391:. Flight is the most costly type of locomotion exemplified in the natural world. The energy expenditure required for flight increases proportionally with body size, which is often why flightlessness coincides with body mass. By reducing large pectoral muscles that require a significant amount of overall metabolic energy, ratites decrease their basal metabolic rate and conserve energy. 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. For ground-feeding birds, a cursorial lifestyle is more economical and allows for easier access to dietary requirements. Flying birds have different wing and feather structures that make flying easier, while flightless birds' wing structures are well adapted to their environment and activities, such as diving in the ocean.
545:
444:. Ratites and tinamous are monogamous and mate only a limited number of times per year. 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. Male size also indicates his protective abilities. Similar to the emperor penguin, male ratites incubate and protect their offspring anywhere between 85 and 92 days while females feed. They can go up to a week without eating and survive only off fat stores. The emu has been documented fasting for as long as 56 days. If no continued pressures warrant the energy expenditure to maintain the structures of flight, selection will tend towards these other traits.
340:. Incongruences between ratite phylogeny and Gondwana geological history indicate the presence of ratites in their current locations is the result of a secondary invasion by flying birds. 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.
47:
2312:
83:
2110:
27:
555:
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1167:
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63:
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371:. Their flat sternum is distinct from the typical sternum of flighted birds because it lacks a keel, like a raft. This structure is the place where flight muscles attach and thus allow for powered flight. However, ratite anatomy presents other primitive characters meant for flight, such as the fusion of wing elements, a cerebellar structure, the presence of a
1498:
1418:
394:
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,
434:
for flight was largely absent, the wing structure has not been lost except in the New
Zealand moas. Ostriches are the fastest running birds in the world and emus have been documented running 50 km/h. At these high speeds, wings are necessary for balance and serving as a parachute apparatus to
220:(all extinct) all evolved similar body shapes – long legs, long necks and big heads – but none of them were closely related. Furthermore, they also share traits of being giant, flightless birds with vestigial wings, long legs, and long necks with some of the ratites, although they are not related.
241:
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
439:
in early ancestral ratites and were thus maintained. This can be seen today in both the rheas and ostriches. These ratites utilize their wings extensively for courtship and displays to other males. Sexual selection also influences the maintenance of large body size, which discourages flight. The
383:
and were free to increase in abundance until the population was limited by food and territory. A study looking at energy conservation and the evolution of flightlessness hypothesized intraspecific competition selected for a reduced individual energy expenditure, which is achieved by the loss of
283:
arrival or because of competitive exclusion. The first flightless bird to arrive in each environment utilized the large flightless herbivore or omnivore niche, forcing the later arrivals to remain smaller. In environments where flightless birds are not present, it is possible that after the K/T
246:
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
2687:
Harshman, J.; Braun, E. L.; Braun, M. J.; Huddleston, C. J.; Bowie, R. C.; 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. (2 September 2008).
3152:
Harshman, J.; Braun, E. L.; Braun, M. J.; Huddleston, C. J.; Bowie, R. C.; 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. (2008).
422:. 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.
269:. However, later evidence suggests this hypothesis first proposed by Joel Cracraft in 1974 is incorrect. Rather ratites arrived in their respective locations via a flighted ancestor and lost the ability to fly multiple times within the lineage.
314:) than any other such location. One reason is that until the arrival of humans roughly a thousand years ago, there were no large mammalian land predators in New Zealand; the main predators of flightless birds were larger birds.
2933:
Mitchell, K. J.; Llamas, B.; Soubrier, J.; Rawlence, N. J.; Worthy, T. H.; Wood, J.; Lee, M. S.; Cooper, A. (2014). "Ancient DNA reveals elephant birds and kiwi are sister taxa and clarifies ratite bird evolution".
336:, and are believed to have evolved flightlessness independently multiple times within their own group. Some birds evolved flightlessness in response to the absence of predators, for example on
355:
Adapting to a cursorial lifestyle causes two inverse morphological changes to occur in the skeleto-muscular system: the pectoral apparatus used to power flight is paedorphically reduced while
451:, wings structure is maintained for use in locomotion underwater. Penguins evolved their wing structure to become more efficient underwater at the cost of their efficiency in the air.
94:'s only mammals were bats and seals, resulting in many bird species evolving to fill the open niches. While many of New Zealand's flightless birds are now extinct, some, such as the
544:
192:
Flightlessness has evolved in many different birds independently, demonstrating repeated convergent evolution. There were families of flightless birds, such as the now-extinct
311:
247:
usually low-lying vegetation, more easily accessed by walking. Traces of these events are reflected in ratite distribution throughout semiarid grasslands and deserts today.
486:(no more than 11,000 years ago). Extinct species are indicated with a cross (†). A number of species suspected, but not confirmed to be flightless, are also included here.
3078:
3760:
Huynen, Leon; Suzuki, Takayuki; Ogura, Toshihiko; Watanabe, Yusuke; Millar, Craig D; Hofreiter, Michael; Smith, Craig; Mirmoeini, Sara; Lambert, David M (December 2014).
261:
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
3256:
Nudds, R. L.; Davidson, J. Slove (2010). "A shortening of the manus precedes the attenuation of other wing-bone elements in the evolution of flightlessness in birds".
395:
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.
