Ornithocheirae

Ornithocheirae is an extinct clade of pteranodontoid pterosaurs from the Early Cretaceous to the Late Cretaceous (Valanginian to Turonian stages) of Asia, Europe, North America and South America.^[1] It was named by Harry Seeley in 1870 as a family that contains Ornithocheirus and its relatives. The name was emended to Ornithocheiridae, to match the requirements of the ICZN Code that a family-ranked clade should end with an -idae suffix. Brian Andres (2010) in his review of pterosaur phylogeny, defined the name Ornithocheirae phylogenetically, as a node-based taxon consisting of the last common ancestor of Anhanguera and Ornithocheirus and all its descendants.^[2] Thus Ornithocheirae is defined to include two families, the Anhangueridae and the Ornithocheiridae, following the opinion of Alexander Kellner^[3] and Andres that these families should not be synonymized based on their original phylogenetical definitions.^[2] However, subsequent studies in 2019 have found Ornithocheirae to be a more inclusive group containing both Anhangueria and Targaryendraconia.

Ornithocheirans Temporal range: Valanginian-Turonian, ~140–92.5 Ma PreꞒ Ꞓ O S D C P T J K Pg N
Skeletal cast of the species Maaradactylus spielbergi
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Order:	†Pterosauria
Suborder:	†Pterodactyloidea
Clade:	†Ornithocheiriformes
Clade:	†Ornithocheirae Seeley, 1870
Subgroups
†Anhangueria †Ornithocheiridae †Targaryendraconia

Classification

Holotypes of Ornithocheirus (A and C) and Tropeognathus (B and D)

Holotype snout of Uktenadactylus

The cladogram below is a topology recovered by Jacobs et al. (2019). Their analysis is similar to the one by Andres & Myers in 2013, though they placed more genera within the families Ornithocheiridae and Anhangueridae, including Camposipterus, Cimoliopterus, Maaradactylus and Siroccopteryx.^[4]

Ornithocheirae

Anhangueridae  Maaradactylus kellneri Maaradactylus spielbergi Anhanguera piscator Anhanguera robustus Anhanguera araripensis  Ornithocheiridae  Coloborhynchus clavirostris Uktenadactylus wadleighi Coloborhynchus reedi Coloborhynchus capito Coloborhynchus fluviferox Coloborhynchus NHMUK R481 Siroccopteryx moroccensis Ornithocheirus simus Tropeognathus mesembrinus Camposipterus segwickii Camposipterus colorhinus Camposipterus nasutus Cimoliopterus cuvieri Cimoliopterus dunni

Many recently subsequent studies have used the term Ornithocheirae for a more inclusive group, which in turn contains the smaller groups Anhangueria and Targaryendraconia.^[5][6][7][8] The cladogram below shows a specific study made by Holgado and Pêgas in 2020.^[9]

Ornithocheirae	Ornithocheirus Targaryendraconia Targaryendraconidae Cimoliopteridae Anhangueria Hamipteridae Hamipterus Iberodactylus Anhangueridae Tropeognathinae Siroccopteryx Tropeognathus Mythunga Ferrodraco Coloborhynchinae Aerodraco Coloborhynchus Nicorhynchus Uktenadactylus Anhanguerinae Caulkicephalus Guidraco Ludodactylus Liaoningopterus Cearadactylus Maaradactylus Anhanguera

References

1. Richard J. Butler; Stephen L. Brusatte; Brian B. Andres; Roger B. J. Benson (2012). "How do geological sampling biases affect studies of morphological evolution in deep time? A case study of the Pterosauria (Reptilia: Archosauria)". Evolution. 66 (1): 147–162. doi:10.1111/j.1558-5646.2011.01415.x. PMID 22220871. S2CID 205783384.
2. Andres, Brian Blake (2010). Systematics of the Pterosauria. Yale University. p. 366. A preview that shows the cladogram without clade names
3. Kellner, A. W. A., (2003): Pterosaur phylogeny and comments on the evolutionary history of the group. pp. 105-137. — in Buffetaut, E. & Mazin, J.-M., (eds.): Evolution and Palaeobiology of Pterosaurs. Geological Society of London, Special Publications 217, London, 1-347
4. Jacobs, M.L.; Martill, D.M.; Ibrahim, N.; Longrich, N. (2019). "A new species of Coloborhynchus (Pterosauria, Ornithocheiridae) from the mid-Cretaceous of North Africa" (PDF). Cretaceous Research. 95: 77–88. doi:10.1016/j.cretres.2018.10.018. S2CID 134439172.
5. 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
6. 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.
7. Pêgas, R.V., Holgado, B., Leal, M.E.C., 2019. "Targaryendraco wiedenrothi gen. nov. (Pterodactyloidea, Pteranodontoidea, Lanceodontia) and recognition of a new cosmopolitan lineage of Cretaceous toothed pterodactyloids", Historical Biology, 1–15. doi:10.1080/08912963.2019.1690482
8. Zhou X., Pêgas R.V., Leal M.E.C. & Bonde N. 2019. "Nurhachius luei, a new istiodactylid pterosaur (Pterosauria, Pterodactyloidea) from the Early Cretaceous Jiufotang Formation of Chaoyang City, Liaoning Province (China) and comments on the Istiodactylidae." PeerJ 7:e7688
9. Holgado, B.; Pêgas, R.V. (2020). "A taxonomic and phylogenetic review of the anhanguerid pterosaur group Coloborhynchinae and the new clade Tropeognathinae". Acta Palaeontologica Polonica. 65. doi:10.4202/app.00751.2020.