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Superregnum: Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Cladus: Craniata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Infraclassis: Archosauromorpha
Cladus: Crurotarsi
Divisio: Archosauria
Subsectio: Ornithodira
Subtaxon: Dinosauromorpha
Cladus: Dinosauria
Ordo: Saurischia
Cladus: Theropoda
Cladus: Neotheropoda
Infraclassis: Aves
Cladus: Euavialae
Cladus: Avebrevicauda
Cladus: Pygostylia
Cladus: Ornithothoraces
Cladus: Euornithes
Cladus: Ornithuromorpha
Cladus: Ornithurae
Cladus: Carinatae
Parvclassis: Neornithes
Cohors: Neognathae
Ordo: Musophagiformes

Familia: Musophagidae
Subfamiliae: Corythaeolinae - Criniferinae - Musophaginae
Name

Musophagidae
Vernacular names
svenska: Turakoer
Türkçe: Muzculgiller

The turacos make up the bird family Musophagidae (/ˌmjuːzoʊˈfædʒɪdiː/ "banana-eaters"), which includes plantain-eaters and go-away-birds. In southern Africa both turacos and go-away-birds are commonly known as loeries. They are semi-zygodactylous: the fourth (outer) toe can be switched back and forth. The second and third toes, which always point forward, are conjoined in some species. Musophagids often have prominent crests and long tails; the turacos are noted for peculiar and unique pigments giving them their bright green and red feathers.

Traditionally, this group has been allied with the cuckoos in the order Cuculiformes, but the Sibley-Ahlquist taxonomy raises this group to a full order Musophagiformes. They have been proposed to link the hoatzin to the other living birds,[1] but this was later disputed.[2] Recent genetic analyses have strongly supported the order ranking of Musophagiformes.[3][4][5]

Musophagidae is one of very few bird families endemic to Africa,[6] one other being the mousebirds, Colliidae. All species are frugivorous, but they also eat leaves, buds, and flowers. Figs are an important part of their diet. They have rounded wings and long tails and strong legs, making them poor fliers, but good runners.[6]

Turacos are medium-sized arboreal birds endemic to sub-Saharan Africa, where they live in forests, woodland and savanna.

They are gregarious, non-migratory birds which move in family groups of up to 10. Many species are noisy, with the go-away-birds being especially noted for their piercing alarm calls, which alert other fauna to the presence of predators; their common name is onomatopoeia of this. Musophagids build large stick nests in trees, and lay 2 or 3 eggs. The young are born with thick down and open, or nearly-open, eyes.[7]

Morphology
Female white-bellied go-away-bird, Crinifer leucogaster

Most turacos are medium-sized birds – an exception being the large great blue turaco – with long tails and short, rounded wings. They range in length from 40 to 75 cm (16–30 in). Their flight is weak, but they are strong climbers and are able to move nimbly on branches and through vegetation. Juveniles have claws on the wings that help them climb.[8][9][10] They have a unique foot arrangement, where the fourth toe can be brought around to the back of the foot where it almost touches the first toe, or brought around so that it is near the second and third. In spite of this flexibility the toe is actually usually held at right angles to the axis of the foot.[11]

The plumage of go-away-birds and plantain-eaters is mainly grey and white. The turacos on the other hand are brightly coloured birds, usually blue, green or purple. The green colour in turacos comes from turacoverdin, the only true green pigment in birds known to date. Other "greens" in bird colors result from a yellow pigment such as some carotenoid, combined with the prismatic physical structure of the feather itself which scatters the light in a particular way and giving a blue colour.

Turaco wings contain the red pigment turacin, unlike in other birds where red colour is due to carotenoids. Both pigments are derived from porphyrins and only known from the Musophagidae into the 21st century, but especially the little-researched turacoverdin might have relatives in other birds. The incidence of turacoverdin in relation to habitat is of interest to scientists, being present in forest species but absent in savanna- and acacia-living species.[11]

Little is known about the longevity of wild turacos, but in captivity they are proving to be exceptionally long-lived, easily living to 30 years in captivity. A bird in the Cotswold Wildlife Park collection in England approached its 37th year.[12]
Evolution and systematics

The fossil genus Veflintornis is known from the Middle Miocene of Grive-Saint-Alban (France). It was established as Apopempsis by Pierce Brodkorb in 1971, but this is pre-empted by Schenkling's 1903 use of the name for some beetles. "Apopempsis" africanus (Early Miocene of Kenya) might also belong there.[13]

