COLUMBAVES Prum et al., 2015
Columbaves contains about 5% of the avian tree. It is the next branch of Neoaves after Mirandornithes, according to Stiller et al. (2024). However, Kuhl et al. (2021) group it with the Gruimorphae (rails and shorebirds), Hoatzin, and Strisores (nightjars, swifts, hummingbirds), partially recalling Fain and Houde's (2004) Metaves hypothesis. However, I'm giving preference to Stiller et al. because they use more data, and have taken steps to avoid the frozen gene problem.
As for Fain and Houde (2004), Jarvis et al. (2014) already made it clear that the original Metaves hypothesis did not hold up in larger datasets. It was in fact an artifact of using the β-fibrogen gene, particularly the 7th intron. Nonetheless, some of the signal was real, and Columbaves remains, gathering the Otidimorphae and Columbimorphae once the Eurypygimorphae, Strisores, and Hoatzin have been moved elsewhere, as they are by Stiller et al.
Jarvis et al. (2014) divided Neoaves into g (including Mirandornithes) and Passerea. This doesn't make sense in the Stiller et al. phylogeny. Had Jarvis et al. used this topology, they would have surely used Passerea for the remainder of the avian tree after the Columbaves split. That is how I will use it here, but with quotation marks to indicate it is not the Jarvis et al. Passerea.
Focusing on Columbaves, we find it is comprised of two superorders: Otidimorphae and Columbimorphae. Stiller et al. (2024) found that each of them is comprised of three orders. Otidimorphae includes Musophagiformes (turacos), Otidiformes (bustards), and Cuculiformes (cuckoos) while Columbimorphae includes Mesitornithiformes (mesites), Pterocliformes (sandgrouse), and Columbiformes (doves and pigeons). This is one place where Kuhl et al. (2021) disagree. They put the cuckoos in Columbimorphae, not Otidimorphae.
Reading off Fig. ED-2, we see that Stiller et al. (2024) estimate the division into Columbaves and "Passerea" occurred 67.5±1.25 million years ago. The split between Otidimorphae and Columbimorphae happened relatively soon after. They estimate it was about 1.7 million years later.
OTIDIMORPHAE Wagler 1830
I follow the arrangment of the orders in Stiller et al. (2024), with the turacos basal in Otidimorphae. In contrast, the cuckoos were basal in Jarvis et al. (2014). No one is surprised by a grouping that includes the turacos and cuckoos. What is surprising is that the bustards snuck in next to the turacos. Even more surpising is Kuhl et al. (2021) which has the turacos and bustards as sister taxa, with the cuckoos exiled to Columbimorphae.
According to Stiller et al.'s point estimates, the splits between all three orders in Otidimorphae happened before 60 mya.
MUSOPHAGIFORMES Seebohm, 1890
Musophagidae: Turacos Lesson, 1828
10 genera, 23 species HBW-4
The arrangement of the Turacos is based on a combination of Veron and Winney (2000) and Njabo and Sorenson (2009). Both use essentially the same data set (except for bannermani), but analyze it differently, with mostly the same results.
As in previous phylogenies of the turacos, there are three subfamilies: Corythaeolinae, Criniferinae, and Musophaginae. Corythaeolinae is monotypic. Musophaginae variously ends up sister to each of the others, so I treat this as an unresolved trichotomy. Within Criniferinae, Veron and Winney (2000) and Njabo and Sorenson (2009) found that the White-bellied Go-away-bird is sister to the rest. This demands a change of genus for it, in this case to Criniferoides (Roberts, 1926).
They also found that the Purple-crested and Ruwenzori Turacos are sister species, and basal in the Musophaginae subfamily. Veron and Winney (2000) also recommend placing them in the same genus. In that case Gallirex has priority.
For the rest, the tree here leaves two main naming options. One is to put them in one genus (Tuaraco). This does a poor job of reflecting phylogeny. The other option is the one followed here, using a more narrowly circumscribed Tuaraco. The name Menelikornis (von Boetticher 1947) applies to the White-cheeked Turaco, while Proturacus (Bates 1923) has priority for the Bannerman's Turaco group. Finally, given the difference in appearance between the two Musophaga and the Yellow-billed Turaco, I perfer to put the latter in a separate genus. It becomes Pseudopoetus (von Boetticher 1947).
