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary".PLOS Biology.16 (3) e2001663.doi:10.1371/journal.pbio.2001663.PMC 5849296.PMID 29534059.
  13. ^Pêgas, Rodrigo V.; Costa, Fabiana R.; Kellner, Alexander W. A. (2018). "New Information on the osteology and a taxonomic revision of the genusThalassodromeus (Pterodactyloidea, Tapejaridae, Thalassodrominae)".Journal of Vertebrate Paleontology.38 (2) e1443273.Bibcode:2018JVPal..38E3273P.doi:10.1080/02724634.2018.1443273.
  14. ^Pêgas, Rodrigo V.; Zhou, Xuanyu; Jin, Xingsheng; Wang, Kai; Ma, Waisum (2023)."A taxonomic revision of theSinopterus complex (Pterosauria, Tapejaridae) from the Early Cretaceous Jehol Biota, with the new genusHuaxiadraco".PeerJ.11 e14829.doi:10.7717/peerj.14829.PMC 9922500.PMID 36788812.
  15. ^abcPêgas, Rodrigo V. (2024). "A taxonomic note on the tapejarid pterosaurs from the Pterosaur Graveyard site (Caiuá Group, ?Early Cretaceous of Southern Brazil): Evidence for the presence of two species".Historical Biology:1–22.doi:10.1080/08912963.2024.2355664.
  16. ^abKellner, Alexander W. A.; Caldwell, Michael W.; Holgado, Borja; Vecchia, Fabio M. Dalla; Nohra, Roy; Sayão, Juliana M.; Currie, Philip J. (2019)."First complete pterosaur from the Afro-Arabian continent: insight into pterodactyloid diversity".Scientific Reports.9 (1): 17875.Bibcode:2019NatSR...917875K.doi:10.1038/s41598-019-54042-z.PMC 6884559.PMID 31784545.
  17. ^abBorja Holgado, R. 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 Reports9: 4940doi:10.1038/s41598-019-41280-4
  18. ^abThomas, Henry N.; McDavid, Skye N. (5 Nov 2025)."Enter the dragons: the phylogeny of Azhdarchoidea (Pterosauria: Pterodactyloidea) and the evolution of giant size in pterosaurs".Journal of Systematic Palaeontology.23 (1) 2569368.doi:10.1080/14772019.2025.2569368. Retrieved5 November 2025.
  19. ^Vidovic, S.U. and Martill, D.M. (2017). The taxonomy and phylogeny ofDiopecephalus kochi (Wagner, 1837) and "Germanodactylus rhamphastinus" (Wagner, 1851). In Hone, D. W. E., Witton, M. P. &Martill, D. M. (eds)New Perspectives on Pterosaur Palaeobiology. Geological Society, London, Special Publications, 455doi:10.1144/SP455.12
  20. ^Kellner, Alexander W. A.; Weinschütz, Luiz C.; Holgado, Borja; Bantim, Renan A. M.; Sayão, Juliana M. (August 19, 2019)."A new toothless pterosaur (Pterodactyloidea) from Southern Brazil with insights into the paleoecology of a Cretaceous desert".Anais da Academia Brasileira de Ciências.91 (suppl 2) e20190768.doi:10.1590/0001-3765201920190768.ISSN 0001-3765.PMID 31432888.
  21. ^Jiang, Shun-Xing; Zhang, Xin-Jun; Cheng, Xin; Wang, Xiao-Lin (2020)."A new pteranodontoid pterosaur forelimb from the upper Yixian Formation, with a revision ofYixianopterus jingangshanensis"(PDF).Vertebrata PalAsiatica.doi:10.19615/j.cnki.1000-3118.201124.
  22. ^Cerqueira, Gabriela; Santos, Mateus; Marks, Maikon; Sayão, Juliana; Pinheiro, Felipe (2021)."A new pterosaur species from the Lower Cretaceous of Brazil and the paleobiogeography of the Tapejaridae (Azhdarchoidea)".Acta Palaeontologica Polonica.66.doi:10.4202/app.00848.2020.
  23. ^Andres, B.; Langston, W. Jr. (2021)."Morphology and taxonomy ofQuetzalcoatlus Lawson 1975 (Pterodactyloidea: Azhdarchoidea)".Journal of Vertebrate Paleontology.41 (sup1): 142.Bibcode:2021JVPal..41S..46A.doi:10.1080/02724634.2021.1907587.ISSN 0272-4634.S2CID 245125409.
  24. ^Kellner, A.W.A. (1996). "Description of new material of Tapejaridae and Anhangueridae (Pterosauria, Pterodactyloidea) and discussion of pterosaur phylogeny".Columbia University.
  25. ^Jaime A. Headden and Hebert B.N. Campos (2014). "An unusual edentulous pterosaur from the Early Cretaceous Romualdo Formation of Brazil".Historical Biology: An International Journal of Paleobiology.27 (7):815–826.doi:10.1080/08912963.2014.904302.S2CID 129306469.
Avemetatarsalia
Pterosauria
    • see below↓
Preondactylia
Caviramidae?
Austriadraconidae
Raeticodactylidae
Eudimorphodontidae
Dimorphodontidae
Campylognathoididae
Rhamphorhynchidae
Scaphognathidae?
Pterodactylomorpha
    • see below↓
Campylognathoides liasicus

Scaphognathus crassirostris

Dorygnathus banthensis
Darwinoptera
Wukongopteridae
Anurognathidae
Pterodactyloidea
Lophocratia
    • see below↓
Jeholopterus ninchengensisKryptodrakon progenitor
Germanodactylidae
Gallodactylidae
Aurorazhdarchia
Aurorazhdarchidae
Ctenochasmatidae
Eupterodactyloidea
Ornithocheiroidea
    • see below↓
Pterodactylus antiquusPlataleorhynchus streptorophorodon
Dsungaripteridae
Thalassodromidae?
Tapejaridae
Dsungaripteromorpha?
Chaoyangopteridae
Azhdarchiformes
Alanqidae?
Azhdarchidae
Pteranodontoidea
    • see below↓
Bakonydraco galaczi

Tupandactylus imperator

Quetzalcoatlus
Pteranodontia
Pteranodontidae
Nyctosauromorpha
Aponyctosauria
Nyctosauridae
Lonchodectidae
Istiodactyliformes
Mimodactylidae
Istiodactylidae
Boreopteridae
Ornithocheiridae
Targaryendraconia?
Cimoliopteridae
Targaryendraconidae
Hamipteridae?
Anhangueridae
Pteranodon longiceps

Nyctosaurus gracilis

Ludodactylus sibbicki
Azhdarchoidea
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