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Neognaths; Temporal range: Late Cretaceous – present, 72–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N Possible early Late Cretaceous origin based on molecular clock
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Aves
Infraclass:	Neognathae; Pycraft, 1900
Subgroups
	Pangalloanserae; Neoaves; †Odontopterygiformes;

Neognathae (/niˈɒɡnəθiː/; from Ancient Greek νέος (néos) 'new, young' and γνάθος (gnáthos) 'jaw') is an infraclass of birds, called neognaths, within the class Aves of the clade Archosauria. Neognathae includes the majority of living birds; the exceptions being the tinamous and the flightless ratites, which belong instead to the sister taxon Palaeognathae. There are nearly 10,000 living species of neognaths.

The earliest fossils are known from the very end of the Cretaceous with the oldest known member being Teviornis from the Nemegt Formation of Mongolia,^[6] but molecular clocks suggest that neognaths originated sometime in the first half of the Late Cretaceous, about 90 million years ago.^[7] Since then, they have undergone adaptive radiation, producing the diversity of form, function, and behavior that exists today. Neognathae includes the order Passeriformes (perching birds), one of the largest orders of land vertebrates, containing some 60% of living birds. Passeriformes is twice as species-rich as Rodentia and about five times as species-rich as Chiroptera (bats), which are the two largest orders of mammals. Neognathae also contains some very small orders, often birds of unclear relationships like the hoatzin.

The neognaths have fused metacarpals, an elongate third finger, and 13 or fewer vertebrae. They differ from the Palaeognathae in features like the structure of their jawbones. Neognathae means "new jaws", but it seems that the supposedly "more ancient" paleognath jaws are among the few apomorphic (more derived) features of the palaeognaths, meaning that the respective jaw structure of these groups is not informative in terms of comparative evolution. However, a neognath-like palate is also present in ornithuran birds like Ichthyornis.^[8]

Taxonomy and systematics

Neognathae was long ranked as a superorder subdivided into orders. Attempts to organise this group further, as in the Conspectus of Charles Lucien Bonaparte, were never accepted by a significant majority of ornithologists. Until the 1980s, there was little subdivision of the Aves in general, and even less of phylogenetic merit. Since then, the availability of massive amounts of new data from fossils (especially Enantiornithes and other Mesozoic birds) and molecular (DNA and protein) sequences allowed scientists to refine the classification. With new groups of neognath orders being verified, the taxonomic rank of the group needed to shift. Most researchers have now employed the unranked taxa of phylogenetic nomenclature.^[9]

Neognathae is now universally accepted to subdivide into two lineages, the "fowl" clade Galloanseres and the Neoaves (sometimes called "higher neognaths"). The formal PhyloCode definition given to Neognathae by George Sangster and colleagues in 2022 is "the least inclusive crown clade containing Gallus gallus and Passer domesticus".^[10]

The subdivisions of the latter are still not well resolved, but several monophyletic lineages have been proposed, such as the Mirandornithes, Cypselomorphae, Metaves, and Coronaves. Although groups such as the former two (uniting a few closely related orders) are robustly supported, this cannot be said for the groups Metaves and Coronaves for which there is no material evidence at present, while the Mesozoic record of Neognathae is at present utterly devoid of birds that should have been present if these proposed clades were real.^[11]

Systematics

The orders are arranged in a sequence that attempts to follow the modern view on neognath phylogeny. It differs from the widely used Clements taxonomy as well as from the Sibley-Ahlquist taxonomy, combining those elements from each that more modern research agrees with while updating those that are refuted. Most of the changes affect those "higher landbirds" that are sometimes united as near passerines.^[12]

Neognathia

Feduccia defined the clade Neognathia as birds whose palatal mobility increased due to the following modifications (Feduccia 1980, 1996):

Loss of the Basipterygoid articulation with the cranium.
Development of a pterygoid/palatine joint.
Reduction of the vomer, such that it does not reach caudally to the pterygoid, or is lost entirely.

Relationships

Neognathae cladogram of modern bird relationships based on Stiller et al. (2024).^[13]

Neognathae comprises 39 extant orders and are categorized as followed:

Neognathae

Galloanserae

	Galliformes (chickens (including the red junglefowl), pheasants, quails, turkeys, peafowl, etc.)

	Anseriformes (ducks, geese, and swans)

Neoaves

Mirandornithes

	Phoenicopteriformes (flamingos)

	Podicipediformes (grebes)

Columbaves

Columbimorphae

Columbiformes (pigeons and doves)

Pteroclimesites

	Mesitornithiformes (mesites)

	Pterocliformes (sandgrouse)

Otidimorphae

Cuculiformes (cuckoos)

Musophagotides

	Otidiformes (bustards)

	Musophagiformes (turacos)

Passerea

Elementaves

Gruae

Opisthocomiformes (hoatzins)

Gruimorphae

	Gruiformes (rails and cranes)

	Charadriiformes (waders, gulls, auks and relatives)

Strisores

Phaethoquornithes

Telluraves

Afroaves

Hieraves

	Strigiformes (owls)

	Accipitriformes (vultures, kites, hawks, and eagles)

Coraciimorphae

Coliiformes (mousebirds)

Cavitaves

Leptosomiformes (cuckoo rollers)

Eucavitaves

Trogoniformes (trogons)

Picocoraciae

Bucerotiformes (hornbills, hoopoes and relatives)

