Ratite - Knowledge

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have communal nests, where they share the incubating duties with others. Ostriches, and great spotted kiwis, are the only ratites where the female incubates; they share the duties, with the males incubating at night. Cassowaries and emu are polyandrous, with males incubating eggs and rearing chicks with no obvious contribution from females. Ostriches and rheas are polygynous with each male courting several females. Male rheas are responsible for building nests and incubating while ostrich males incubate only at night. Kiwis stand out as the exception with extended monogamous reproductive strategies where either the male alone or both sexes incubate a single egg. Unlike most birds, male ratites have a

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richness and no mammalian predators, the need for large, powerful flight muscles that make for a quick escape decreases. Moreover, raptor species tend to become generalist predators on islands with low species richness, as opposed to specializing in the predation of birds. An increase in leg size compensates for a reduction in wing length in insular birds that have not lost flight by providing a longer lever to increase force generated during the thrust that initiates takeoff.

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the combination of rapid early radiation of the group and long terminal branches. A morphological analysis that created a basal New Zealand clade has not been corroborated by molecular studies. A 2008 study of nuclear genes shows ostriches branching first, followed by rheas and tinamous, then kiwi splitting from emus and cassowaries. In more recent studies, moas and tinamous were shown to be

1600:, concentrating on broad-leafed plants. However, they will eat insects if the opportunity arises. Emus have tracts of 7 m (23 ft) length, and have a more omnivorous diet, including insects and other small animals. Cassowaries have next to the shortest tracts at 4 m (13 ft). Finally, kiwi have the shortest tracts and eat earthworms, insects, and other similar creatures. 3300: 1253:

require dispersal across oceans by flight, as apparently does the colonization of New Zealand by the moa and possibly the back-dispersal of tinamous to South America, if the latter occurred. The phylogeny as a whole suggests not only multiple independent origins of flightlessness, but also of gigantism (at least five times).

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Antarctica, (by the same route marsupials are thought to have used to reach Australia) and then reached New Zealand and Madagascar via "sweepstakes" dispersals (rare low probability dispersal methods, such as long distance rafting) across the oceans. Gigantism would have evolved subsequent to trans-oceanic dispersals.

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position and among extant ratites, placed rheas in the second most basal position, with Australo-Pacific ratites splitting up last; they have also shown that both the latter groups are monophyletic. Early mitochondrial genetic studies that failed to make ostriches basal were apparently compromised by

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Ratites and humans have had a long relationship starting with the use of the egg for water containers, jewelry, or other art medium. Male ostrich feathers were popular for hats during the 18th century, which led to hunting and sharp declines in populations. Ostrich farming grew out of this need, and

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appear to be the most basal members of the clade. The various ratite lineages were probably descended from flying ancestors that independently colonised South America and Africa from the north, probably initially in South America. From South America they could have traveled overland to Australia via

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While the ratites share a lot of similarities, they also have major differences. Ostriches have only two toes, with one being much larger than the other. Cassowaries have developed long inner toenails, used defensively. Ostriches and rheas have prominent wings; although they do not use them to fly,

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among ratites at least three times. More recent evidence suggests this happened at least six times, or once in each major ratite lineage. Re-evolution of flight in the tinamous would be an alternative explanation, but such a development is without precedent in avian history, while loss of flight is

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Ratites are different from the flying birds in that they needed to adapt or evolve certain features to protect their young. First and foremost is the thickness of the shells of their eggs. Their young are hatched more developed than most and they can run or walk soon thereafter. Also, most ratites

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Kiwi and tinamous are the only palaeognath lineages not to evolve gigantism, perhaps because of competitive exclusion by giant ratites already present on New Zealand and South America when they arrived or arose. The fact that New Zealand has been the only land mass to recently support two major

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split-up of Gondwana followed by continental drift would predict that the deepest phylogenetic split would be between African and all other ratites, followed by a split between South American and Australo-Pacific ratites, roughly as observed. However, the elephant bird–kiwi relation appears to

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created a window of time with large predators absent that may have allowed the ancestors of extant flightless ratites to evolve flightlessness. They subsequently underwent selection for large size. One hypothesis suggests that as predation pressure decreases on islands with low raptor species

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Research on flightless rails indicates the flightless condition evolved in the absence of predators. This shows flight to be generally necessary for survival and dispersal in birds. In apparent contradiction to this, many landmasses occupied by ratites are also inhabited by predatory mammals.

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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.;

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Harshman, J.; Braun, E. L.; Braun, M. J.; Huddleston, C. J.; Bowie, R. C. K.; Chojnowski, J. L.; Hackett, S. J.; Han, K.-L.; Kimball, R. T.; Marks, B. D.; Miglia, K. J.; Moore, W. S.; Reddy, S.; Sheldon, F. H.; Steadman, D. W.; Steppan, S. J.; Witt, C. C.; Yuri, T. (September 2008).

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and use a highly developed sense of smell to find small insects and grubs in the soil. Kiwi are notable for laying eggs that are very large in relation to their body size. A kiwi egg may equal 15 to 20 percent of the body mass of a female kiwi. The smallest species of kiwi is the

671:, cassowary eggs are brought back to villages and the chicks raised for eating as a much-prized delicacy, despite (or perhaps because of) the risk they pose to life and limb. They reach up to 1.8 metres (5 ft 11 in) tall and weigh as much as 85 kilograms (187 lb) 1314:

mass, hollow bones and a light build, et cetera. The basal metabolic rate of flighted species is much higher than that of flightless terrestrial birds. But energetic efficiency can only help explain the loss of flight when the benefits of flying are not critical to survival.