242:
existence by altering them morphologically and behaviorally. The successful acquisition and protection of a claimed territory selected for large size and
234:
158:(length 12.5 cm, weight 34.7 g). The largest (both heaviest and tallest) flightless bird, which is also the largest living bird in general, is the
3685:
3105:"Multiple nuclear genes and retroposons support vicariance and dispersal of the palaeognaths, and an Early Cretaceous origin of modern birds"
2912:
2601:
348:
Two key differences between flying and flightless birds are the smaller wing bones of flightless birds and the absent (or greatly reduced)
46:
3906:
2591:
363:
across ratites suggests these adaptions comprise a more efficient use of energy in adulthood. The name "ratite" comes from the Latin
3625:
379:
on the wing. These morphological traits suggest some affinities to volant groups. Palaeognathes were one of the first colonizers of
2748:
3828:
Diamond, Jared (July 1991). "A New
Species of Rail from the Solomon Islands and Convergent Evolution of Insular Flightlessness".
965:
279:, even though they coexisted with the moa and rheas that both exhibit gigantism. This could be the result of different ancestral
3361:
Cubo, Jorge; Arthur, Wallace (2000). "Patterns of correlated character evolution in flightless birds: A phylogenetic approach".
3082:
2311:
3701:
Elliott, Kyle H.; Ricklefs, Robert E.; Gaston, Anthony J.; Hatch, Scott A.; Speakman, John R.; Davoren, Gail K. (4 June 2013).
3668:
Ksepka, Daniel T.; Ando, Tatsuro (2011). "Penguins Past, Present, and Future: Trends in the
Evolution of the Sphenisciformes".
3473:
McCall, Robert A.; Nee, Sean; Harvey, Paul H. (July 1998). "The role of wing length in the evolution of avian flightlessness".
2504:
1304:
2823:"Tinamous and moa flock together: Mitochondrial genome sequence analysis reveals independent losses of flight among ratites"
398:
A number of bird species appear to be in the process of losing their powers of flight to various extents. These include the
3414:
Elliott, Kyle H.; Ricklefs, Robert E.; Gaston, Anthony J.; Hatch, Scott A.; Speakman, John R.; Davoren, Gail K. (2013).
3283:
2265:
2256:
285:
182:
3641:
Handford, Paul; Mares, Michael A. (1985). "The mating systems of ratites and tinamous: An evolutionary perspective".
3703:"High flight costs, but low dive costs, in auks support the biomechanical hypothesis for flightlessness in penguins"
3565:
Terrill, Ryan S. (2020-12-01). "Simultaneous Wing Molt as a
Catalyst for the Evolution of Flightlessness in Birds".
3416:"High flight costs, but low dive costs, in auks support the biomechanical hypothesis for flightlessness in penguins"
387:
Some flightless varieties of island birds are closely related to flying varieties, implying flight is a significant
1688:
1379:
330:
189:; the birds were bred to grow massive breast meat that weighs too much for the bird's wings to support in flight.
3927:
3762:"Reconstruction and in vivo analysis of the extinct tbx5 gene from ancient wingless moa (Aves: Dinornithiformes)"
20:
2361:
1726:
155:
454:
The only known species of flightless bird in which wings completely disappeared was the gigantic, herbivorous
947:
558:
1997:
82:
3378:
1893:
1716:
1506:
1448:
1326:
779:
769:
721:
2895:
Palombo, Maria Rita; Moncunill-Solé, Blanca (2023). "Dwarfing and gigantism in quaternary vertebrates".
2525:
2341:
2247:
1468:
1284:
1274:
1248:
583:
574:
554:
3863:
Hunter, Laurie A. (November 1988). "Status of the
Endemic Atitlan Grebe of Guatemala: Is It Extinct?".
1825:
1631:
3004:"Genomic support for a moa-tinamou clade and adaptive morphological convergence in flightless ratites"
1785:
3773:
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2701:
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2007:
1940:
1805:
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2109:
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2165:
1987:
1949:
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1136:
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26:
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3340:
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1883:
1755:
1391:
938:
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741:
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186:
1536:
1215:
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3621:
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3455:
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514:
360:
213:
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3732:
3722:
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3650:
3609:
3574:
3529:
3490:
3445:
3435:
3388:
3324:
3315:
McNab, Brian K. (1994). "Energy
Conservation and the Evolution of Flightlessness in Birds".
3265:
3228:
3184:
3174:
3124:
3116:
3060:
3015:
2959:
2951:
2900:
2834:
2782:
2719:
2709:
2648:
2638:
2283:
2183:
2070:
2033:
1968:
1815:
1641:
1593:
1224:
1187:
1177:
829:
819:
809:
761:
537:
436:
250:
166:
2464:
196:, that evolved to be powerful terrestrial predators. Taking this to a greater extreme, the
181:, respectively, are capable of extended flight. A few particularly bred birds, such as the
173:, have lost the ability to fly for extended periods, although their ancestral species, the
2870:
2292:
2219:
2201:
2120:
2113:
2102:
2082:
1930:
1845:
1745:
1546:
1371:
730:
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467:
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388:
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1706:
99:
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3431:
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3170:
2947:
2705:
2634:
3796:
3761:
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3702:
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3450:
3415:
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3129:
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1338:
1316:
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901:
644:
613:
565:
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498:
494:
337:
275:
is not a requirement for flightlessness. The kiwi do not exhibit gigantism, along with
217:
193:
174:
159:
78:
are the largest extant flightless birds as well as the largest extant birds in general.
