| Neoavians Temporal range: Paleocene – Holocene, Possible Late Cretaceous origin based on molecular clock [2] [3] | |
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| Great crested grebe (Podiceps cristatus) | |
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| House sparrow (Passer domesticus) | |
Scientific classification  | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Aves |
| Infraclass: | Neognathae |
| Clade: | Neoaves Sibley et al., 1988 |
| Clades | |
Neoaves is a clade that consists of all modern birds (Neornithes or Aves) with the exception of Palaeognathae (ratites and kin) and Galloanserae (ducks, chickens and kin). [4] This group is defined in the PhyloCode by George Sangster and colleagues in 2022 as "the most inclusive crown clade containing Passer domesticus , but not Gallus gallus ". [5] Almost 95% of the roughly 10,000 known species of extant birds belong to the Neoaves. [6]
The early diversification of the various neoavian groups occurred very rapidly around the Cretaceous–Paleogene extinction event, [7] [8] and attempts to resolve their relationships with each other have resulted initially in much controversy. [9] [10]
The early diversification of the various neoavian groups occurred very rapidly around the Cretaceous–Paleogene extinction event. [11] As a result of the rapid radiation, attempts to resolve their relationships have produced conflicting results, some quite controversial, especially in the earlier studies. [12] [13] [14] Nevertheless, some recent large phylogenomic studies of Neoaves have led to much progress on defining orders and supraordinal groups within Neoaves. Still, the studies have failed to produce to a consensus on an overall high order topology of these groups. [15] [16] [17] [14] A genomic study of 48 taxa by Jarvis and colleagues in 2014 divided Neoaves into two main clades, Columbea and Passerea, but an analysis of 198 taxa by Prum and colleagues in 2015 recovered different groupings for the earliest split in Neoaves. [15] [16] A reanalysis with an extended dataset by Reddy and colleagues in 2017 suggested this was due to the type of sequence data, with coding sequences favouring the Prum topology. [17] The disagreement on topology even with large phylogenomic studies led Alexander Suh in 2016 to propose a hard polytomy of nine clades as the base of Neoaves. [18] An analysis by Houde and colleagues in 2019 recovered Columbea and a reduced hard polytomy of six clades within Passerea. [19]
Despite other disagreements, these studies do agree on a number of supraorderal groups, which Reddy and colleagues in 2017 dubbed the "magnificent seven", which together with three "orphaned orders" make up Neoaves. [17] Significantly, they both include a large waterbird clade (Aequornithes) and a large landbird clade (Telluraves). The groups defined by Reddy and colleagues (2017) are as follows:
| Jarvis et al. (2014) [4] | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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| Prum et al. (2015) [20] | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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| Suh (2016) — a hard polytomy [18] | ||||||||||||||||||||||||||||||||||||
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| Reddy et al. (2017) [17] | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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| Houde et al. (2019) —polytomy in Passerea [19] | |||||||||||||||||||||||||||||||||||||||||||||||||||
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| Kuhl et al. (2021) [2] | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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| Braun & Kimball (2021) — soft polytomies at the base of Neoaves and in Passerea [21] | ||||||||||||||||||||||||||||||||||||||||||
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| Wu et al. (2024) [22] | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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| Stiller et al (2024) [23] | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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The following cladogram illustrates the proposed relationships between all neoavian bird clades. [24]
| Neoaves |
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Otidae is a clade that includes the superorders Otidimorphae and Strisores. It was identified in 2014 by genome analysis. Earlier it was thought that Strisores was closely related to birds such as pigeons, flamingos, tropicbirds, and the sunbittern and kagu in the taxon Metaves, but subsequent work has provided evidence that Metaves is polyphyletic. Although analyses of genome data provided relatively high support for monophyly of Otidae, indicating that it is sister to all other Passerea clades, other analyses of large data matrices have not recovered a clade comprising Otidimorphae and Strisores, raising questions about the monophyly of Otidae.
Neognathae 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.
