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Flightless bird

Flightless birds are birds that cannot fly, as they have, through evolution, lost the ability. There are over 60 extant species, including the well-known ratites, and penguins (Sphenisciformes). The smallest flightless bird is the Inaccessible Island rail. The largest flightless bird, which is also the largest living bird in general, is the common ostrich.

History
Origins of flightlessness Divergences and losses of flight within ratite lineage occurred right after the K-Pg extinction event wiped out all non-avian dinosaurs and large vertebrates 66 million years ago. The immediate evacuation of niches following the mass extinction provided opportunities for Palaeognathes to distribute and occupy novel environments. New ecological influences selectively pressured different taxa to converge on flightless modes of existence by altering them morphologically and behaviorally. The successful acquisition and protection of a claimed territory selected for large size and cursoriality in Tertiary ancestors of ratites. Temperate rainforests dried out throughout the Miocene and transformed into semiarid deserts, causing habitats to be widely spread across the growingly disparate landmasses. Cursoriality was an economic means of traveling long distances to acquire food that was usually low-lying vegetation, more easily accessed by walking. Gigantism and flightlessness in birds are almost exclusively correlated due to islands lacking mammalian or reptilian predators and competition. However, ratites occupy environments that are mostly occupied by a diverse number of mammals. It is thought that they first originated through allopatric speciation caused by breakup of the supercontinent Gondwana. Rather ratites arrived in their respective locations via a flighted ancestor and lost the ability to fly multiple times within the lineage. Gigantism is not a requirement for flightlessness. The kiwi do not exhibit gigantism, along with tinamous, even though they coexisted with the moa and rheas that both exhibit gigantism. This could be the result of different ancestral flighted birds arrival or because of competitive exclusion. Independent evolution of flightlessness in Palaeognathes Ratites belong to the superorder Palaeognathae, which include the volant tinamou, and are believed to have evolved flightlessness independently multiple times within their own group. It remains possible that the most recent common ancestor of ratites was flightless and the tinamou regained the ability to fly. However, it is believed that the loss of flight is an easier transition for birds than the loss and regain of flight, which has never been documented in avian history. Moreover, tinamou nesting within flightless ratites indicates ancestral ratites were volant and multiple losses of flight occurred independently throughout the lineage. This indicates that the distinctive flightless nature of ratites is the result of convergent evolution. ==Morphological changes and energy conservation==
Morphological changes and energy conservation
Two key differences between flying and flightless birds are the smaller wing bones of flightless birds and the absent (or greatly reduced) keel on their breastbone, which anchors muscles needed for wing movement. Repeated selection for cursorial traits across ratites suggests these adaptions comprise a more efficient use of energy in adulthood. However, ratite anatomy presents other primitive characters meant for flight, such as the fusion of wing elements, a cerebellar structure, the presence of a pygostyle for tail feathers, and an alula on the wing. Some flightless varieties of island birds are closely related to flying varieties, implying flight is a significant