66:
3921:
3594:
3269:
3048:
2981:
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2486:
2437:
2385:
2351:
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1854:
1775:
1410:
1234:
1206:
1170:
690:
622:
592:
529:
522:
502:
356:
326:
253:
and flightlessness in birds are almost exclusively correlated due to islands lacking
201:
170:
3502:
3344:
1166:
2619:"Anthropogenic extinctions conceal widespread evolution of flightlessness in birds"
2405:
2377:
2129:
1864:
1835:
1670:
1660:
1611:
1584:
1399:
1148:
1050:
1041:
1032:
1023:
1014:
1005:
986:
510:
407:
209:
197:
51:
30:
3400:
2617:
Sayol, F.; Steinbauer, M. J.; Blackburn, T. M.; Antonelli, A.; Faurby, S. (2020).
2040:
3901:
1765:
1653:
1621:
1108:
1088:
1079:
1070:
1061:
889:
839:
799:
482:; this list shows species that are either still extant or became extinct in the
459:
415:
399:
295:
291:
280:
233:
Divergences and losses of flight within ratite lineage occurred right after the
147:
143:
123:
95:
91:
62:
3159:
Proceedings of the
National Academy of Sciences of the United States of America
2694:
Proceedings of the
National Academy of Sciences of the United States of America
3677:
3494:
3392:
2534:
2395:
2304:
1978:
1958:
1480:
1116:
1101:
997:
849:
479:
359:
leads to enlargement of the pelvic girdle for running. Repeated selection for
284:
Boundary there were no niches for them to fill. They were pushed out by other
262:
3818:
Roots, Clive. Flightless Birds. Westport, CT: Greenwood, 2006. 136-37. Print.
3543:
3287:
3786:
3727:
3440:
3233:
3216:
3179:
3020:
3003:
2955:
2839:
2822:
2787:
2770:
2714:
2090:
2044:
1602:
879:
869:
506:
372:
272:
243:
205:
139:
119:
3805:
3746:
3586:
3551:
3459:
3242:
3198:
3138:
3120:
3029:
2973:
2848:
2796:
2733:
2662:
2643:
1903:
352:
on their breastbone. (The keel anchors muscles needed for wing movement.)
303:
107:
2017:
1516:
1268:
1156:
789:
483:
462:, hunted to extinction by humans by the 15th century. In moa, the entire
435:
help the bird slow down. Wings are hypothesized to have played a role in
266:
1487:
3884:
3849:
3534:
3517:
3336:
1876:
1497:
448:
333:
299:
276:
258:
178:
151:
131:
127:
75:
39:
2964:
3155:"Phylogenomic evidence for multiple losses of flight in ratite birds"
2690:"Phylogenomic evidence for multiple losses of flight in ratite birds"
2566:
419:
322:
254:
3876:
3841:
3578:
3328:
3002:
Baker, A. J.; Haddrath, O.; McPherson, J. D.; Cloutier, A. (2014).
2463:
2310:
2108:
2039:
1496:
1486:
1417:
1416:
1165:
553:
543:
440:
large size of ratites leads to greater access to mates and higher
411:
376:
115:
81:
61:
45:
25:
489:
Longer-extinct groups of flightless birds include the Cretaceous
1556:
1491:
1428:
1421:
403:
368:
307:
103:
3217:"Ratite nonmonophyly: Independent evidence from 40 novel Loci"
2821:
Phillips, M. J.; Gibb, G. C.; Crimp, E. A.; Penny, D. (2010).
2771:"Ratite nonmonophyly: Independent evidence from 40 novel Loci"
661:
652:
455:
135:
87:
126:. There are over 60 extant species, including the well-known
2897:
Reference Module in Earth Systems and Environmental Sciences
318:
Independent evolution of flightlessness in Palaeognathes
3051:(2008). "Phylogeny and Evolution of the Ratite Birds".
19:"Flightless" redirects here. For the record label, see
2816:
2814:
2812:
2810:
2808:
2806:
1336:
California flightless sea-duck or Law's diving goose,
3310:
3308:
3306:
3304:
3210:
3208:
3043:
3041:
3039:
2997:
2995:
2993:
2991:
2928:
2926:
2924:
294:
had more species of flightless birds (including the
3215:Smith, J. V.; Braun, E. L.; Kimball, R. T. (2013).
2864:
2862:
2860:
2858:
2769:Smith, J. V.; Braun, E. L.; Kimball, R. T. (2013).
2749:"Bird evolutionary tree given a shake by DNA study"
265:speciation caused by breakup of the supercontinent
2538:tapaculos (possibly flightless, never seen flying)
3902:TerraNature pages on New Zealand flightless birds
3518:"Ecomorphological variation of the penguin wing"
185:, have become totally flightless as a result of
3707:Proceedings of the National Academy of Sciences
3420:Proceedings of the National Academy of Sciences
3356:
3354:
2871:"On ratites and their interactions with plants"
1405:(possibly flightless, has not been seen flying)
426:Continued presence of wings in flightless birds
3913:Te Ara – the Encyclopedia of New Zealand
344:Morphological changes and energy conservation
42:are a well-known example of flightless birds.
8:
2596:. Westport: Greenwood Press. pp. XIV.
1323:† (possibly flightless or very weak flier)
3795:
3785:
3736:
3726:
3643:Biological Journal of the Linnean Society
3533:
3449:
3439:
3382:
3232:
3188:
3178:
3128:
3019:
2963:
2838:
2786:
2723:
2713:
2652:
2642:
666:Dromaius novaehollandiae novaehollandiae
2558:
521:(mihirungs or "demon ducks"), and the
154:. The smallest flightless bird is the
2336:(pelicans, herons, ibises and allies)
1965:(adults only; immature birds can fly)
165:Many domesticated birds, such as the
7:
3616:. Westport, CT: Greenwood. pp.