Further fossil material of putative musophagids was found in Egypt as well as in Late Oligocene deposits at Gaimersheim in Germany and Middle Miocene deposits at Grive-Saint-Alban[14] and Vieux-Collonges (each in France).[13] While it is not entirely certain that these fossils are indeed of turacos, it nonetheless appears as if the family evolved in the Oligocene of central Europe or perhaps northern Africa, and later on shifted its distribution southwards. The climate of those European regions during the late Paleogene was not too dissimilar to that of (sub)tropical Africa today; the Saharan desert was not yet present and the distance across the Mediterranean was not much more than what it is today. Thus such a move south may well have been a very slow and gradual shifting of a large and continuous range.
Great blue turaco
Corythaeola cristata

The Early Eocene Promusophaga was initially believed to be the oldest record of the turacos; it was eventually reconsidered a distant relative of the ostrich and is now in the ratite family Lithornithidae. Filholornis from the Late Eocene or Early Oligocene of France is occasionally considered a musophagid, but its relationships have always been disputed. It is not often considered a turaco in more recent times and has been synonymised with the presumed gruiform Talantatos, though it is not certain whether this will become widely accepted.[13]

The phylogenetic analysis conducted by Field & Hsiang (2018) indicated that Eocene (Wasatchian) species Foro panarium known from the Fossil Butte Member of the Green River Formation (Wyoming, United States) was a stem-turaco.[15]
Phylogeny

The IOC World Bird List (version 10.1) recognises 23 species of turaco in six genera.[16] However, a phylogenetic analysis by Perktaş et al (2020) found genus Tauraco polyphyletic and a revised classification has been proposed based on molecular, morphological and biogeographic analysis.[17] This study recognised 33 species-level taxa in seven genera corresponding to the major clades. The following phylogenetic tree is based on this proposal and uses their proposed genus and species names.[17]


Corythaeolinae

Corythaeola cristata (great blue turaco)

Criniferinae

Crinifer leucogaster (white-bellied go-away-bird)

Crinifer

Crinifer piscator (western plantain-eater)

Crinifer zonurus (eastern plantain-eater)

sensu stricto

Crinifer personatus (bare-faced go-away-bird)

Crinifer concolor (gray go-away-bird)

(Crinifer sensu lato)[a]
Musophaginae
Gallirex[h]

Gallirex porphyreolophus (southern purple-crested turaco)

Gallirex chlorochlamys (northern purple-crested turaco)[b]

Gallirex kivuensis (Kivu turaco)[b]

Gallirex johnstoni (Rwenzori turaco)

Menelikornis[g]

Menelikornis ruspolii (Prince Ruspoli's turaco)

Menelikornis leucotis (white-cheeked turaco)

Menelikornis donaldsoni (Donaldson's turaco)[b]

Musophaga

Musophaga macrorhynchus (western yellow-billed turaco)

Musophaga verreauxii (eastern yellow-billed turaco)[b]

Musophaga violacea (violet turaco)

Musophaga rossae (Ross's turaco)

Tauraco
Proturacus[f]

Proturacus bannermani (Bannerman's turaco)

Proturacus erythrolophus (red-crested turaco)

Proturacus leucolophus (white-crested turaco)

Tauraco

Tauraco emini (eastern black-billed turaco)[c]

Tauraco hartlaubi (Hartlaub's turaco)

Tauraco persa (Guinea turaco or eastern green turaco)

Tauraco buffoni (western green turaco)[b]

Tauraco fischeri (Fischer's turaco)

Tauraco reichenowi (Reichenow's turaco)[d]

Tauraco corythaix (Knysna turaco)

Tauraco livingstonii (Livingstone's turaco)

Tauraco schuettii (black-billed turaco)

Tauraco chalcolophus (Ngorongoro turaco)[e]

Tauraco schalowi (Schalow's turaco)

Tauraco marungensis (Zambia turaco)[e]

Tauraco loitanus (Loita turaco)[e]

Notes:[17]

Broader definition of Crinifer proposed, because Corythaixoides leucogaster is recovered with the Crinifer species rather than the other Corythaixoides species
Proposed species split based on phylogenetic species principle.
Elevation of former subspecies to new species proposed because Tauraco schuettii was found to be polyphyletic.
Elevation of former subspecies to new species proposed because Tauraco livingstonii was found to be polyphyletic.
Proposed species split of Tauraco schalowi complex into four species based on phylogenetic species principle.
Proposed recognition of genus Proturacus for a clade of Tauraco bannermani, Tauraco erythrolophus and Tauraco leucolophus.
Proposed genus for clade of former Taurico species