Corythaeolinae: Great Blue Turaco Verheyen, 1956
Criniferinae: Go-away-birds and Plantain-eaters Verheyen, 1956
- White-bellied Go-away-bird, Criniferoides leucogaster
- Gray Go-away-bird, Corythaixoides concolor
- Bare-faced Go-away-bird, Corythaixoides personatus
- Western Plantain-eater, Crinifer piscator
- Eastern Plantain-eater, Crinifer zonurus
Musophaginae: Turacos Lesson, 1828
- Purple-crested Turaco, Gallirex porphyreolophus
- Ruwenzori Turaco, Gallirex johnstoni
- White-cheeked Turaco, Menelikornis leucotis
- Yellow-billed Turaco, Pseudopoetus macrorhynchus
- Violet Turaco, Musophaga violacea
- Ross's Turaco, Musophaga rossae
- Bannerman's Turaco, Proturacus bannermani
- White-crested Turaco, Proturacus leucolophus
- Red-crested Turaco, Proturacus erythrolophus
- Black-billed Turaco, Tauraco schuettii
- Schalow's Turaco, Tauraco schalowi
- Hartlaub's Turaco, Tauraco hartlaubi
- Ruspoli's Turaco, Tauraco ruspolii
- Guinea Turaco, Tauraco persa
- Knysna Turaco, Tauraco corythaix
- Livingstone's Turaco, Tauraco livingstonii
- Fischer's Turaco, Tauraco fischeri
OTIDIFORMES Wagler 1830
The bustards have been reorganized using Cohen (2011, esp. Figs 2.4-2.5), who included all the bustards in a multigene analysis. The arrangment is rather different from the cytochrome-b results of Pitra et al. (2002), which I had previously used. Some uncertainty remains concerning the exact position of Lissotis, Tetrax, and the Otis-Chlamydotis clade.
Both Pitra et al. (2002) and Cohen (2011) support merging Neotis (Sharpe, 1893) into Ardeotis (Le Maout, 1853) and separating Heterotetrax (Sharpe, 1894, type vigorsii) from Eupodotis. Those three species have sometimes been considered a separate genus under this name.
Following Hockey et al. (2005, aka Roberts VII), Barrow's Korhaan, Eupodotis barrowii, is considered a subspecies of White-bellied Bustard, Eupodotis senegalensis.
Otididae: Bustards Rafinesque, 1815
11 genera, 26 species HBW-3
- Black-bellied Bustard, Lissotis melanogaster
Click for Otididae tree - Hartlaub's Bustard, Lissotis hartlaubii
- Nubian Bustard, Ardeotis nuba
- Denham's Bustard, Ardeotis denhami
- Ludwig's Bustard, Ardeotis ludwigii
- Heuglin's Bustard, Ardeotis heuglinii
- Arabian Bustard, Ardeotis arabs
- Kori Bustard, Ardeotis kori
- Great Indian Bustard, Ardeotis nigriceps
- Australian Bustard, Ardeotis australis
- Little Bustard, Tetrax tetrax
- Great Bustard, Otis tarda
- Houbara Bustard, Chlamydotis undulata
- Macqueen's Bustard, Chlamydotis macqueenii
- Bengal Florican, Houbaropsis bengalensis
- Lesser Florican, Sypheotides indicus
- Red-crested Korhaan, Lophotis ruficrista
- Savile's Bustard, Lophotis savilei
- Buff-crested Bustard, Lophotis gindiana
- Little Brown Bustard, Heterotetrax humilis
- Rueppell's Korhaan, Heterotetrax rueppelii
- Karoo Korhaan, Heterotetrax vigorsii
- Northern Black-Korhaan, Afrotis afraoides
- Southern Black-Korhaan, Afrotis afra
- Blue Korhaan, Eupodotis caerulescens
- White-bellied Bustard, Eupodotis senegalensis
CUCULIFORMES Wagler 1830
There is no real question that the cuckoos form a clade. The Cuculiformes are placed here following Jarvis et al. (2014). Some divisions between the branches of Cuculiformes are old, and it may make sense to promotes some subfamilies to family level.
Cuculidae: Cuckoos Leach, 1820
32 genera, 147 species HBW-4
Very complete information is available on cuckoo taxonomy. Sorenson and Payne (2005) carried out a very extensive study of Cuckoo DNA. The result is the sequence used in Payne's book (2005). After adjusting the species limits for a couple of couas, it is the same sequence that is used here. Click on the graphic below for the genus-level tree.
Although Burchell's Coucal, Centropus superciliosus burchellii is sometimes treated as a separate species, the genetic data examined by Sorenson and Payne (2005) does not support this. The same is true of the Kai Coucal, Centropus phasianinus spilopterus.
The race aeruginosus has been transferred from Rusty-breasted Cuckoo, Cacomantis sepulcralis, to Moluccan Cuckoo. As aeruginosus (Salvadori, 1878) has priority over heinrichi (Stresemann, 1931), the Moluccan Cuckoo becomes Cacomantis aeruginosus.