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rather than monophyletic, if we exclude the tinamous. Since tinamous are weak fliers, this raises interesting questions about the evolution of flightlessness in this group. The branching of the tinamous within the ratite radiation suggests flightlessness

839:. Cracraft's 1974 biogeographic vicariance hypothesis suggested that ancestral flightless paleognaths, the ancestors of ratites, were present and widespread in Gondwana during the Late Cretaceous. As the supercontinent fragmented due to 860:

Recent analyses of genetic variation between the ratites do not support this simple picture. The ratites may have diverged from one another too recently to share a common Gondwanan ancestor. Also, the Middle Eocene ratites such as

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Le Duc, D.; Renaud, G.; Krishnan, A.; Almén, M. S.; Huynen, L.; Prohaska, S. J.; Ongyerth, M.; Bitarello, B. D.; Schiöth, H. B.; Hofreiter, M.; Stadler, P. F.; Prüfer, K.; Lambert, D.; Kelso, J.; Schöneberg, T. (23 July 2015).

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Most recently, studies on genetic and morphological divergence and fossil distribution show that paleognaths as a whole probably had an origin in the northern hemisphere. Early Cenozoic northern hemisphere paleognaths such as

1673:(FSIS) began a voluntary, fee-for-service ratite inspection program in 1995 to help the fledgling industry improve the marketability of the meat. A provision in the FY2001 USDA appropriations act (P.L. 106–387) amended the 576:', a vessel which has no keel — in contradistinction to extant flighted birds with a keel). Without this to anchor their wing muscles, they could not have flown even if they had developed suitable wings. Ratites are a 1588:; similarities in adults end with feeding, as they all vary in diet and length of digestive tract, which is indicative of diet. Ostriches, with the longest tracts at 14 m (46 ft), are primarily 1273:. However, various other landmasses such as South America and Europe have supported multiple lineages of flightless ratites that evolved independently, undermining this competitive exclusion hypothesis. 2135:

Sackton, Timothy B.; Grayson, Phil; Cloutier, Alison; Hu, Zhirui; Liu, Jun S.; Wheeler, Nicole E.; Gardner, Paul P.; Clarke, Julia A.; Baker, Allan J.; Clamp, Michele; Edwards, Scott V. (2019-04-05).

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Buffetaut, E.; Angst, D. (November 2014). "Stratigraphic distribution of large flightless birds in the Palaeogene of Europe and its palaeobiological and palaeogeographical implications".

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is next in height, reaching up to 1.9 metres (6 ft 3 in) tall and about 50 kilograms (110 lb). Like the ostrich, it is a fast-running, powerful bird of the open plains and

1352:. Their tail and flight feathers have retrogressed or have become decorative plumes. They have no feather vanes, which means they do not need to oil their feathers, hence they have no 641:

is the largest living ratite. A large member of this species can be nearly 2.8 metres (9 ft 2 in) tall, weigh as much as 156 kilograms (344 lb), and can outrun a horse.

1265:, in which ratites were able to fill vacant herbivorous niches before mammals attained large size. Some authorities, though, have been skeptical of the new findings and conclusions. 3453:

Corfield, J. R.; Wild, J. M.; Hauber, M. E.; Parsons, S.; Kubke, M. F. (2007-11-21). "Evolution of Brain Size in the Palaeognath Lineage, with an Emphasis on New Zealand Ratites".

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branch, followed by rheas, then a clade consisting of moas and tinamous, followed by the final two branches: a clade of emus plus cassowaries and one of elephant birds plus kiwis.

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McNab, B. K.; Ellis, H. I. (November 2006). "Flightless rails endemic to islands have lower energy expenditures and clutch sizes than flighted rails on islands and continents".

885:, and elephant birds were shown to be most closely related to the New Zealand kiwi. Additional support for the latter relationship was obtained from morphological analysis. 1661:. Rhea feathers are popular for dusters, and eggs and meat are used for chicken and pet feed in South America. Ratite hides are popular for leather products like shoes. 888:

The finding that tinamous nest within this group, originally based on twenty nuclear genes and corroborated by a study using forty novel nuclear loci makes 'ratites'

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are known from several million years earlier, and the classification and membership of the Ratitae itself is uncertain. Some of the earliest ratites occur in Europe.

766:. All these species went into decline following the arrival of humans on Madagascar around 2,000 years ago, and were gone by the 17th or 18th century if not earlier. 3035:

Agnolin, F. L. (2016-07-05). "Unexpected diversity of ratites (Aves, Palaeognathae) in the early Cenozoic of South America: palaeobiogeographical implications".

835:, which carried them to their present locations. Supporting this idea, some studies based on morphology, immunology and DNA sequencing reported that ratites are 2402: 843:, they were carried by plate movements to their current positions and evolved into the species present today. The earliest known ratite fossils date to the 2962: 1945:"Moa's Ark or volant ghosts of Gondwana? Insights from nineteen years of ancient DNA research on the extinct moa (Aves: Dinornithiformes) of New Zealand" 461: 2523:

Davies, S.J.J.F. (2003). "Struthioniformes (Tinamous and Ratites)". In Hutchins, Michael; Jackson, Jerome A.; Bock, Walter J.; Olendorf, Donna (eds.).

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The understanding of relationships within the paleognath clade has been in flux. Previously, all the flightless members had been assigned to the order

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with a height of 3.7 metres (12 ft 2 in) and weighing about 230 kilograms (510 lb). They became extinct by A.D. 1400 due to hunting by

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could weigh over 400 kilograms (880 lb) and stand up to 3 metres (9 ft 10 in) tall. Accompanying it were three other species of

2011: 610:, although the genus is thought to have originated in Africa. However, the ostrich order may have evolved in Eurasia. A recent study posits a 3711: 1348:) are almost absent. They have simplified wing skeletons and musculature. Their legs are stronger and do not have air chambers, except the 1269:

lineages of flightless ratites may reflect the near total absence of native mammals, which allowed kiwi to occupy a mammal-like nocturnal

3706: 1487: 1261:; however, a ten-million-year-long window of opportunity for evolution of avian gigantism on continents may have existed following the 2929: 3496:

Alvarenga, H. M. F. (1983). "Uma ave ratitae do Paleoceno Brasileiro: bacia calcária de Itaboraí, Estado do Rio de Janeiro, Brasil".