2764:
2762:
2682:
2680:
695:Dromaius novaehollandiae diemenensis
685:Dromaius novaehollandiae baudinianus
3103:Haddrath, O.; Baker, A. J. (2012).
2878:Revista Chilena de Historia Natural
1880:crakes from various Pacific islands
3655:10.1111/j.1095-8312.1985.tb00387.x
3516:Haidr, Nadia Soledad (June 2023).
3065:10.1111/j.1474-919X.1974.tb07648.x
2905:10.1016/B978-0-323-99931-1.00012-X
1657:rails from various Pacific islands
14:
3284:"The Bird Site: Flightless Birds"
470:, which is the size of a finger.
110:have survived to the present day.
3270:10.1111/j.1463-6395.2009.00391.x
3109:Proceedings. Biological Sciences
2567:"New Zealand Ecology – Moa"
2523:South Island stout-legged wren,
2514:North Island stout-legged wren,
2307:(boobies, cormorants and allies)
976:South Island little spotted kiwi
966:North Island little spotted kiwi
3079:"New Zealand's Icon:Flightless"
3008:Molecular Biology and Evolution
1702:Dryolimnas (cuvieri) aldabranus
90:. Until the arrival of humans,
1305:Maui Nui large-billed moa-nalo
1117:New Caledonian giant scrubfowl
735:Casuarius bennetti westermanni
675:Dromaius novaehollandiae minor
1:
1698:Aldabra (white-throated) rail
501:("terror birds") and related
312:several other extinct species
2509:Dendroscansor decurvirostris
2392:spp. † (possibly flightless)
2266:Southern rockhopper penguin
2257:Northern rockhopper penguin
932:Apteryx australis australis
870:Hildebrandt's elephant bird
726:Casuarius bennetti bennetti
183:Broad Breasted White turkey
3944:
2747:Holmes, Bob (2008-06-26).
2362:Hawaiian flightless ibises
1689:New Guinea flightless rail
1618:(flightless, or almost so)
1483:(cranes, rails, and coots)
1384:Aegotheles novaezealandiae
1380:New Zealand owlet-nightjar
755:Casuarius unappendiculatus
597:Struthio camelus massaicus
588:Struthio camelus australis
478:Many flightless birds are
375:for tail feathers, and an
18:
3678:10.1002/9781119990475.ch6
2077:† (reportedly flightless)
1636:Gallirallus dieffenbachii
1570:Gallirallus lafresnayanus
1551:Nesoclopeus poecilopterus
1463:Dysmoropelia dekarchiskos
1093:Rhea pennata tarapacensis
1055:Rhea americana araneipes
1028:Rhea americana intermedia
606:Struthio camelus syriacus
367:, raft, a vessel with no
325:belong to the superorder
229:Origins of flightlessness
200:(and their relatives the
58:), also known as the rowi
21:Flightless (record label)
3766:BMC Evolutionary Biology
2012:Nesotrochis picapicensis
1727:Inaccessible Island rail
1589:Gallirallus calayanensis
1543:(most likely flightless)
1309:Thambetochen chauliodous
1279:Chelychelynechen quassus
1131:Megavitornis altirostris
1121:Sylviornis neocaledoniae
1037:Rhea americana albescens
1019:Rhea americana americana
923:Apteryx australis lawryi
794:Anomalopteryx didiformis
774:Dinornis novaezealandiae
657:Dromaius novaehollandiae
579:Struthio camelus camelus
474:List of flightless birds
156:Inaccessible Island rail
3787:10.1186/1471-2148-14-75
3728:10.1073/pnas.1304838110
3567:The American Naturalist
3495:10.1023/A:1006508826501
3441:10.1073/pnas.1304838110
3393:10.1023/A:1011695406277
3317:The American Naturalist
3180:10.1073/pnas.0803242105
2956:10.1126/science.1251981
2715:10.1073/pnas.0803242105
2440:(falcons and caracaras)
2420:(hornbills and hoopoes)
2412:† (possibly flightless)
2402:† (possibly flightless)
2179:Spheniscus magellanicus
2134:Aptenodytes patagonicus
2085:(shorebirds and allies)
1973:Diaphorapteryx hawkinsi
1874:numerous other unnamed
1651:numerous other unnamed
1553:† (probably flightless)
1473:Gallicolumba leonpascoi
1262:Tachyeres leucocephalus
1188:Bermuda flightless duck
1046:Rhea americana nobilis
970:Apteryx owenii iredalei
948:North Island brown kiwi
824:Pachyornis elephantopus
559:North Island brown kiwi
466:is reduced to a paired
35:Aptenodytes patagonicus
3121:10.1098/rspb.2012.1630
2644:10.1126/sciadv.abb6095
2471:
2406:Andros Island barn owl
2356:Xenicibis xymphithecus
2318:
2234:Eudyptes pachyrhynchus
2116:
2048:
2002:Nesotrochis steganinos
1983:Capellirallus karamu †
1908:Porphyrio hochstetteri
1760:Porzana astrictocarpus
1719:or Cheke's wood rail,
1580:Gallirallus sylvestris
1507:Cuban flightless crane
1502:
1494:
1449:Viti Levu giant pigeon
1424:
1253:Tachyeres brachypterus
1173:
919:Stewart Island tokoeka
874:Aepyornis hildebrandti
780:South Island giant moa
770:North Island giant moa
731:Papuan dwarf cassowary
647:(cassowaries and emus)
618:Struthio molybdophanes
561:
551:
162:(2.7 m, 156 kg).
122:, lost the ability to
111:
79:
59:
43:
3608:Roots, Clive (2006).