Mirandornithes is a clade that consists of flamingos and grebes. Many scholars use the term Phoenicopterimorphae for the superorder containing flamingoes and grebes.
Aequornithes, or core water birds, are defined in the PhyloCode as "the least inclusive crown clade containing Pelecanus onocrotalus and Gavia immer".
The order Falconiformes is represented by the extant family Falconidae and a handful of enigmatic Paleogene species. Traditionally, the other bird of prey families Cathartidae, Sagittariidae (secretarybird), Pandionidae (ospreys), Accipitridae (hawks) were classified in Falconiformes. A variety of comparative genome analyses published since 2008, however, found that falcons are part of a clade of birds called Australaves, which also includes seriemas, parrots and passerines. Within Australaves falcons are more closely related to the parrot-passerine clade than they are to the seriemas. The hawks, vultures and owls are placed in the clade Afroaves.
Metaves is a controversial group proposed by Fain & Houde (2004) and later rescued on the studies of Ericson et al. (2006) and Hackett et al. (2008). This group consists of several lineages that diversified early in Neornithes evolution. These lineages include Strisores, pigeons, sandgrouses, mesites, Eurypygae, tropicbirds and Mirandornithes, but the exact members of Metaves and their relationship differs between those studies, and the group is only supported by the β-fibrinogen gene.
Strisores, sometimes called nightbirds, is a clade of birds that includes the living families and orders Caprimulgidae, Nyctibiidae (potoos), Steatornithidae (oilbirds), Podargidae (frogmouths), Apodiformes, as well as the Aegotheliformes (owlet-nightjars) whose distinctness was only recently realized. The Apodiformes and the Aegotheliformes form the Daedalornithes.
Australaves is a clade of birds, defined in 2012, consisting of the Eufalconimorphae as well as the Cariamiformes. They appear to be the sister group of Afroaves. This clade was defined in the PhyloCode by George Sangster and colleagues in 2022 as "the least inclusive crown clade containing Cariama cristata and Passer domesticus".
Afroaves is a clade of birds, consisting of the kingfishers and kin (Coraciiformes), woodpeckers and kin (Piciformes), hornbills and kin (Bucerotiformes), trogons (Trogoniformes), cuckoo roller (Leptosomiformes), mousebirds (Coliiformes), owls (Strigiformes) and raptors (Accipitriformes). The most basal clades are predatory, suggesting the last common ancestor of Afroaves was also a predatory bird. This group was defined in the PhyloCode by George Sangster and colleagues in 2022 as "the least inclusive crown clade containing Accipiter nisus, Colius colius, and Picus viridis, but not Passer domesticus".
Telluraves is a recently defined clade of birds defined by their arboreality. Based on most recent genetic studies, the clade unites a variety of bird groups, including the australavians as well as the afroavians. This grouping was defined in the PhyloCode by George Sangster and colleagues in 2022 as "the least inclusive crown clade containing Accipiter nisus and Passer domesticus". They appear to be the sister group of the Phaethoquornithes.
Passerea is a clade of neoavian birds that was proposed by Jarvis et al. (2014). Their genomic analysis recovered two major clades within Neoaves, Passerea and Columbea, and concluded that both clades appear to have many ecologically driven convergent traits.
Eurypygimorphae or Phaethontimorphae is a clade of birds that contains the orders Phaethontiformes (tropicbirds) and Eurypygiformes recovered by genome analysis. The relationship was first identified in 2013 based on their nuclear genes. This group was defined in the PhyloCode by George Sangster and colleagues in 2022 as "the least inclusive crown clade containing Phaethon aethereus, Eurypyga helias, and Rhynochetos jubatus". Historically these birds were placed at different parts of the tree, with tropicbirds in Pelecaniformes and the kagu and sunbittern in Gruiformes. Some genetic analyses have placed the eurypygimorph taxa in the controversial and obsolete clade Metaves, with uncertain placement within that group. More recent molecular studies support their grouping together in Eurypygimorphae, which is usually recovered as the sister taxon to Aequornithes within Ardeae.