2803: 2799:"A new Transantarctic relationship: morphological evidence for a Rheidae–Dromaiidae–Casuariidae clade (Aves, Palaeognathae, Ratitae)" 1657:

humans harvested feathers, hides, eggs, and meat from the ostrich. Emu farming also became popular for similar reasons and for their

1262: 470: 3576: 2199:"Complete mitochondrial DNA genome sequences of extinct birds: ratite phylogenetics and the vicariance biogeography hypothesis" 1686: 2479:

suggest palaeognathous affinities of the Geranoididae and other "crane-like" birds from the Eocene of the Northern Hemisphere"

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Mitchell, K. J.; Llamas, B.; Soubrier, J.; Rawlence, N. J.; Worthy, T. H.; Wood, J.; Lee, M. S. Y.; Cooper, A. (23 May 2014).

3559: 3532: 3437: 1944: 1874:"Tinamous and moa flock together: mitochondrial genome sequence analysis reveals independent losses of flight among ratites" 1792: 1608:

were the largest native herbivores in their faunas, far larger than contemporary herbivorous mammals in the latter's case.

2527:. Vol. 8 Birds I Tinamous and Ratites to Hoatzins (2 ed.). Farmington Hills, MI: Gale Group. pp. 56–105. 811:, while the other supposes that the lineages evolved mostly independently and thus elevates the families to order rank ( 3432:. Vol. 8 Birds I Tinamous and Ratites to Hoatzins (2 ed.). Farmington Hills, MI: Gale Group. pp. 53–55. 2532: 1670: 588:. This implies that flightlessness is a trait that evolved independently multiple times in different ratite lineages. 1429: 1356:. They have no separation of pterylae (feathered areas) and apteria (non-feathered areas), and finally, they have 1674: 689:

grows to about 1.4 metres (4 ft 7 in) tall and usually weighs 15 to 40 kilograms (33–88 lb).

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Loss of flight allows birds to eliminate the costs of maintaining various flight-enabling adaptations like high

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Ratites in general have many physical characteristics in common, although many are not shared by the family

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The longstanding story of ratite evolution was that they share a common flightless ancestor that lived in

163: 385: 376: 367: 358: 340: 331: 545: 3213: 2087:"Genomic Support for a Moa-Tinamou Clade and Adaptive Morphological Convergence in Flightless Ratites" 1473: 1396:. Kiwis are exceptions to this trend, and possess proportionally larger brains comparable to those of 1374:

during the breeding season. Emus, cassowaries, and kiwis show some dimorphism, predominantly in size.

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Nilsson, M.A.; Churakov, G.; Sommer, M.; Van Tran, N.; Zemann, A.; Brosius, J.; Schmitz, J. (2010).

2426:"Phylogenomics and Morphology of Extinct Paleognaths Reveal the Origin and Evolution of the Ratites" 3301:"A New Species of Rail from the Solomon-Islands and Convergent Evolution of Insular Flightlessness" 1818: 1736: 349: 3610: 3169:

McNab, B. K. (October 1994). "Energy Conservation and the Evolution of Flightlessness in Birds".

2012:"Ancient DNA reveals elephant birds and kiwi are sister taxa and clarifies ratite bird evolution" 1563: 1337: 439: 421: 17: 2249:

Cooper, A.; Lalueza-Fox, C.; Anderson, S.; Rambaut, A.; Austin, J.; Ward, R. (8 February 2001).

713:, at 0.9 to 1.9 kilograms (2.0–4.2 lb) and 35 to 45 centimetres (14–18 in). 3673: 3621: 3228: 3078:

Tambussi, C.P.; Noriega, J.I.; Gazdzicki, A.; Tatur, A.; Reguero, M.A.; Vizcaino, S.F. (1994).

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to make federal inspection of ratite meat mandatory as of April 2001 (21 U.S.C. 451 et seq.).

3668: 3549: 2634:"Testing gradual and speciational models of evolution in extant taxa: the example of ratites" 2396: 1637: 1529: 1239: 877: 394: 1515: 3353: 3044: 2886: 2751: 2563: 2554: 2437: 2360: 2318: 2262: 2148: 2026: 1780:. Vol. XXVII. Red Lion Court Fleet Street, London UK: Taylor and Francis. p. 570. 1645: 1557: 1311: 659:. Shorter than an emu, but heavier and solidly built, cassowaries prefer thickly vegetated 457: 448: 1459: 1445: 8: 3214:"Patterns of correlated character evolution in flightless birds: a phylogenetic approach" 1549: 710: 3583: 3357: 3048: 2890: 2755: 2567: 2441: 2364: 2322: 2266: 2152: 2030: 1415: 1238:

By 2014, a mitochondrial DNA phylogeny including fossil members placed ostriches on the

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from Central Europe may imply that the "out-of-Gondwana" hypothesis is oversimplified.

797: 310: 158: 147: 2251:"Complete Mitochondrial Genome Sequences of Two Extinct Moas Clarify Ratite Evolution" 1501: 663:

forest. They can be dangerous when surprised or cornered because of their razor-sharp

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lived in New Zealand before the arrival of humans, ranging from turkey-sized to the

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Some extinct ratites might have had odder lifestyles, such as the narrow-billed

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Proceedings of the National Academy of Sciences of the United States of America

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Also native to Australia and the islands to the north are the three species of

592: 525:. They are mostly large, long-necked, and long-legged, the exception being the 412: 403: 272: 246: 3524: 3340:

Wright, Natalie A.; Steadman, David W.; Witt, Christopher C. (26 April 2016).

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Laurin, M.; Gussekloo, S.W.S.; Marjanovic, D.; Legendre, L.; Cubo, J. (2012).