biological cost. A study looking at the basal rates of birds found a significant correlation between low basal rate and pectoral muscle mass in kiwis. On the contrary, flightless penguins exhibit an intermediate basal rate. This is likely because penguins have well-developed pectoral muscles for hunting and diving in the water. Species with certain characteristics are more likely to evolve flightlessness. For example, species that already have shorter wings are more likely to lose flight ability. Some species will evolve flatter wings so that they move more efficiently underwater at the cost of their flight. Additionally, birds that undergo simultaneous wing molt, in which they replace all of the feathers in their wings at once during the year, are more likely to evolve flight loss. A number of bird species appear to be in the process of losing their powers of flight to various extents. These include the Zapata rail of Cuba, the Okinawa rail of Japan, and the Laysan duck of Hawaii. All of these birds show adaptations common to flightlessness, and evolved recently from fully flighted ancestors, but have not yet completely given up the ability to fly. They are, however, weak fliers and are incapable of traveling long distances by air. Continued presence of wings in flightless birds Although selection pressure for flight was largely absent, the wing structure has not been lost except in the New Zealand moas. High parental involvement denotes the necessity for choosing a reliable mate. In a climatically stable habitat providing year-round food supply, a male's claimed territory signals to females the abundance of resources readily available to her and her offspring. Penguins evolved their wing structure to become more efficient underwater at the cost of their efficiency in the air. The only known species of flightless bird in which wings completely disappeared was the gigantic, herbivorous moa of New Zealand, hunted to extinction by humans by the 15th century. In moa, the entire pectoral girdle is reduced to a paired scapulocoracoid, which is the size of a finger. ==List of flightless birds==
List of flightless birds
Many flightless birds are extinct; this list shows species that are either still extant or became extinct in the Holocene (no more than 11,000 years ago). A number of species suspected, but not confirmed to be flightless, are also included here. (†) - Indicates an extinct species. Longer-extinct groups of flightless birds include the Cretaceous patagopterygiformes, hesperornithids, the Cenozoic phorusrhacids ("terror birds") and related bathornithids, the unrelated eogruids, geranoidids, gastornithiforms, and dromornithids (mihirungs or "demon ducks"), and the plotopterids. ===Palaeognathae (ratites)=== ====Struthioniformes (ostriches)==== • Common ostrich, Struthio camelusNorth African ostrich, Struthio camelus camelusSouth African ostrich, Struthio camelus australisMasai ostrich, Struthio camelus massaicusArabian ostrich, Struthio camelus syriacus † • Somali ostrich, Struthio molybdophanesAsian ostrich, Struthio asiaticus † • East Asian ostrich, Struthio anderssoni † ====Casuariiformes (cassowaries and emus)==== • Common emu, Dromaius novaehollandiaeMainland emu, Dromaius novaehollandiae novaehollandiaeKing Island emu, Dromaius novaehollandiae minor † • Kangaroo Island emu, Dromaius novaehollandiae baudinianus † • Tasmanian emu, Dromaius novaehollandiae diemenensis † • Pygmy cassowary, Casuarius lydekkeri † • Dwarf cassowary, Casuarius bennettiBennett's cassowary, Casuarius bennetti bennettiPapuan dwarf cassowary, Casuarius bennetti westermanniSouthern cassowary, Casuarius casuariusNorthern cassowary, Casuarius unappendiculatus ====Dinornithiformes (moa) † ==== • North Island giant moa, Dinornis novaezealandiae † • South Island giant moa, Dinornis robustus † • Bush moa, Anomalopteryx didiformis † • Eastern moa, Emeus crassus † • Broad-billed moa, Euryapteryx curtus † • Heavy-footed moa, Pachyornis elephantopus † • Mantell's moa, Pachyornis geranoides † • Crested moa, Pachyornis australis † • Upland moa, Megalapteryx didinus † ====Aepyornithiformes (elephant birds) † ==== • Hildebrandt's elephant bird, Aepyornis hildebrandti † • Giant elephant bird, Aepyornis maximus † • Lesser elephant bird, Mullerornis modestus † ====Apterygiformes (kiwi)==== • Southern brown kiwi, Apteryx australisStewart Island tokoeka, Apteryx australis lawryiFiordland tokoeka, Apteryx australis australisGreat spotted kiwi, Apteryx haastiiNorth Island brown kiwi, Apteryx mantelliLittle spotted kiwi, Apteryx oweniiNorth Island little spotted kiwi, Apteryx owenii iredalei † • South Island little spotted kiwi, Apteryx owenii oweniiOkarito kiwi, Apteryx rowi ====Rheiformes (rheas)==== • Greater rhea, Rhea americanaAmerican rhea, Rhea americana americanaIntermediate rhea, Rhea americana intermediaArgentine rhea, Rhea americana albescensParaguayan rhea, Rhea americana nobilis Brodkorb's rhea, Rhea americana araneipesLesser rhea, Rhea pennataDarwin's rhea, Rhea pennata pennataGarlepp's rhea, Rhea pennata garleppiPuna rhea, Rhea pennata tarapacensis ===Neognathae=== ====Galliformes (landfowl)==== • New Caledonian giant scrubfowl, Sylviornis neocaledoniae † • Noble megapode, Megavitiornis altirostris † • Viti Levu scrubfowl, Megapodius amissus † ====Anseriformes (waterfowl)==== • Mihirung, Genyornis newtoni † • Amsterdam wigeon, Mareca marecula † • Bermuda flightless duck, Anas pachyscelus † • Auckland Island teal, Anas aucklandicaCampbell teal, Anas nesiotisEaton's pintail, Anas eatoniFinsch's duck, Chenonetta finschi † • Steamer ducksFuegian steamer duck, Tachyeres pteneresFalkland steamer duck, Tachyeres brachypterusChubut steamer duck, Tachyeres leucocephalusMoa-nalo † • Turtle-jawed moa-nalo, Chelychelynechen quassus † • Small-billed moa-nalo, Ptaiochen pau † • O'ahu moa-nalo, Thambetochen xanion † • Maui Nui large-billed moa-nalo, Thambetochen chauliodous † • Nēnē-nui, Branta hylobadistes † (possibly flightless or very weak flier) • Giant Hawaiʻi goose, Branta rhuax † • California flightless sea-duck or Law's diving goose, Chendytes lawi † • Kaua'i mole duck, Talpanas lippa † • New Zealand goose, Cnemiornis gracilis and C. calcitrans † ====Aegotheliformes (owlet-nightjars)==== • New Zealand owlet-nightjar, Aegotheles novaezealandiae † ====Mesitornithiformes (mesites)==== • Brown mesite Mesitornis unicolor (possibly flightless, has not been seen flying) ====Columbiformes (pigeons, doves)==== • Dodo, Raphus cucullatus † • Rodrigues solitaire, Pezophaps solitaria † • Viti Levu giant pigeon, Natunaornis gigoura † • Saint Helena dove, Dysmoropelia dekarchiskos † • Henderson ground dove, Gallicolumba leonpascoi † ====Gruiformes (cranes, rails, and coots)==== • Cuban flightless crane, Grus cubensis † • Red rail, Aphanapteryx bonasia † • Rodrigues rail, Erythromachus leguati † • Woodford's rail, Hypotaenidia woodfordi (most likely flightless) • Bar-winged rail, Hypotaenidia poeciloptera † (probably flightless) • Weka, Gallirallus australisNew Caledonian rail, Gallirallus lafresnayanus (likely †) • Lord Howe woodhen, Gallirallus sylvestrisCalayan rail, Gallirallus calayanensisPink-legged rail, Gallirallus insignisGuam rail, Gallirallus owstoniRoviana rail, Gallirallus rovianae (flightless, or almost so) • Tahiti rail, Gallirallus pacificus † • Dieffenbach's rail, Gallirallus dieffenbachii † • Wake Island rail, Gallirallus wakensis † • numerous other unnamed Gallirallus rails from various Pacific islands • Chatham rail, Cabalus modestus † • Snoring rail, Aramidopsis