Crotophaginae: Anis Swainson, 1837
- Guira Cuckoo, Guira guira
Click for Cuckoo tree - Greater Ani, Crotophaga major
- Smooth-billed Ani, Crotophaga ani
- Groove-billed Ani, Crotophaga sulcirostris
Neomorphinae: Ground-Cuckoos, Roadrunners Shelley, 1891
- Striped Cuckoo, Tapera naevia
- Pheasant Cuckoo, Dromococcyx phasianellus
- Pavonine Cuckoo, Dromococcyx pavoninus
- Lesser Ground-Cuckoo, Morococcyx erythropygus
- Lesser Roadrunner, Geococcyx velox
- Greater Roadrunner, Geococcyx californianus
- Banded Ground-Cuckoo, Neomorphus radiolosus
- Rufous-winged Ground-Cuckoo, Neomorphus rufipennis
- Red-billed Ground-Cuckoo, Neomorphus pucheranii
- Rufous-vented Ground-Cuckoo, Neomorphus geoffroyi
- Scaled Ground-Cuckoo, Neomorphus squamiger
Couinae: Couas Bonaparte, 1854
- Sumatran Ground-Cuckoo, Carpococcyx viridis
- Bornean Ground-Cuckoo, Carpococcyx radiceus
- Coral-billed Ground-Cuckoo, Carpococcyx renauldi
- Crested Coua, Coua cristata
- Verreaux's Coua, Coua verreauxi
- Blue Coua, Coua caerulea
- Red-capped Coua, Coua ruficeps
- Red-fronted Coua, Coua reynaudii
- Coquerel's Coua, Coua coquereli
- Running Coua, Coua cursor
- Giant Coua, Coua gigas
- Snail-eating Coua, Coua delalandei
- Red-breasted Coua, Coua serriana
Centropodinae: Coucals Horsfield, 1823
- Buff-headed Coucal, Centropus milo
- White-necked Coucal, Centropus ateralbus
- Ivory-billed Coucal, Centropus menbeki
- Biak Coucal, Centropus chalybeus
- Rufous Coucal, Centropus unirufus
- Green-billed Coucal, Centropus chlororhynchos
- Black-faced Coucal, Centropus melanops
- Black-hooded Coucal, Centropus steerii
- Short-toed Coucal, Centropus rectunguis
- Bay Coucal, Centropus celebensis
- Gabon Coucal, Centropus anselli
- Black-throated Coucal, Centropus leucogaster
- Senegal Coucal, Centropus senegalensis
- Blue-headed Coucal, Centropus monachus
- Coppery-tailed Coucal, Centropus cupreicaudus
- White-browed Coucal, Centropus superciliosus
- Sunda Coucal, Centropus nigrorufus
- Greater Coucal, Centropus sinensis
- Andaman Coucal, Centropus andamanensis
- Goliath Coucal, Centropus goliath
- Malagasy Coucal, Centropus toulou
- Black Coucal, Centropus grillii
- Philippine Coucal, Centropus viridis
- Lesser Coucal, Centropus bengalensis
- Violaceous Coucal, Centropus violaceus
- Black-billed Coucal, Centropus bernsteini
- Pheasant Coucal, Centropus phasianinus
Cuculinae: Cuckoos Leach 1820
Rhinorthini: Raffles's Malkoha Informal
- Raffles's Malkoha, Rhinortha chlorophaea
Phaenicophaeini: Malkohas, Clamator, American Cuckoos Horsfield, 1822
- Green Malkoha, Ceuthmochares australis
- Blue Malkoha, Ceuthmochares aereus
- Sirkeer Malkoha, Taccocua leschenaultii
- Red-billed Malkoha, Zanclostomus javanicus
- Chestnut-breasted Malkoha, Phaenicophaeus curvirostris
- Chestnut-bellied Malkoha, Phaenicophaeus sumatranus
- Red-faced Malkoha, Phaenicophaeus pyrrhocephalus
- Blue-faced Malkoha, Phaenicophaeus viridirostris
- Black-bellied Malkoha, Phaenicophaeus diardi
- Green-billed Malkoha, Phaenicophaeus tristis
- Yellow-billed Malkoha, Rhamphococcyx calyorhynchus
- Rough-crested Malkoha, Dasylophus superciliosus
- Scale-feathered Malkoha, Dasylophus cumingi
- Chestnut-winged Cuckoo, Clamator coromandus
- Great Spotted Cuckoo, Clamator glandarius
- Levaillant's Cuckoo, Clamator levaillantii