2549: 2450: 2425: 1404:, though evidence for similar advanced cognitive skills is currently lacking. 3690: 3375: 2583: 2380: 2170: 1746: 1624: 1585: 1451: 1291: 950: 686: 674: 598:

have ratites, or did have until the fairly recent past. So did Europe in the

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Members of the four genera of large extant ratites. Clockwise from top left:

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Molecular phylogenies of the ratites have generally placed ostriches in the

3644: 3474: 3393: 3285: 3155: 3118: 3021: 2955:"A Theory on How Flightless Birds Spread Across the World: They Flew There" 2906: 2859: 2783: 2659: 2495: 2474: 2459: 2282: 2232: 2214: 2178: 2137:"Convergent regulatory evolution and loss of flight in paleognathous birds" 2112: 2056: 1971: 1899: 1371: 882: 854: 836: 581: 577: 295: 139: 3114:"Tracking Marsupial Evolution Using Archaic Genomic Retroposon Insertions" 1913:

Fowler, Murray E. (June 1991). "Comparative Clinical Anatomy of Ratites".

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Without exception, ratite chicks are capable of swimming and even diving.

774: 3653: 1845: 1613: 1589: 1357: 1353: 1158: 1125: 849: 779: 762: 693: 678: 559: 526: 55: 1926: 3326: 3190: 2986:"Kiwi genome provides insights into evolution of a nocturnal lifestyle" 2605:

Cracraft, J (October 1974). "Phylogeny and evolution of ratite birds".

2575: 2372: 1597: 1581: 1389: 1385: 1245: 1174: 928: 747: 668: 615: 555: 544:. The modern bird superorder Palaeognathae consists of ratites and the 285: 238: 100: 65: 3466: 2047: 2740:"Phylogenomic evidence for multiple losses of flight in ratite birds" 2706: 2274: 1619: 1401: 1333: 1297: 1279: 1057: 904: 889: 869: 863: 844: 756: 726: 701: 656: 599: 548: 530: 175: 105: 49: 39: 3615: 3342:"Predictable evolution toward flightlessness in volant island birds" 3317: 2192: 2190: 2188: 3638: 3182: 2085:

Baker, A. J.; Haddrath, O.; McPherson, J. D.; Cloutier, A. (2014).

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Catalogue of British Birds in the Collection of the British Museum

853:, a possible early relative of the rhea). However, more primitive 2631: 2185: 1658: 1421: 1397: 1345: 1341: 991: 783: 697: 638: 563: 551: 541: 143: 110: 85: 3406: 2248: 800:

approaches to ratite classification: one combines the groups as

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Akishinonomiya, F.; Willerslev, E.; Hasegawa, M. (2016-12-15).

2349:"A Miocene ostrich fossil from Gansu Province, northwest China" 1641: 1593: 1360: 705: 682: 603: 185: 2834:"Ratite Nonmonophyly: Independent Evidence from 40 Novel Loci" 2084: 2009: 3111: 3077: 1349: 3428:

Bruning, D. F. (2003). "Rheas". In Hutchins, Michael (ed.).

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Smith, J. V.; Braun, E. L.; Kimball, R. T. (January 2012).

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10.1206/0003-0082(2005)491[0001:ANSOTF]2.0.CO;2

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Bourdon, Estelle; De Ricqles, Armand; Cubo, Jorge (2009).

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Leonard, L.; Dyke, G. J.; Van Tuinen, M. (October 2005).

1521: 722: 645: 618:, which may have been ratites, existed in North America. 585: 540:, which is more recently regarded as containing only the 500: 151: 2550:"The evolution of flightlessness: Is history important?" 2422: 2134: 1872:

Phillips MJ, Gibb GC, Crimp EA, Penny D (January 2010).

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they do use them in courtship and predator distraction.

1871: 1825:. Zwaag, The Netherlands: Universal Taxonomic Services 1819:"Systema Naturae 2000 / Taxon: Order Struthioniformes" 558:). Unlike other flightless birds, the ratites have no 2796: 2680: 831:, whose descendants were isolated from each other by 3577:

Agriculture: A Glossary of Terms, Programs, and Laws

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Boletim do Museu Nacional (Rio de Janeiro), Geologia

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Alcheringa: An Australasian Journal of Palaeontology