Proposed genus for clade of former Taurico and Ruwenzorornis species

Species

The species of Musophagidae, arranged in taxonomic sequence and Paleofile.com websites are:[18][19]

Order Musophagiformes Seebohm 1890

Genus †Foro Olson, 1992?
†F. panarium Olson, 1992
Family Musophagidae Lesson 1828 [Apopempsidae Brodkorb, 1971b; Veflintornithidae Kašin, 1976]
Genus †Veflintornis Kašin 1976 [Apopempsis Brodkorb 1971 non Schenkling 1903]
†V. meini (Ballmann 1969) Kašin 1976 [Musophaga meini Ballmann 1969; Apopempsis meini (Ballmann 1969) Brodkorb 1971]
†V. africanus (Harrison 1980) [Musophaga africanus Harrison 1980; Apopempsis africanus (Harrison 1980)]
Subfamily Corythaeolinae
Genus Corythaeola Heine 1860
Great blue turaco, Corythaeola cristata (Vieillot 1816) Heine 1860
Subfamily Criniferinae
Genus Crinifer
Western plantain-eater, Crinifer piscator (Boddaert 1783)
Eastern plantain-eater, Crinifer zonurus (Rüppell 1835)
White-bellied go-away-bird, Crinifer leucogaster (Rüppell 1842) Roberts 1926
Grey go-away-bird, Crinifer concolor
Bare-faced go-away-bird, Crinifer personatus
Subfamily Musophaginae
Genus Gallirex
Purple-crested turaco, Gallirex porphyreolophus
Rwenzori turaco, Gallirex johnstoni
Genus Menelikornis
White-cheeked turaco, Menelikornis leucotis (Rüppell 1835)
Ruspoli's turaco, Menelikornis ruspolii Salvadori 1896
Genus Tauraco
Bannerman's turaco, Tauraco bannermani (Bates 1923)
White-crested turaco, Tauraco leucolophus (Heuglin 1855)
Red-crested turaco, Tauraco erythrolophus (Vieillot 1819)
Guinea turaco, Tauraco persa (Linnaeus 1758)
Knysna turaco, Tauraco corythaix (Wagler 1827)
Livingstone's turaco, Tauraco livingstonii Gray 1864
Fischer's turaco, Tauraco fischeri (Reichenow 1878)
Black-billed turaco, Tauraco schuettii (Cabanis 1879)
Schalow's turaco, Tauraco schalowi (Reichenow 1891)
Hartlaub's turaco, Tauraco hartlaubi (Fischer & Reichenow 1884)
Yellow-billed turaco, Tauraco macrorhynchus (Fraser 1839)
Violet turaco, Tauraco violaceus Isert 1788
Ross's turaco, Tauraco rossae Gould 1852

Interaction with humans

The crimson flight feathers of turacos have been treasured as status symbols to royalty and paramount chiefs all over Africa. They are recorded as being valued by the Swazi and Zulu royal families.[20] British ornithologist Constantine Walter Benson, who collected heavily in Africa, is alleged to have tasted every species he collected; he claimed that turacos tasted the best.[21]
Footnotes