plateniInvisible rail, Habroptila wallaciiNew Guinea flightless rail, Megacrex ineptaAldabra (white-throated) rail, Dryolimnas (cuvieri) aldabranusRéunion rail, Dryolimnas augusti † • Sauzier's wood rail or Cheke's wood rail, Dryolimnas chekei † • Inaccessible Island rail, Atlantisia rogersiSaint Helena rail, Aphanocrex podarces † • Ascension crake, Mundia elpenor † • Saint Helena crake, Porzana astrictocarpus † • Laysan rail, Porzana palmeri † • Hawaiian rail, Porzana sandwichensis † • Small Maui crake, Porzana keplerorum † • Liliput crake, Porzana menehune † • Great Oʻahu crake, Porzana ralphorum † • Great Maui crake, Porzana severnsi † • Small Oʻahu crake, Porzana ziegleri † • Kosrae crake, Porzana monasa † • Henderson crake, Porzana atraMangaia crake, Porzana rua † • Tahiti crake, Porzana nigra † • numerous other unnamed Porzana crakes from various Pacific islands • Lord Howe swamphen, Porphyrio albus † • North Island takahē, Porphyrio mantelli † • Takahē, Porphyrio hochstetteriSamoan woodhen, Gallinula pacificaMakira woodhen, Gallinula silvestrisTristan moorhen, Gallinula nesiotis † • Gough Island moorhen, Gallinula comeriTasmanian native hen, Tribonyx mortieriiGiant coot, Fulica gigantea (adults only; immature birds can fly) • Hawkins' rail, Diaphorapteryx hawkinsi † • Snipe-rail, Capellirallus karamu †Antillean cave rail, Nesotrochis debooyi † • Hispaniolan cave rail, Nesotrochis steganinos † • Cuban cave rail, Nesotrochis picapicensis † • Adzebills, Aptornis otidiformis and A. defossor † • Auckland rail, Lewinia muelleri (possibly flightless) ====Podicipediformes (grebes)==== • Junín grebe, Podiceps taczanowskiiTiticaca grebe, Rollandia micropteraAtitlán grebe, Podilymbus gigas † (reportedly flightless) ====Charadriiformes (shorebirds and allies)==== † • Great auk, Pinguinus impennis † ====Sphenisciformes (penguins)==== • Emperor penguin, Aptenodytes forsteriKing penguin, Aptenodytes patagonicusAdélie penguin, Pygoscelis adeliaeChinstrap penguin, Pygoscelis antarcticaGentoo penguin, Pygoscelis papuaLittle blue penguin, Eudyptula minorMagellanic penguin, Spheniscus magellanicusHumboldt penguin, Spheniscus humboldtiGalapagos penguin, Spheniscus mendiculusAfrican penguin, Spheniscus demersusYellow-eyed penguin, Megadyptes antipodesWaitaha penguin, Megadyptes waitahaFiordland penguin, Eudyptes pachyrhynchusSnares penguin, Eudyptes robustusErect-crested penguin, Eudyptes sclateriNorthern rockhopper penguin, Eudyptes moseleyiSouthern rockhopper penguin, Eudyptes chrysocomeRoyal penguin, Eudyptes schlegeliMacaroni penguin, Eudyptes chrysolophusChatham penguin, Eudyptes warhami † ====Suliformes (boobies, cormorants and allies)==== • Flightless cormorant, Nannopterum harrisi ====Pelecaniformes (pelicans, herons, ibises and allies)==== • Ascension night heron, Nycticorax olsoni † • Jamaican ibis, Xenicibis xymphithecus † • Hawaiian flightless ibises, Apteribis glenos and A. brevis † ====Strigiformes (owls)==== • Cuban giant owl, Ornimegalonyx spp. † (possibly flightless) • Cretan owl, Athene cretensis † (possibly flightless) • Andros Island barn owl, Tyto pollens † (possibly flightless) ====Bucerotiformes (hornbills and hoopoes)==== • Saint Helena hoopoe, Upupa antaios † ====Falconiformes (falcons and caracaras)==== • Jamaican caracara, Caracara tellustris † ====Psittaciformes (parrots)==== • Kākāpō, Strigops habroptilus ====Passeriformes (perching birds)==== • Lyall's wren, Xenicus lyalli † • Long-billed wren, Dendroscansor decurvirostris † • North Island stout-legged wren, Pachyplichas jagmi † • South Island stout-legged wren, Pachyplichas yaldwyni † • some Scytalopus tapaculos (possibly flightless, never seen flying) • Long-legged bunting, Emberiza alcoveri † ==References==
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