509: 503: 497: 2930:"Bird evolutionary tree given a shake by DNA study" 2347:Hou, L.; Zhou, Z.; Zhang, F.; Wang, Z. (Aug 2005). 494: 3339: 2831: 2401:: CS1 maint: DOI inactive as of September 2024 ( 1942: 3688: 3548:Farner, Donald S.; King, James R. (2013-09-03). 2976: 2346: 2304: 2302: 2300: 1943:Allentoft, M. E.; Rawlence, N. J. (2012-01-20). 1844:Harshman, John; Brown, Joseph W. (13 May 2010). 2873:Hackett, Shannon J.; et al. (2008-06-27). 2744:Proceedings of the National Academy of Sciences 2674: 2418: 2416: 2414: 2412: 2005: 2003: 2001: 1938: 1936: 1627:, and could imply similar animalivorous diets. 2625: 2308: 1999: 1997: 1995: 1993: 1991: 1989: 1987: 1985: 1983: 1981: 1867: 1865: 1863: 1861: 1859: 760:as well as three species of the smaller genus 580:group; tinamous fall within them, and are the 2297: 2196: 1843: 692:The smallest ratites are the five species of 3446: 3071: 2518: 2516: 2514: 2512: 2510: 2508: 2506: 2409: 2080: 2078: 2076: 2074: 1933: 1777:Catalogue of the Birds in the British Museum 1407: 1340:are underdeveloped. They do not have keeled 3292: 3105: 3028: 2732: 2730: 2728: 2726: 2724: 1978: 1856: 1766: 1764: 1762: 1760: 1758: 1756: 1741:. London, UK: British Museum. p. 133. 1664: 567: 3568: 3547: 3423: 3421: 3419: 3417: 3415: 3333: 3263: 2946: 2683:"A New Specimen of the Fossil Palaeognath 2342: 2340: 2130: 2128: 2126: 2124: 2122: 129: 3519:. Springer Science & Business Media. 3495: 3489: 3383: 3365: 3316: 3257: 3232: 3162: 3145: 3135: 3011: 3001: 2849: 2825: 2816: 2804:Zoological Journal of the Linnean Society 2773: 2763: 2649: 2503: 2494: 2449: 2244: 2242: 2222: 2160: 2102: 2071: 2046: 1952:Annals of Anatomy - Anatomischer Anzeiger 1889: 716: 288:included but traditionally excluded taxa 2721: 2604: 2598: 2466: 1753: 847:epoch about 56 million years ago (e.g., 773: 738:settlers, who arrived around A.D. 1280. 3427: 3412: 3298: 3034: 2872: 2866: 2790: 2337: 2119: 1797:Integrated Taxonomic Information System 1727: 1725: 1723: 1721: 1570: 14: 3689: 3574: 3409:for definitions of the two latin words 2952: 2927: 2921: 2522: 2239: 1912: 1816: 1719: 1717: 1715: 1713: 1711: 1709: 1707: 1705: 1703: 1701: 1687:List of Struthioniformes by population 644:Of the living species, the Australian 3620: 3619: 3168: 1906: 1837: 1810: 1651: 1263:extinction of the non-avian dinosaurs 3611:Websites With Information On Ratites 3514: 3211: 3205: 2547: 2472: 1915:Journal of Zoo and Wildlife Medicine 1790: 1731: 946: 3712:Taxa named by William Plane Pycraft 2197:Haddrath, O.; Baker, A. J. (2001). 1784: 1698: 1640:that is inserted into the female's 1575: 1392:have generally smaller brains than 24: 3707:Extant Thanetian first appearances 3430:Grzimek's Animal Life Encyclopedia 2619:10.1111/j.1474-919X.1974.tb07648.x 2525:Grzimek's Animal Life Encyclopedia 1671:Food Safety and Inspection Service 681:, large fast-running birds of the 25: 3733: 3604: 2687:from the Lower Eocene of Denmark" 1623:, compared to the shorebird-like 1305: 769: 566:— hence the name, from the Latin 2818:10.1111/j.1096-3642.2008.00509.x 2651:10.1111/j.1420-9101.2011.02422.x 2203:Proceedings of the Royal Society 1556: 1542: 1528: 1514: 1500: 1486: 1472: 1458: 1444: 1428: 1414: 490: 162: 53: 3541: 3508: 3400: 2638:Journal of Evolutionary Biology 2541: 2331:10.1016/j.earscirev.2014.07.001 2091:Molecular Biology and Evolution 1817:Brands, Sheila J., ed. (2020). 1675:Poultry Products Inspection Act 1630: 632: 3582:(Report). 2005. Archived from 1580:Ratite chicks tend to be more 1327: 517:) is any of a group of mostly 13: 1: 3455:Brain, Behavior and Evolution 3407:http://www.freedictionary.com 3057:10.1080/03115518.2016.1184898 2483:Acta Palaeontologica Polonica 1774:; Sharpe, R. Bowdler (1895). 1692: 987: 3137:10.1371/journal.pbio.1000436 1850:The Tree of Life Web Project 1366:Ostriches have the greatest 822: 521:birds within the infraclass 7: 3212:Cubo, Arthur (4 May 2001). 1964:10.1016/j.aanat.2011.04.002 1680: 1161:(cassowaries, 3 spp.) 696:from New Zealand. Kiwi are 27:Polyphyletic group of birds 10: 3738: 3278:10.1016/j.cbpa.2006.02.025 2928:Holmes, Bob (2008-06-26). 1336:, or tinamous. First, the 625: 621: 3628: 3551:Avian Biology: Volume III 3525:10.1007/978-3-540-89628-9 3299:Diamond, J. (July 1991). 