3522:Journal of Morphology
3234:10.1093/sysbio/sys067
3021:10.1093/molbev/msu153
2869:Noble, J. C. (1991).
2840:10.1093/sysbio/syp079
2788:10.1093/sysbio/sys067
2526:Pachyplichas yaldwyni
2467:
2342:Ascension night heron
2314:
2288:Eudyptes chrysolophus
2248:Erect-crested penguin
2197:Spheniscus mendiculus
2152:Pygoscelis antarctica
2112:
2057:Podiceps taczanowskii
2043:
1998:Hispaniolan cave rail
1780:Porzana sandwichensis
1626:Gallirallus pacificus
1561:Gallirallus australis
1541:Nesoclopeus woodfordi
1531:Erythromachus leguati
1500:
1490:
1469:Henderson ground dove
1420:
1285:Small-billed moa-nalo
1275:Turtle-jawed moa-nalo
1249:Falkland steamer duck
1169:
1084:Rhea pennata garleppi
980:Apteryx owenii owenii
834:Pachyornis geranoides
584:South African ostrich
575:North African ostrich
557:
547:
298:, several species of
235:K-Pg extinction event
85:
65:
49:
29:
16:Birds that cannot fly
3672:. pp. 155–186.
3475:Evolutionary Ecology
3363:Evolutionary Ecology
2480:Strigops habroptilus
2323:Flightless cormorant
2316:Flightless cormorant
2215:Megadyptes antipodes
2188:Spheniscus humboldti
2125:Aptenodytes forsteri
2066:Rollandia microptera
2022:Aptornis otidiformis
1950:Tasmanian native hen
1941:Gough Island moorhen
1926:Gallinula silvestris
1646:Gallirallus wakensis
1616:Gallirallus rovianae
1598:Gallirallus insignis
1521:Aphanapteryx bonasia
1240:Fuegian steamer duck
1198:Auckland Island teal
1075:Rhea pennata pennata
1071:Darwin's lesser rhea
894:Mullerornis modestus
890:Lesser elephant bird
854:Megalapteryx didinus
844:Pachyornis australis
442:reproductive success
329:, which include the
3778:2014BMCEE..14...75H
3719:2013PNAS..110.9380E
3487:1998EvEco..12..569M
3432:2013PNAS..110.9380E
3375:2000EvEco..14..693C
3171:2008PNAS..10513462H
2948:2014Sci...344..898M
2706:2008PNAS..10513462H
2700:(36): 13462–13467.
2635:2020SciA....6.6095S
2542:Long-legged bunting
2450:Caracara tellustris
2426:Saint Helena hoopoe
2327:Nannopterum harrisi
2270:Eudyptes chrysocome
2211:Yellow-eyed penguin
2206:Spheniscus demersus
2166:Little blue penguin
1992:Nesotrochis debooyi
1988:Antillean cave rail
1894:North Island takahē
1740:Aphanocrex podarces
1717:Sauzier's wood rail
1684:Habroptila wallacii
1675:Aramidopsis plateni
1607:Gallirallus owstoni
1566:New Caledonian rail
1453:Natunaornis gigoura
1443:Pezophaps solitaria
1439:Rodrigues solitaire
1403:Mesitornis unicolor
1360:Cnemiornis gracilis
1327:Giant Hawaiʻi goose
1321:Branta hylobadistes
1299:Thambetochen xanion
1258:Chubut steamer duck
1160:, Genyornis newtoni
1137:Viti Levu scrubfowl
957:Little spotted kiwi
910:Southern brown kiwi
880:Giant elephant bird
746:Casuarius casuarius
722:Bennett's cassowary
707:Casuarius lydekkeri
681:Kangaroo Island emu
637:Struthio anderssoni
491:patagopterygiformes
286:herbivorous mammals
239:non-avian dinosaurs
3535:10.1002/jmor.21588
3221:Systematic Biology
2827:Systematic Biology
2775:Systematic Biology
2517:Pachyplichas jagmi
2472:
2319:
2279:Eudyptes schlegeli
2224:Megadyptes waitaha
2175:Magellanic penguin
2143:Pygoscelis adeliae
2117:
2095:Pinguinus impennis
2049:
1954:Tribonyx mortierii
1935:Gallinula nesiotis
1917:Gallinula pacifica
1898:Porphyrio mantelli
1884:Lord Howe swamphen
1790:Porzana keplerorum
1756:Saint Helena crake
1731:Atlantisia rogersi
1711:Dryolimnas augusti
1632:Dieffenbach's rail
1503:
1495:
1425:
1392:Mesitornithiformes
1244:Tachyeres pteneres
1229:Chenonetta finschi
1174:
1141:Megapodius amissus
939:Great spotted kiwi
864:(elephant birds) †
814:Euryapteryx curtus
751:Northern cassowary
742:Southern cassowary
717:Casuarius bennetti
633:East Asian ostrich
627:Struthio asiaticus
562:
552:
432:selection pressure
187:selective breeding
112:
80:
60:
44:
3713:(23): 9380–9384.
3687:978-0-470-65666-2
3426:(23): 9380–9384.
3115:(1747): 4617–25.
2942:(6186): 898–900.
2914:978-0-12-409548-9
2603:978-0-313-33545-7
2590:Roots C. (2006).