Hughes & Baker (1999)
Sorenson et al. (2003)
Ericson, P.G.P.; et al. (2006). "Diversification of Neoaves: integration of molecular sequence data and fossils" (PDF). Biology Letters. 2 (4): 543–547. doi:10.1098/rsbl.2006.0523. PMC 1834003. PMID 17148284. Archived from the original (PDF) on 2008-03-07.
Hackett, S.J.; et al. (2008). "A Phylogenomic Study of Birds Reveals Their Evolutionary History". Science. 320 (5884): 1763–1768. Bibcode:2008Sci...320.1763H. doi:10.1126/science.1157704. PMID 18583609. S2CID 6472805.
Jarvis, E.D.; et al. (2014). "Whole-genome analyses resolve early branches in the tree of life of modern birds". Science. 346 (6215): 1320–1331. Bibcode:2014Sci...346.1320J. doi:10.1126/science.1253451. PMC 4405904. PMID 25504713.
Holzman, Barbara A. (2008). Tropical forest biomes. Greenwood Press. ISBN 978-0-313-33840-3. OCLC 470649845.
Marchant, S. (1991). Forshaw, Joseph (ed.). Encyclopaedia of Animals: Birds. London: Merehurst Press. p. 125. ISBN 978-1-85391-186-6.
Fain, Matthew G. & Houde, Peter (2004). "Parallel radiations in the primary clades of birds" (PDF). Evolution. 58 (11): 2558–2573. doi:10.1554/04-235. PMID 15612298. S2CID 1296408. Archived from the original (PDF) on 2017-07-09. Retrieved 2016-07-08.
"Violaceous Touraco" (PDF). rosamondgiffordzoo.org. 2010-03-01. Archived from the original (PDF) on 2011-01-05. Retrieved 2016-07-08.
"TURACO TAG HUSBANDRY MANUAL" (PDF). aviansag.org. 1998-08-18. Retrieved 2016-07-08.
Turner, Donald (1997), "Family Musophagidae (Turacos)", in del Hoyo, Josep; Elliott, Andrew; Sargatal, Jordi (eds.), Handbook of the Birds of the World. Volume 4, Sandgrouse to Cuckoos, Barcelona: Lynx Edicions, pp. 480–508, ISBN 978-84-87334-22-1
Originally from Nigel Hewston, discussed at the ITS AGM in spring 2012 (at the same venue)
Mlíkovský (2002)
"TT 149", a proximal left and a distal right tibiotarsus of a bird similar in size to living Tauraco: Ballmann (1969)
Daniel J. Field; Allison Y. Hsiang (2018). "A North American stem turaco, and the complex biogeographic history of modern birds". BMC Evolutionary Biology. 18 (1): 102. doi:10.1186/s12862-018-1212-3. PMC 6016133. PMID 29936914.
Gill, F.; Donsker, D.; Rasmussen, P. (eds.). "Family Musophagidae". IOC World Bird List. 10.1. International Ornithological Congress. Retrieved 19 June 2020.
Perktaş, Utku; Groth, Jeff G.; Barrowclough, George F. (April 2020). "Phylogeography, Species Limits, Phylogeny, and Classification of the Turacos (Aves: Musophagidae) Based on Mitochondrial and Nuclear DNA Sequences". American Museum Novitates. 2020 (3949): 1–61. doi:10.1206/3949.1. ISSN 0003-0082. S2CID 214763342.
"Taxonomic lists- Aves". Paleofile.com (net, info). Retrieved 30 December 2015.
Çınar, Ümüt (November 2015). "05 → Oᴛɪᴅᴀᴇ: Mᴜsᴏᴘʜᴀɢɪfᴏʀᴍᴇs, Oᴛɪᴅɪfᴏʀᴍᴇs, Cᴜᴄᴜʟɪfᴏʀᴍᴇs, Cᴀᴘʀɪᴍᴜʟɢɪfᴏʀᴍᴇs, Sᴛᴇᴀᴛᴏʀɴɪᴛʜɪfᴏʀᴍᴇs, Nʏᴄᴛɪʙɪɪfᴏʀᴍᴇs, Pᴏᴅᴀʀɢɪfᴏʀᴍᴇs, Aᴘᴏᴅɪfᴏʀᴍᴇs". www.kmoksy.com. Retrieved 30 December 2015.
ITS Magazine, autumn 2003 (20), www.turacos.org

Brewer, David (2018). Birds New to Science: Fifty Years of Avian Discoveries. London: Christopher Helm. p. 105. ISBN 978-1-4729-0628-1.

References

Ballmann, Peter (1969). "Les Oiseaux miocènes de la Grive-Saint-Alban (Isère) [The Miocene birds of Grive-Saint-Alban, Isère]". Geobios. 2: 157–204. doi:10.1016/S0016-6995(69)80005-7. (French with English abstract)
Hughes, Janice M.; Baker, Allan J. (1999). "Phylogenetic relationships of the enigmatic hoatzin (Opisthocomus hoazin) resolved using mitochondrial and nuclear gene sequences" (PDF). Molecular Biology and Evolution. 16 (9): 1300–1307. doi:10.1093/oxfordjournals.molbev.a026220. PMID 10486983.
International Turaco Society (Magazines 1993–2012), also website 2001, www.turacos.org
Mlíkovský, Jirí (2002): Cenozoic Birds of the World, Part 1: Europe. Ninox Press, Prague. ISBN 80-901105-3-8 PDF fulltext Archived 2011-05-20 at the Wayback Machine
Newton, Alfred (1911). "Touracou" . In Chisholm, Hugh (ed.). Encyclopædia Britannica. 27 (11th ed.). Cambridge University Press. p. 102. This is based on a now outdated classification, but does provide a detailed description of the morphology of some species.
Sorenson, Michael D.; Oneal, Elen; García-Moreno, Jaime; Mindell, David P. (2003). "More Taxa, More Characters: The Hoatzin Problem is Still Unresolved". Molecular Biology and Evolution. 20 (9): 1484–1499. doi:10.1093/molbev/msg157. PMID 12777516. S2CID 24173060. Supplementary Material

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