3128:Public Library of Science 3003:10.1186/s13059-015-0711-4 2953:Zimmer, C. (2014-05-22). 2691:American Museum Novitates 2451:10.1016/j.cub.2016.10.029 1408:Gallery of living species 1172: 1156: 1148: 1123: 1106: 1098: 1090: 1060:(tinamous, 46 spp.) 1055: 1038: 1030: 1022: 1015: 989: 981: 974: 953:(ostriches, 2 spp.) 948: 940: 933: 746:, the "elephant bird" of 721:At least nine species of 591:Most parts of the former 529:, which is also the only 316: 309: 292: 284: 226: 221: 159:Scientific classification 157: 137: 128: 34: 2475:"Hindlimb morphology of 2353:Chinese Science Bulletin 1665:United States regulation 3575:Womach, Jasper (2005). 3367:10.1073/pnas.1522931113 3243:10.1023/a:1011695406277 3171:The American Naturalist 2899:10.1126/science.1157704 2765:10.1073/pnas.0803242105 2548:Roff, Derek A. (1994). 2375:(inactive 2024-09-12). 2205:. Biological Sciences. 2162:10.1126/science.aat7244 2039:10.1126/science.1251981 1596:. They are also mainly 1564:North Island brown kiwi 994:(rheas, 2~3 spp.) 3517:Paleogene Fossil Birds 3266:Comp Biochem Physiol A 2496:10.4202/app.00650.2019 2215:10.1098/rspb.2001.1587 793: 717:Holocene extinct forms 614:origin for the clade. 568: 338:Fissipedes bidactyles 250:(cassowaries and emus) 3669:Paleobiology Database 3515:Mayr, Gerald (2009). 3087:Polish Polar Research 2851:10.1093/sysbio/sys067 2311:Earth-Science Reviews 2104:10.1093/molbev/msu153 1891:10.1093/sysbio/syp079 1321:K–Pg extinction event 1257:in birds tends to be 895:evolved independently 777: 3221:Evolutionary Ecology 2555:Evolutionary Ecology 1571:Behavior and ecology 1437:S. camelus massaicus 1246:Vicariant speciation 1128:(kiwi, 5 spp.) 704:. They nest in deep 475:Struthiornithiformes 3722:Paraphyletic groups 3717:Polyphyletic groups 3358:2016PNAS..113.4765W 3049:2017Alch...41..101A 2891:2008Sci...320.1763H 2885:(5884): 1763–1768. 2756:2008PNAS..10513462H 2750:(36): 13462–13467. 2568:1994EvEco...8..639R 2442:2017CBio...27...68Y 2365:2005ChSBu..50.1808H 2323:2014ESRv..138..394B 2267:2001Natur.409..704C 2153:2019Sci...364...74S 2031:2014Sci...344..898M 1733:Gray, George Robert 1550:Little spotted kiwi 1344:. Their wishbones ( 1177:(emus, 1 sp.) 927: recent 792:, each with its egg 711:little spotted kiwi 677:has two species of 2959:The New York Times 2838:Systematic Biology 2576:10.1007/BF01237847 2373:10.1360/982005-575 1878:Systematic Biology 1793:"Struthioniformes" 1772:Salvadori, Tomasso 1652:Ratites and humans 1536:Great spotted kiwi 1494:Northern cassowary 1480:Southern cassowary 1370:, rheas show some 907:based on Mitchell 794: 148:southern cassowary 3684: 3683: 3622:Taxon identifiers 3561:978-1-4832-6943-6 3534:978-3-540-89628-9 3467:10.1159/000111456 3439:978-0-7876-5784-0 3352:(17): 4765–4770. 2473:Mayr, G. (2019). 2359:(16): 1808–1810. 2261:(6821): 704–707. 2209:(1470): 939–945. 2025:(6186): 898–900. 1236: 1235: 1231: 1230: 1222: 1221: 1213: 1212: 1204: 1203: 1195: 1194: 1186: 1185: 1137: 1136: 1112:(elephant birds) 1078: 1077: 1069: 1068: 1003: 1002: 962: 961: 833:continental drift 743:Aepyornis maximus 731:Dinornis robustus 700:-sized, shy, and 482: 481: 474: 465: 452: 443: 434: 425: 416: 407: 398: 389: 380: 371: 362: 353: 344: 335: 326: 302:Lithornithiformes 277: 269: 260: 256:Aepyornithiformes 251: 243: 235: 123: 122: 16:(Redirected from 3729: 3702:Flightless birds 3677: 3676: 3664: 3663: 3662: 3649: 3648: 3647: 3617: 3616: 3599: 3598: 3596: 3594: 3588: 3581: 3572: 3566: 3565: 3545: 3539: 3538: 3512: 3506: 3505: 3493: 3487: 3486: 3450: 3444: 3443: 3425: 3410: 3404: 3398: 3397: 3387: 3369: 3337: 3331: 3330: 3320: 3296: 3290: 3289: 3261: 3255: 3254: 3236: 3218: 3209: 3203: 3202: 3166: 3160: 3159: 3149: 3139: 3109: 3103: 3102: 3100: 3098: 3084: 3075: 3069: 3068: 3032: 3026: 3025: 3015: 3005: 2980: 2974: 2973: 2971: 2970: 2961:. Archived from 2950: 2944: 2943: 2941: 2940: 2925: 2919: 2918: 2870: 2864: 2863: 2853: 2829: 2823: 2822: 2820: 2794: 2788: 2787: 2777: 2767: 2734: 2719: 2718: 2678: 2672: 2671: 2653: 2629: 2623: 2622: 2602: 2596: 2595: 2545: 2539: 2538: 2520: 2501: 2500: 2498: 2470: 2464: 2463: 2453: 2420: 2407: 2406: 2400: 2392: 2344: 2335: 2334: 2306: 2295: 2294: 2275:10.1038/35055536 2246: 2237: 2236: 2226: 2194: 2183: 2182: 2164: 2132: 2117: 2116: 2106: 2097:(7): 1686–1696. 