Picodynastornithes

	Coraciiformes (kingfishers, rollers, bee-eaters and relatives)

	Piciformes (woodpeckers, toucans and relatives)

Australaves

Cariamiformes (seriemas)

Eufalconimorphae

Falconiformes (falcons and caracaras)

Psittacopasserae

	Psittaciformes (parrots)

	Passeriformes (passerines)

Footnotes

^ Field, Daniel J.; Benito, Juan; Chen, Albert; Jagt, John W. M.; Ksepka, Daniel T. (March 2020). "Late Cretaceous neornithine from Europe illuminates the origins of crown birds". Nature. 579 (7799): 397–401. Bibcode:2020Natur.579..397F. doi:10.1038/s41586-020-2096-0. ISSN 0028-0836. PMID 32188952. S2CID 212937591.
^ De Pietri, Vanesa L.; Scofield, R. Paul; Zelenkov, Nikita; Boles, Walter E.; Worthy, Trevor H. (February 2016). "The unexpected survival of an ancient lineage of anseriform birds into the Neogene of Australia: the youngest record of Presbyornithidae". Royal Society Open Science. 3 (2): 150635. Bibcode:2016RSOS....350635D. doi:10.1098/rsos.150635. PMC 4785986. PMID 26998335.
^ Kuhl., H.; Frankl-Vilches, C.; Bakker, A.; Mayr, G.; Nikolaus, G.; Boerno, S. T.; Klages, S.; Timmermann, B.; Gahr, M. (2020). "An unbiased molecular approach using 3'UTRs resolves the avian family-level tree of life". Molecular Biology and Evolution. 38: 108–127. doi:10.1093/molbev/msaa191. PMC 7783168. PMID 32781465.
^ Yonezawa, T.; Segawa, T.; Mori, H.; Campos, P. F.; Hongoh, Y.; Endo, H.; Akiyoshi, A.; Kohno, N.; Nishida, S.; Wu, J.; Jin, H.; Adachi, J.; Kishino, H.; Kurokawa, K.; Nogi, Y.; Tanabe, H.; Mukoyama, H.; Yoshida, K.; Rasoamiaramanana, A.; Yamagishi, S.; Hayashi, Y.; Yoshida, A.; Koike, H.; Akishinonomiya, F.; Willerslev, E.; Hasegawa, M. (2016-12-15). "Phylogenomics and Morphology of Extinct Paleognaths Reveal the Origin and Evolution of the Ratites". Current Biology. 27 (1): 68–77. doi:10.1016/j.cub.2016.10.029. PMID 27989673.
^ Mayr, Gerald; De Pietri, Vanesa L.; Love, Leigh; Mannering, Al; Scofield, Richard Paul (2019). "Oldest, smallest and phylogenetically most basal pelagornithid, from the early Paleocene of New Zealand, sheds light on the evolutionary history of the largest flying birds". Papers in Palaeontology. 7 (1): 217–233. doi:10.1002/spp2.1284.
^ Marjanović, D. (2021). "The Making of Calibration Sausage Exemplified by Recalibrating the Transcriptomic Timetree of Jawed Vertebrates". Frontiers in Genetics. 12. 521693. doi:10.3389/fgene.2021.521693. PMC 8149952.
^ Claramunt, S.; Cracraft, J. (Dec 2015). "A new time tree reveals Earth history's imprint on the evolution of modern birds". Sci Adv. 1 (11): e1501005. Bibcode:2015SciA....1E1005C. doi:10.1126/sciadv.1501005. PMC 4730849. PMID 26824065.
^ Torres, Christopher R.; Norell, Mark A.; Clarke, Julia A. (2021). "Bird neurocranial and body mass evolution across the end-Cretaceous mass extinction: The avian brain shape left other dinosaurs behind". Science Advances. 7 (31). Bibcode:2021SciA....7.7099T. doi:10.1126/sciadv.abg7099. PMC 8324052. PMID 34330706.
^ Mindell & Brown (2005)
^ Sangster, George; Braun, Edward L.; Johansson, Ulf S.; Kimball, Rebecca T.; Mayr, Gerald; Suh, Alexander (2022-01-01). "Phylogenetic definitions for 25 higher-level clade names of birds" (PDF). Avian Research. 13: 100027. Bibcode:2022AvRes..1300027S. doi:10.1016/j.avrs.2022.100027. ISSN 2053-7166.
^ For a draft phylogeny of Neoaves that is based on a review of massive amounts of published sources, and probably rather close to "the real thing", see Mindell et al. (2005)
^ Mindell et al. (2005)
^ Stiller, J., Feng, S., Chowdhury, AA. et al. Complexity of avian evolution revealed by family-level genomes. Nature (2024). https://doi.org/10.1038/s41586-024-07323-1

References

Claramunt, S.; Cracraft, J. (2015). "A new time tree reveals Earth history's imprint on the evolution of modern birds". Sci Adv. 1 (11): e1501005. Bibcode:2015SciA....1E1005C. doi:10.1126/sciadv.1501005. PMC 4730849. PMID 26824065.
Mindell, David P. & Brown, Joseph W. (2005): The Tree of Life Web Project - Neornithes. Version of 2005-DEC-14. Retrieved 2008-JAN-08.
Mindell, David P.; Brown, Joseph W. & Harshman, John (2005): The Tree of Life Web Project - Neoaves. Version of 2005-DEC-14. Retrieved 2008-JAN-08.