2546:Emberiza alcoveri
2446:Jamaican caracara
2346:Nycticorax olsoni
2261:Eudyptes moseleyi
2252:Eudyptes sclateri
2243:Eudyptes robustus
2230:Fiordland penguin
2193:Galapagos penguin
2148:Chinstrap penguin
1826:Small Oʻahu crake
1810:Porzana ralphorum
1806:Great Oʻahu crake
1736:Saint Helena rail
1721:Dryolimnas chekei
1576:Lord Howe woodhen
1459:Saint Helena dove
1433:Raphus cucullatus
1374:(owlet-nightjars)
1356:New Zealand goose
1024:Intermediate rhea
928:Fiordland tokoeka
914:Apteryx australis
884:Aepyornis maximus
862:Aepyornithiformes
784:Dinornis robustus
3935:
3928:Flightless birds
3889:
3888:
3860:
3854:
3853:
3825:
3819:
3816:
3810:
3809:
3799:
3789:
3757:
3751:
3750:
3740:
3730:
3698:
3692:
3691:
3670:Living Dinosaurs
3665:
3659:
3658:
3638:
3632:
3631:
3615:
3612:Flightless Birds
3605:
3599:
3598:
3562:
3556:
3555:
3537:
3513:
3507:
3506:
3470:
3464:
3463:
3453:
3443:
3411:
3405:
3404:
3386:
3358:
3349:
3348:
3312:
3299:
3298:
3296:
3295:
3286:. Archived from
3280:
3274:
3273:
3253:
3247:
3246:
3236:
3212:
3203:
3202:
3192:
3182:
3149:
3143:
3142:
3132:
3100:
3094:
3093:
3091:
3090:
3081:. Archived from
3075:
3069:
3068:
3045:
3034:
3033:
3023:
2999:
2986:
2985:
2967:
2930:
2919:
2918:
2892:
2886:
2885:
2875:
2866:
2853:
2852:
2842:
2818:
2801:
2800:
2790:
2766:
2757:
2756:
2744:
2738:
2737:
2727:
2717:
2684:
2675:
2674:
2656:
2646:
2623:Science Advances
2614:
2608:
2607:
2593:Flightless Birds
2587:
2581:
2580:
2578:
2577:
2563:
2505:Long-billed wren
2489:(perching birds)
2400:Athene cretensis
2366:Apteribis glenos
2297:Eudyptes warhami
2284:Macaroni penguin
2184:Humboldt penguin
2161:Pygoscelis papua
2075:Podilymbus gigas
2034:Podicipediformes
1945:Gallinula comeri
1830:Porzana ziegleri
1820:Porzana severnsi
1816:Great Maui crake
1800:Porzana menehune
1786:Small Maui crake
1665:Cabalus modestus
1642:Wake Island rail
1594:Pink-legged rail
1413:(pigeons, doves)
1346:Kaua'i mole duck
1202:Anas aucklandica
1192:Anas pachyscelus
1178:Amsterdam wigeon
952:Apteryx mantelli
820:Heavy-footed moa
810:Broad-billed moa
762:Dinornithiformes
570:Struthio camelus
538:Struthioniformes
515:gastornithiforms
505:, the unrelated
437:sexual selection
361:cursorial traits
214:gastornithiforms
167:domestic chicken
71:Struthio camelus
3943:
3942:
3938:
3937:
3936:
3934:
3933:
3932:
3918:
3917:
3898:
3893:
3892:
3877:10.2307/1368847
3862:
3861:
3857:
3842:10.2307/4088088
3827:
3826:
3822:
3817:
3813:
3759:
3758:
3754:
3700:
3699:
3695:
3688:
3667:
3666:
3662:
3640:
3639:
3635:
3628:
3607:
3606:
3602:
3564:
3563:
3559:
3515:
3514:
3510:
3472:
3471:
3467:
3413:
3412:
3408:
3384:10.1.1.115.1294
3360:
3359:
3352:
3314:
3313:
3302:
3293:
3291:
3282:
3281:
3277:
3255:
3254:
3250:
3214:
3213:
3206:
3165:(36): 13462–7.
3151:
3150:
3146:
3102:
3101:
3097:
3088:
3086:
3077:
3076:
3072:
3047:
3046:
3037:
3001:
3000:
2989:
2932:
2931:
2922:
2915:
2894:
2893:
2889:
2873:
2868:
2867:
2856:
2820:
2819:
2804:
2768:
2767:
2760:
2746:
2745:
2741:
2686:
2685:
2678:
2616:
2615:
2611:
2604:
2589:
2588:
2584:
2575:
2573:
2565:
2564:
2560:
2555:
2491:
2462:
2442:
2422:
2386:Cuban giant owl
2382:
2338:
2309:
2293:Chatham penguin
2220:Waitaha penguin
2202:African penguin
2170:Eudyptula minor
2121:Emperor penguin
2114:Emperor penguin
2107:
2103:Sphenisciformes
2087:
2083:Charadriiformes
2038:
2008:Cuban cave rail
1963:Fulica gigantea
1931:Tristan moorhen
1888:Porphyrio albus
1846:Henderson crake
1770:Porzana palmeri
1746:Ascension crake
1693:Megacrex inepta
1547:Bar-winged rail
1537:Woodford's rail
1485:
1415:
1396:
1376:
1372:Aegotheliformes
1216:Eaton's pintail
1153:
1113:
1105:
1051:Brodkorb's rhea
1042:Paraguayan rhea
1002:
943:Apteryx haastii
906:
866:
766:
713:Dwarf cassowary
703:Pygmy cassowary
671:King Island emu
649:
602:Arabian ostrich
542:
534:
497:, the Cenozoic
495:hesperornithids
476:
468:scapulocoracoid
464:pectoral girdle
428:
389:biological cost
346:
338:oceanic islands
320:
310:, the moa, and
231:
226:
74:
38:
24:
17:
12:
11:
5:
3941:
3939:
3931:
3930:
3920:
3919:
3916:
3915:
3904:
3897:
3896:External links
3894:
3891:
3890:
3871:(4): 906–912.