2082: 2069: 2068: 2050: 2016: 2007: 1976: 1975: 1949: 1940: 1931: 1930: 1910: 1904: 1903: 1893: 1869: 1854: 1853: 1841: 1835: 1834: 1832: 1830: 1814: 1808: 1807: 1805: 1803: 1788: 1782: 1781: 1768: 1751: 1750: 1729: 1576:Feeding and diet 1560: 1546: 1532: 1518: 1504: 1490: 1476: 1462: 1448: 1432: 1418: 1151: 1150: 1101: 1100: 1093: 1092: 1042:Dinornithiformes 1033: 1032: 1025: 1024: 1018: 1017: 984: 983: 977: 976: 943: 942: 936: 935: 924: 923: 901: 900: 809:Struthioniformes 778:Comparison of a 571: 538:Struthioniformes 516: 515: 512: 511: 508: 505: 502: 499: 496: 469: 456: 447: 438: 429: 420: 411: 402: 393: 384: 375: 366: 357: 348: 339: 330: 321: 275: 267: 265:Dinornithiformes 259:(elephant birds) 258: 249: 241: 233: 231:Struthioniformes 222:Groups included 167: 166: 133: 116: 115: 52: 38:Temporal range: 37: 32: 31: 21: 3737: 3736: 3732: 3731: 3730: 3728: 3727: 3726: 3687: 3686: 3685: 3680: 3672: 3667: 3658: 3657: 3652: 3643: 3642: 3637: 3624: 3607: 3602: 3592: 3590: 3586: 3579: 3573: 3569: 3562: 3546: 3542: 3535: 3513: 3509: 3494: 3490: 3451: 3447: 3440: 3426: 3413: 3405: 3401: 3338: 3334: 3318:10.2307/4088088 3297: 3293: 3262: 3258: 3234:10.1.1.115.1294 3216: 3210: 3206: 3167: 3163: 3110: 3106: 3096: 3094: 3082: 3076: 3072: 3033: 3029: 2981: 2977: 2968: 2966: 2951: 2947: 2938: 2936: 2926: 2922: 2871: 2867: 2830: 2826: 2795: 2791: 2735: 2722: 2679: 2675: 2630: 2626: 2603: 2599: 2546: 2542: 2535: 2521: 2504: 2471: 2467: 2430:Current Biology 2421: 2410: 2394: 2393: 2345: 2338: 2307: 2298: 2247: 2240: 2195: 2186: 2147:(6435): 74–78. 2133: 2120: 2083: 2072: 2014: 2008: 1979: 1947: 1941: 1934: 1911: 1907: 1870: 1857: 1846:"Palaeognathae" 1842: 1838: 1828: 1826: 1823:The Taxonomicon 1815: 1811: 1801: 1799: 1789: 1785: 1769: 1754: 1730: 1699: 1695: 1683: 1667: 1654: 1633: 1578: 1573: 1566: 1561: 1552: 1547: 1538: 1533: 1524: 1519: 1510: 1508:Dwarf cassowary 1505: 1496: 1491: 1482: 1477: 1468: 1463: 1454: 1449: 1440: 1433: 1424: 1419: 1410: 1330: 1312:pectoral muscle 1308: 1286:Pseudocrypturus 1232: 1223: 1214: 1205: 1196: 1187: 1138: 1079: 1070: 1004: 963: 912: 841:plate tectonics 825: 772: 719: 635: 630: 628:List of ratites 624: 584:of the extinct 533:extant ratite. 493: 489: 478: 280: 161: 124: 119:Late Cretaceous 114: 113: 108: 103: 98: 93: 88: 83: 78: 73: 68: 63: 58: 47: 46: 28: 23: 22: 15: 12: 11: 5: 3735: 3725: 3724: 3719: 3714: 3709: 3704: 3699: 3682: 3681: 3679: 3678: 3665: 3650: 3634: 3632: 3626: 3625: 3614: 3613: 3606: 3605:External links 3603: 3601: 3600: 3567: 3560: 3540: 3533: 3507: 3500:. Nova Série. 3488: 3445: 3438: 3411: 3399: 3332: 3311:(3): 461–470. 3291: 3272:(3): 628–648. 3256: 3227:(8): 693–702. 3204: 3183:10.1086/285697 3177:(4): 628–648. 3161: 3104: 3070: 3043:(1): 101–111. 3027: 2996:(1): 147–162. 2990:Genome Biology 2975: 2945: 2920: 2865: 2824: 2811:(3): 641–663. 2789: 2720: 2693:(3491): 1–11. 2673: 2644:(2): 293–303. 2624: 2613:(4): 494–521. 2597: 2562:(6): 639–657. 2540: 2534:978-0787657840 2533: 2502: 2465: 2408: 2336: 2296: 2238: 2184: 2118: 2070: 1977: 1932: 1921:(2): 204–227. 1905: 1855: 1836: 1809: 1783: 1752: 1696: 1694: 1691: 1690: 1689: 1682: 1679: 1666: 1663: 1653: 1650: 1632: 1629: 1606:elephant birds 1577: 1574: 1572: 1569: 1568: 1567: 1562: 1555: 1553: 1548: 1541: 1539: 1534: 1527: 1525: 1520: 1513: 1511: 1506: 1499: 1497: 1492: 1485: 1483: 1478: 1471: 1469: 1464: 1457: 1455: 1450: 1443: 1441: 1435:Ostrich herd ( 1434: 1427: 1425: 1420: 1413: 1409: 1406: 1358:palaeognathous 1338:breast muscles 1329: 1326: 1307: 1306:Loss of flight 1304: 1250:plate tectonic 1234: 1233: 1229: 1228: 1225: 1224: 1220: 1219: 1216: 1215: 1211: 1210: 1207: 1206: 1202: 1201: 1198: 1197: 1193: 1192: 1189: 1188: 1184: 1183: 1180: 1179: 1171: 1168: 1167: 1164: 1163: 1155: 1149: 1147: 1144: 1143: 1140: 1139: 1135: 1134: 1131: 1130: 1122: 1119: 1118: 1115: 1114: 1110:Aepyornithidae 1105: 1099: 1097: 1091: 1089: 1085: 1084: 1081: 1080: 1076: 1075: 1072: 1071: 1067: 1066: 1063: 1062: 1054: 1051: 1050: 1047: 1046: 1037: 1031: 1029: 1023: 1021: 1016: 1014: 1010: 1009: 1006: 1005: 1001: 1000: 997: 996: 988: 982: 980: 975: 973: 969: 968: 965: 964: 960: 959: 956: 955: 947: 941: 939: 934: 932: 922: 919: 918: 824: 821: 796:There are two 771: 770:Classification 768: 718: 715: 634: 631: 623: 620: 593:supercontinent 480: 479: 477: 476: 