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 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.

Azhdarchoids Temporal range: Early - Late Cretaceous, 143–66 Ma PreꞒ Ꞓ O S D C P T J K Pg N Possible Late Jurassic record[1]
Reconstructed skeleton of Quetzalcoatlus northropi
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Order:	†Pterosauria
Suborder:	†Pterodactyloidea
Clade:	†Ornithocheiroidea
Clade:	†Azhdarchoidea Nesov, 1984
Subgroups
†Inabtanin[2] †Neoazhdarchia †Tapejaromorpha †Azhdarchomorpha †Cretornis? †Ornithostoma? †Dsungaripteromorpha? †Microtuban? †Dsungaripteridae †Thalassodromidae? †Alanqidae? †Alanqa †Argentinadraco? †Keresdrakon? †Leptostomia? †Xericeps? †Neopterodactyloidea †Eoazhdarcho †Chaoyangopteridae †Azhdarchiformes †Radiodactylus †Montanazhdarcho †Azhdarchidae

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.^[3]

There have been several competing views of azhdarchoid relationships. The first, presented by Felipe Pinheiro and colleagues in 2011, considered the tapejarids to be a monophyletic clade including the thalassodromines and chaoyangopterines.^[4] 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.^[5] 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.^[3]

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 the family Dsungaripteridae, but as separate groups.^[6][7] A lot of recent studies have followed this concept.^{[8][9][10][11]}

There are competing theories of azhdarchoid phylogeny; it is either recovered as closely related to the clade Ctenochasmatoidea,^[3][12] 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.^{[13][9][11][10]} Below 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.^[13]

Azhdarchoidea

Tapejaromorpha Bennettazhia oregonensis Eopteranodon lii "Sinopterus" gui Nemicolopterus crypticus Huaxiapterus jii Tapejaridae Neoazhdarchia Dsungaripteromorpha Dsungaripteridae Dsungaripterinae Dsungaripterus weii Domeykodactylus ceciliae Noripterinae Noripterus parvus Noripterus complicidens Thalassodrominae Thalassodromeus sethi Tupuxuara longicristatus Tupuxuara leonardii Neopterodactyloidea Chaoyangopteridae Eoazhdarcho liaoxiensis Chaoyangopterinae Shenzhoupterus chaoyangensis Chaoyangopterus zhangi Jidapterus edentus Azhdarchidae Azhdarcho lancicollis Quetzalcoatlinae Zhejiangopterus linhaiensis Arambourgiania philadelphiae Quetzalcoatlus northropi

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.^[11]

Tapejaroidea

Dsungaripteridae Dsungaripterus weii Noripterus parvus Azhdarchoidea Azhdarchidae Azhdarcho lancicollis Quetzalcoatlus northropi Zhejiangopterus linhaiensis Chaoyangopteridae Chaoyangopterus zhangi Jidapterus edentus Shenzhoupterus chaoyangensis Tapejaromorpha Keresdrakon vilsoni Tapejaridae Thalassodrominae Thalassodromeus sethi Tupuxuara leonardii Tapejarinae Caupedactylus ybaka Aymberedactylus cearensis Eopteranodon lii "Huaxiapterus" benxiensis "Huaxiapterus" corollatus Sinopterus dongi Tapejarini Europejara olcadesorum Caiuajara dobruskii Tapejara wellnhoferi Tupandactylus imperator

In 2022, Pêgas et al.^[14] 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 original concept of Alanqidae was later refuted and updated by Pêgas^[15]. The updated phylogeny of Pêgas (2024)^[15] is shown below:

Tapejaroidea

Dsungaripteridae Azhdarchoidea Tapejaromorpha Thalassodromidae Tupuxuara Kariridraco Thalassodromeus oberlii Thalassodromeus sethi Tapejariformes Caupedactylia Aymberedactylus Caupedactylus Tapejaridae Sinopterinae Wightia Afrotapejara Bakonydraco Huaxiadraco Eopteranodon Sinopterus Tapejarinae Europejara Tupandactylus Tapejara Caiuajara Torukjara Azhdarchomorpha Keresdrakon Chaoyangopteridae Azhdarchiformes Alanqidae Alanqa Argentinadraco Xericeps Leptostomia Azhdarchidae

References

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: 121–134.
2. Rosenbach, K. L.; Goodvin, D. M.; Albshysh, M. G.; Azzam, H. A.; Smadi, A. A.; Mustafa, H. A.; Zalmout, I. S. A.; Wilson Mantilla, J. A. (2024). "New pterosaur remains from the Late Cretaceous of Afro-Arabia provide insight into flight capacity of large pterosaurs". Journal of Vertebrate Paleontology. e2385068. doi:10.1080/02724634.2024.2385068.
3. 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.
4. 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. doi:10.4202/app.2010.0057
5. 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.
6. Kellner, A.W.A. (1996). "Description of new material of Tapejaridae and Anhangueridae (Pterosauria, Pterodactyloidea) and discussion of pterosaur phylogeny". Columbia University.
7. Buffetaut, Eric; Mazin, Jean-Michel (2003). Evolution and Palaeobiology of pterosaurs. Geological Society of London. ISBN 9781862391437.
8. Vidovic, S. U.; Martill, D. M. (2014). "Pterodactylus scolopaciceps Meyer, 1860 (Pterosauria, Pterodactyloidea) from the Upper Jurassic of Bavaria, Germany: The Problem of Cryptic Pterosaur Taxa in Early Ontogeny". PLOS ONE. 9 (10): e110646. Bibcode:2014PLoSO...9k0646V. doi:10.1371/journal.pone.0110646. PMC 4206445. PMID 25337830.
9. 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 doi:10.1038/s41598-019-41280-4
10. Kellner, 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.
11. Kellner, Alexander W. A.; Weinschütz, Luiz C.; Holgado, Borja; Bantim, Renan A. M.; Sayão, Juliana M. (19 August 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.
12. Bestwick, J.; Unwin, D.M.; Butler, R.J.; Henderson, D.M.; Purnell, M.A. (2018). "Pterosaur dietary hypotheses: a review of ideas and approaches". Biological Reviews. 93 (4): 2021–2048. doi:10.1111/brv.12431. PMC 6849529. PMID 29877021.
13. Andres, B.; Clark, J.; Xu, X. (2014). "The Earliest Pterodactyloid and the Origin of the Group". Current Biology. 24 (9): 1011–6. doi:10.1016/j.cub.2014.03.030. PMID 24768054.
14. Pêgas, R.V.; Holgado, B.; Ortiz David, L.D.; Baiano, M.A.; Costa, F.R. (2022). "On the pterosaur Aerotitan sudamericanus (Neuquén Basin, Upper Cretaceous of Argentina), with comments on azhdarchoid phylogeny and jaw anatomy". Cretaceous Research. 129: Article 104998. doi:10.1016/j.cretres.2021.104998. ISSN 0195-6671. S2CID 238725853.
15. Pêgas, Rodrigo V. (2024-06-10). "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. ISSN 0891-2963.