3855:
3836:(3): 461–470.
3820:
3811:
3752:
3693:
3686:
3660:
3633:
3626:
3600:
3579:10.1086/711416
3573:(6): 775–784.
3557:
3508:
3481:(5): 569–580.
3465:
3406:
3369:(8): 693–702.
3350:
3329:10.1086/285697
3323:(4): 628–642.
3300:
3275:
3258:Acta Zoologica
3248:
3204:
3144:
3095:
3070:
3059:(4): 494–521.
3049:Cracraft, Joel
3035:
3014:(7): 1686–96.
2987:
2920:
2913:
2887:
2854:
2802:
2758:
2739:
2676:
2609:
2602:
2582:
2557:
2556:
2554:
2551:
2550:
2549:
2539:
2530:
2521:
2512:
2502:
2499:Xenicus lyalli
2490:
2484:
2483:
2482:
2461:
2458:Psittaciformes
2455:
2454:
2453:
2441:
2435:
2434:
2433:
2421:
2418:Bucerotiformes
2415:
2414:
2413:
2403:
2393:
2381:
2375:
2374:
2373:
2359:
2349:
2337:
2334:Pelecaniformes
2331:
2330:
2329:
2308:
2302:
2301:
2300:
2290:
2281:
2272:
2263:
2254:
2245:
2239:Snares penguin
2236:
2227:
2217:
2208:
2199:
2190:
2181:
2172:
2163:
2157:Gentoo penguin
2154:
2145:
2139:Adélie penguin
2136:
2127:
2106:
2100:
2099:
2098:
2086:
2080:
2079:
2078:
2068:
2062:Titicaca grebe
2059:
2037:
2031:
2030:
2029:
2015:
2005:
1995:
1985:
1976:
1966:
1956:
1947:
1938:
1928:
1922:Makira woodhen
1919:
1913:Samoan woodhen
1910:
1901:
1891:
1881:
1872:
1862:
1852:
1843:
1840:Porzana monasa
1833:
1823:
1813:
1803:
1793:
1783:
1773:
1763:
1753:
1750:Mundia elpenor
1743:
1733:
1724:
1714:
1704:
1695:
1686:
1680:Invisible rail
1677:
1668:
1658:
1649:
1639:
1629:
1619:
1609:
1600:
1591:
1582:
1573:
1563:
1554:
1544:
1534:
1527:Rodrigues rail
1524:
1514:
1484:
1478:
1477:
1476:
1466:
1456:
1446:
1436:
1414:
1408:
1407:
1406:
1395:
1389:
1388:
1387:
1375:
1369:
1368:
1367:
1353:
1350:Talpanas lippa
1343:
1339:Chendytes lawi
1334:
1324:
1314:
1313:
1312:
1302:
1295:O'ahu moa-nalo
1292:
1282:
1266:
1265:
1264:
1255:
1246:
1232:
1222:
1213:
1204:
1195:
1185:
1164:
1163:
1152:
1146:
1145:
1144:
1134:
1127:Noble megapode
1124:
1112:
1106:
1104:
1099:
1098:
1097:
1096:
1095:
1086:
1080:Garlepp's rhea
1077:
1059:
1058:
1057:
1048:
1039:
1033:Argentine rhea
1030:
1021:
1010:Rhea americana
1001:
995:
994:
993:
984:
983:
982:
973:
961:Apteryx owenii
954:
945:
936:
935:
934:
925:
905:
902:Apterygiformes
899:
898:
897:
887:
877:
865:
859:
858:
857:
847:
837:
827:
817:
807:
797:
787:
777:
765:
759:
758:
757:
748:
739:
738:
737:
728:
710:
700:
699:
698:
688:
678:
668:
648:
645:Casuariiformes
642:
641:
640:
630:
620:
614:Somali ostrich
611:
610:
609:
599:
590:
581:
566:Common ostrich
549:Common ostrich
541:
535:
533:
527:
475:
472:
427:
424:
345:
342:
319:
316:
281:flighted birds
237:wiped out all
230:
227:
225:
222:
194:Phorusrhacidae
175:red junglefowl
160:common ostrich
118:have, through
67:Common ostrich
15:
13:
10:
9:
6:
4:
3:
2:
3940:
3929:
3926:
3925:
3923:
3914:
3910:
3909:
3905:
3903:
3900:
3899:
3895:
3886:
3882:
3878:
3874:
3870:
3866:
3859:
3856:
3851:
3847:
3843:
3839:
3835:
3831:
3824:
3821:
3815:
3812:
3807:
3803:
3798:
3793:
3788:
3783:
3779:
3775:
3771:
3767:
3763:
3756:
3753:
3748:
3744:
3739:
3734:
3729:
3724:
3720:
3716:
3712:
3708:
3704:
3697:
3694:
3689:
3683:
3679:
3675:
3671:
3664:
3661:
3656:
3652:
3648:
3644:
3637:
3634:
3629:
3627:9780313335457
3623:
3619:
3614:
3613:
3604:
3601:
3596:
3592:
3588:
3584:
3580:
3576:
3572:
3568:
3561:
3558:
3553:
3549:
3545:
3541:
3536:
3531:
3528:(6): e21588.