466: 453: 444: 435: 426: 417: 410:Pressirostres 408: 399: 390: 381: 372: 363: 354: 345: 336: 327: 317: 314: 313: 307: 306: 305: 304: 298: 290: 289: 282: 281: 279: 278: 273:Apterygiformes 270: 261: 252: 247:Casuariiformes 244: 236: 227: 224: 223: 219: 218: 213: 209: 208: 203: 199: 198: 193: 189: 188: 183: 179: 178: 173: 169: 168: 155: 154: 135: 134: 126: 125: 109: 104: 99: 94: 89: 84: 79: 74: 69: 64: 59: 54: 36: 26: 9: 6: 4: 3: 2: 3734: 3723: 3720: 3718: 3715: 3713: 3710: 3708: 3705: 3703: 3700: 3698: 3695: 3694: 3692: 3675: 3670: 3666: 3661: 3655: 3651: 3646: 3640: 3636: 3635: 3633: 3631: 3627: 3623: 3618: 3612: 3609: 3608: 3589:on 2011-08-10 3585: 3578: 3571: 3563: 3557: 3553: 3552: 3544: 3536: 3530: 3526: 3522: 3518: 3511: 3503: 3499: 3492: 3484: 3480: 3476: 3472: 3468: 3464: 3460: 3456: 3449: 3441: 3435: 3431: 3424: 3422: 3420: 3418: 3416: 3408: 3403: 3395: 3391: 3386: 3381: 3377: 3373: 3368: 3363: 3359: 3355: 3351: 3347: 3343: 3336: 3328: 3324: 3319: 3314: 3310: 3306: 3302: 3295: 3287: 3283: 3279: 3275: 3271: 3267: 3260: 3252: 3248: 3244: 3240: 3235: 3230: 3226: 3222: 3215: 3208: 3200: 3196: 3192: 3188: 3184: 3180: 3176: 3172: 3165: 3157: 3153: 3148: 3143: 3138: 3133: 3129: 3125: 3121: 3120: 3115: 3108: 3092: 3088: 3081: 3074: 3066: 3062: 3058: 3054: 3050: 3046: 3042: 3038: 3031: 3023: 3019: 3014: 3009: 3004: 2999: 2995: 2991: 2987: 2979: 2965:on 2014-05-23 2964: 2960: 2956: 2949: 2935: 2934:New Scientist 2931: 2924: 2916: 2912: 2908: 2904: 2900: 2896: 2892: 2888: 2884: 2880: 2876: 2869: 2861: 2857: 2852: 2847: 2843: 2839: 2835: 2828: 2819: 2814: 2810: 2806: 2805: 2800: 2793: 2785: 2781: 2776: 2771: 2766: 2761: 2757: 2753: 2749: 2745: 2741: 2733: 2731: 2729: 2727: 2725: 2716: 2712: 2708: 2704: 2700: 2696: 2692: 2688: 2686: 2677: 2669: 2665: 2661: 2657: 2652: 2647: 2643: 2639: 2635: 2628: 2620: 2616: 2612: 2608: 2601: 2593: 2589: 2585: 2581: 2577: 2573: 2569: 2565: 2561: 2557: 2556: 2551: 2544: 2536: 2530: 2526: 2519: 2517: 2515: 2513: 2511: 2509: 2507: 2497: 2492: 2488: 2484: 2480: 2478: 2469: 2461: 2457: 2452: 2447: 2443: 2439: 2435: 2431: 2427: 2419: 2417: 2415: 2413: 2404: 2398: 2390: 2386: 2382: 2378: 2374: 2370: 2366: 2362: 2358: 2354: 2350: 2343: 2341: 2332: 2328: 2324: 2320: 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1626: 1625:lithornithids 1622: 1621: 1616: 1615: 1609: 1607: 1603: 1599: 1595: 1591: 1587: 1586:insectivorous 1583: 1565: 1559: 1554: 1551: 1545: 1540: 1537: 1531: 1526: 1523: 1517: 1512: 1509: 1503: 1498: 1495: 1489: 1484: 1481: 1475: 1470: 1467: 1466:Darwin's rhea 1461: 1456: 1453: 1452:American rhea 1447: 1442: 1438: 1431: 1426: 1423: 1417: 1412: 1411: 1405: 1403: 1399: 1395: 1391: 1387: 1382: 1379: 1375: 1373: 1369: 1364: 1362: 1359: 1355: 1351: 1347: 1343: 1339: 1335: 1325: 1322: 1319:However, the 1316: 1313: 1303: 1300: 1299: 1294: 1293: 1292:Paracathartes 1288: 1287: 1282: 1281: 1274: 1272: 1266: 1264: 1260: 1256: 1251: 1248:based on the 1247: 1243: 1241: 1227: 1226: 1218: 1217: 1209: 1208: 1200: 1199: 1191: 1190: 1182: 1181: 1178: 1176: 1170: 1169: 1166: 1165: 1162: 1160: 1153: 1152: 1146: 1145: 1142: 1141: 1133: 1132: 1129: 1127: 1121: 1120: 1117: 1116: 1113: 1111: 1103: 1102: 1095: 1094: 1087: 1086: 1083: 1082: 1074: 1073: 1065: 1064: 1061: 1059: 1053: 1052: 1049: 1048: 1045: 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The larger 684: 680: 676: 675:South America 672: 670: 666: 662: 658: 653: 651: 647: 642: 640: 629: 619: 617: 613: 609: 605: 601: 597: 594: 589: 587: 583: 579: 575: 570: 565: 561: 557: 553: 550: 547: 543: 539: 534: 532: 528: 524: 523:Palaeognathae 520: 514: 487: 472: 467: 463: 459: 454: 450: 445: 441: 436: 432: 427: 423: 418: 414: 409: 405: 400: 396: 391: 387: 382: 378: 373: 369: 364: 360: 355: 351: 346: 342: 337: 333: 328: 324: 319: 318: 315: 312: 308: 303: 299: 297: 294: 293: 291: 287: 286:Cladistically 283: 274: 271: 266: 262: 257: 253: 248: 245: 240: 237: 232: 229: 228: 225: 220: 217: 216:Palaeognathae 214: 211: 210: 207: 204: 201: 200: 197: 194: 191: 190: 187: 184: 181: 180: 177: 174: 171: 170: 165: 160: 156: 153: 149: 145: 141: 136: 132: 127: 120: 112: 107: 102: 97: 92: 87: 82: 77: 72: 67: 62: 57: 51: 45: 41: 33: 30: 19: 3629: 3591:. Retrieved 3584:the original 3570: 3554:. 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