- Jacobin Cuckoo, Clamator jacobinus
- Little Cuckoo, Coccycua minuta
- Dwarf Cuckoo, Coccycua pumila
- Ash-colored Cuckoo, Coccycua cinerea
- Squirrel Cuckoo, Piaya cayana
- Black-bellied Cuckoo, Piaya melanogaster
- Dark-billed Cuckoo, Coccyzus melacoryphus
- Yellow-billed Cuckoo, Coccyzus americanus
- Pearly-breasted Cuckoo, Coccyzus euleri
- Mangrove Cuckoo, Coccyzus minor
- Cocos Cuckoo, Coccyzus ferrugineus
- Black-billed Cuckoo, Coccyzus erythropthalmus
- Gray-capped Cuckoo, Coccyzus lansbergi
- Chestnut-bellied Cuckoo, Coccyzus pluvialis
- Bay-breasted Cuckoo, Coccyzus rufigularis
- Jamaican Lizard-Cuckoo, Coccyzus vetula
- Puerto Rican Lizard-Cuckoo, Coccyzus vieilloti
- Great Lizard-Cuckoo, Coccyzus merlini
- Hispaniolan Lizard-Cuckoo, Coccyzus longirostris
Cuculini: Old World Parasitic Cuckoos Wagler 1830
- Thick-billed Cuckoo, Pachycoccyx audeberti
- Dwarf Koel, Microdynamis parva
- Asian Koel, Eudynamys scolopaceus
- Black-billed Koel, Eudynamys melanorhynchus
- Pacific Koel, Eudynamys orientalis
- Long-tailed Koel / Long-tailed Cuckoo, Urodynamis taitensis
- Channel-billed Cuckoo, Scythrops novaehollandiae
- Asian Emerald-Cuckoo, Chrysococcyx maculatus
- Violet Cuckoo, Chrysococcyx xanthorhynchus
- Diederik Cuckoo, Chrysococcyx caprius
- Klaas's Cuckoo, Chrysococcyx klaas
- Yellow-throated Cuckoo, Chrysococcyx flavigularis
- African Emerald-Cuckoo, Chrysococcyx cupreus
- Long-billed Cuckoo, Chrysococcyx megarhynchus
- Horsfield's Bronze-Cuckoo, Chrysococcyx basalis
- Black-eared Cuckoo, Chrysococcyx osculans
- Rufous-throated Bronze-Cuckoo, Chrysococcyx ruficollis
- Shining Bronze-Cuckoo, Chrysococcyx lucidus
- White-eared Bronze-Cuckoo, Chrysococcyx meyerii
- Little Bronze-Cuckoo, Chrysococcyx minutillus
- Pallid Cuckoo, Cacomantis pallidus
- White-crowned Cuckoo, Cacomantis leucolophus
- Chestnut-breasted Cuckoo, Cacomantis castaneiventris
- Fan-tailed Cuckoo, Cacomantis flabelliformis
- Banded Bay Cuckoo, Cacomantis sonneratii
- Plaintive Cuckoo, Cacomantis merulinus
- Gray-bellied Cuckoo, Cacomantis passerinus
- Brush Cuckoo, Cacomantis variolosus
- Rusty-breasted Cuckoo, Cacomantis sepulcralis
- Moluccan Cuckoo, Cacomantis aeruginosus
- Dusky Long-tailed Cuckoo, Cercococcyx mechowi
- Olive Long-tailed Cuckoo, Cercococcyx olivinus
- Barred Long-tailed Cuckoo, Cercococcyx montanus
- Fork-tailed Drongo-Cuckoo, Surniculus dicruroides
- Philippine Drongo-Cuckoo, Surniculus velutinus
- Square-tailed Drongo-Cuckoo, Surniculus lugubris
- Moluccan Drongo-Cuckoo, Surniculus musschenbroeki
- Moustached Hawk-Cuckoo, Hierococcyx vagans
- Dark Hawk-Cuckoo, Hierococcyx bocki
- Large Hawk-Cuckoo, Hierococcyx sparverioides
- Common Hawk-Cuckoo, Hierococcyx varius
- Rufous Hawk-Cuckoo, Hierococcyx hyperythrus
- Philippine Hawk-Cuckoo, Hierococcyx pectoralis
- Malaysian Hawk-Cuckoo, Hierococcyx fugax
- Hodgson's Hawk-Cuckoo, Hierococcyx nisicolor
- Black Cuckoo, Cuculus clamosus
- Red-chested Cuckoo, Cuculus solitarius
- Lesser Cuckoo, Cuculus poliocephalus
- Sulawesi Cuckoo, Cuculus crassirostris
- Indian Cuckoo, Cuculus micropterus
- Madagascan Cuckoo, Cuculus rochii
- African Cuckoo, Cuculus gularis
- Oriental Cuckoo, Cuculus optatus
- Himalayan Cuckoo, Cuculus saturatus
- Sunda Cuckoo, Cuculus lepidus
- Common Cuckoo, Cuculus canorus