Otidimorphae is a clade of birds that contains the orders Cuculiformes (cuckoos), Musophagiformes (turacos), and Otidiformes (bustards) identified in 2014 by genome analysis. George Sangster and colleagues in 2022 named the clade uniting turacos and bustards as Musophagotides, defining it in the PhyloCode as "the least inclusive crown clade containing Otis tarda and Musophaga violacea, but not Grus grus or Mesitornis variegatus".
Picocoraciae is a clade that contains the order Bucerotiformes and the clade Picodynastornithes supported by various genetic analysis and morphological studies. While these studies supported a sister grouping of Coraciiformes and Piciformes, a large scale, sparse supermatrix has suggested alternative sister relationship between Bucerotiformes and Piciformes instead. This group was defined in the PhyloCode by George Sangster and colleagues in 2022 as "the least inclusive crown clade containing Buceros rhinoceros, Coracias garrulus, and Picus viridis".
Columbimorphae is a clade/superorder discovered by genome analysis that includes birds of the orders Columbiformes, Pterocliformes (sandgrouse), and Mesitornithiformes (mesites). This group was defined in the PhyloCode by George Sangster and colleagues in 2022 as "the least inclusive crown clade containing Columba oenas, Mesitornis variegatus, and Pterocles alchata". Previous analyses had also recovered this grouping, although the exact relationships differed. Some studies indicated a sister relationship between sandgrouse and pigeons while other studies favored a sister grouping of mesites and sandgrouse instead. This sister relationship of the sandgrouses and mesites was named by George Sangster and colleagues in 2022 as the clade Pteroclimesites and defined in the PhyloCode as "the least inclusive crown clade containing Mesitornis variegatus and Pterocles alchata".
Columbaves is a clade that contains Columbimorphae and Otidimorphae discovered by genomic analysis by Prum et al. (2015). This conflicts with the Columbea and Otidae hypotheses which Mirandornithes are the sister taxon to Columbimorphae and Cypselomorphae the sister taxon to Otidimorphae, respectively, found by Jarvis et al. (2014). Neither hypothesis supports the two subdivisions of Metaves and Coronoaves as previous studies had found.
Aequorlitornithes is a clade of waterbirds recovered in a comprehensive genomic systematic study using nearly 200 species in 2015. It contains the clades Charadriiformes, Mirandornithes and Phaethoquornithes. Previous studies have found different placement for the clades in the tree.
Inopinaves is a clade of neoavian birds recovered in a compressive genomic systematic study using nearly 200 species in 2015. It contains the clades Opisthocomiformes (hoatzin) and Telluraves ; the study shows that the hoatzin diverged from other birds 64 million years ago. Previous studies have placed the hoatzin in different parts of the bird family tree; however, despite its unusual morphology, genetic studies have shown the hoatzin is not as primitive or as ancient as once thought; it could be a very derived bird that reverted to or retains some plesiomorphic traits.
Vanescaves is a probable clade of strisorean birds that include the clades Steatornithiformes, Nyctibiiformes, Podargiformes (frogmouths), and Apodimorphae. Some molecular studies do support the grouping of these birds, others offer conflicting positions of the non-apodimorphaean strisoreans. In 2019 the authors Chen et al. performed a combined analysis using 2289 ultra-conserved elements [UCEs], 117 morphological characters from extant and fossil taxa found support in this clade. The authors then proposed to name this group, which its meaning is Latin for "vanish birds" in reference to the disparate nature of their geographic distribution, as well as to the poem "A Route of Evanescence" by the American poet Emily Dickinson which features a hummingbird as the main subject. In 2020 Chen & Field named the two major subclades of this group, with Sedentaves and Letornithes for their crown-groups.
Litusilvanae is a proposed clade of birds, position as the sister clade to Aequorlitornithes. This clade comprises Gruimorphae and Strisores. While different lines of evidence from molecular, morphology and the fossil record has found support in the clades Gruimorphae and Strisores Wu et al. (2024) was the first to find support in such a novel sister group relationship between these two taxa.