3527:
3523:
3519:
3512:
3509:
3504:
3500:
3496:
3492:
3488:
3484:
3480:
3476:
3469:
3466:
3461:
3457:
3452:
3447:
3442:
3437:
3433:
3429:
3425:
3421:
3417:
3410:
3407:
3402:
3398:
3394:
3390:
3385:
3380:
3376:
3372:
3368:
3364:
3357:
3355:
3351:
3346:
3342:
3338:
3334:
3330:
3326:
3322:
3318:
3311:
3309:
3307:
3305:
3301:
3290:on 2007-07-13
3289:
3285:
3279:
3276:
3271:
3267:
3263:
3259:
3252:
3249:
3244:
3240:
3235:
3230:
3226:
3222:
3218:
3211:
3209:
3205:
3200:
3196:
3191:
3186:
3181:
3176:
3172:
3168:
3164:
3160:
3156:
3148:
3145:
3140:
3136:
3131:
3126:
3122:
3118:
3114:
3110:
3106:
3099:
3096:
3085:on 2007-08-18
3084:
3080:
3074:
3071:
3066:
3062:
3058:
3054:
3050:
3044:
3042:
3040:
3036:
3031:
3027:
3022:
3017:
3013:
3009:
3005:
2998:
2996:
2994:
2992:
2988:
2983:
2979:
2975:
2971:
2966:
2961:
2957:
2953:
2949:
2945:
2941:
2937:
2929:
2927:
2925:
2921:
2916:
2910:
2906:
2902:
2898:
2891:
2888:
2883:
2879:
2872:
2865:
2863:
2861:
2859:
2855:
2850:
2846:
2841:
2836:
2833:(1): 90–107.
2832:
2828:
2824:
2817:
2815:
2813:
2811:
2809:
2807:
2803:
2798:
2794:
2789:
2784:
2780:
2776:
2772:
2765:
2763:
2759:
2754:
2753:New Scientist
2750:
2743:
2740:
2735:
2731:
2726:
2721:
2716:
2711:
2707:
2703:
2699:
2695:
2691:
2683:
2681:
2677:
2672:
2668:
2664:
2660:
2655:
2650:
2645:
2640:
2636:
2632:
2628:
2624:
2620:
2613:
2610:
2605:
2599:
2595:
2594:
2586:
2583:
2572:
2568:
2562:
2559:
2552:
2547:
2543:
2540:
2537:
2536:
2531:
2528:
2527:
2522:
2519:
2518:
2513:
2510:
2506:
2503:
2500:
2496:
2493:
2492:
2488:
2487:Passeriformes
2485:
2481:
2477:
2474:
2473:
2470:
2466:
2459:
2456:
2451:
2447:
2444:
2443:
2439:
2438:Falconiformes
2436:
2431:
2430:Upupa antaios
2427:
2424:
2423:
2419:
2416:
2411:
2407:
2404:
2401:
2397:
2394:
2391:
2390:Ornimegalonyx
2387:
2384:
2383:
2379:
2376:
2371:
2367:
2363:
2360:
2357:
2353:
2352:Jamaican ibis
2350:
2347:
2343:
2340:
2339:
2335:
2332:
2328:
2324:
2321:
2320:
2317:
2313:
2306:
2303:
2298:
2294:
2291:
2289:
2285:
2282:
2280:
2276:
2275:Royal penguin
2273:
2271:
2267:
2264:
2262:
2258:
2255:
2253:
2249:
2246:
2244:
2240:
2237:
2235:
2231:
2228:
2225:
2221:
2218:
2216:
2212:
2209:
2207:
2203:
2200:
2198:
2194:
2191:
2189:
2185:
2182:
2180:
2176:
2173:
2171:
2167:
2164:
2162:
2158:
2155:
2153:
2149:
2146:
2144:
2140:
2137:
2135:
2131:
2128:
2126:
2122:
2119:
2118:
2115:
2111:
2104:
2101:
2096:
2092:
2089:
2088:
2084:
2081:
2076:
2072:
2071:Atitlán grebe
2069:
2067:
2063:
2060:
2058:
2054:
2051:
2050:
2046:
2042:
2035:
2032:
2027:
2023:
2019:
2016:
2013:
2009:
2006:
2003:
1999:
1996:
1993:
1989:
1986:
1984:
1980:
1977:
1974:
1970:
1969:Hawkins' rail
1967:
1964:
1960:
1957:
1955:
1951:
1948:
1946:
1942:
1939:
1936:
1932:
1929:
1927:
1923:
1920:
1918:
1914:
1911:
1909:
1905:
1902:
1899:
1895:
1892:
1889:
1885:
1882:
1879:
1878:
1873:
1870:
1869:Porzana nigra
1866:
1863:
1860:
1856:
1855:Mangaia crake
1853:
1851:
1847:
1844:
1841:
1837:
1834:
1831:
1827:
1824:
1821:
1817:
1814:
1811:
1807:
1804:
1801:
1797:
1796:Liliput crake
1794:
1791:
1787:
1784:
1781:
1777:
1776:Hawaiian rail
1774:
1771:
1767:
1764:
1761:
1757:
1754:
1751:
1747:
1744:
1741:
1737:
1734:
1732:
1728:
1725:
1722:
1718:
1715:
1712:
1708:
1705:
1703:
1699:
1696:
1694:
1690:
1687:
1685:
1681:
1678:
1676:
1672:
1669:
1666:
1662:
1659:
1656:
1655:
1650:
1647:
1643:
1640:
1637:
1633:
1630:
1627:
1623:
1620:
1617:
1613:
1610:
1608:
1604:
1601:
1599:
1595:
1592:
1590:
1586:
1583:
1581:
1577:
1574:
1571:
1567:
1564:
1562:
1558:
1555:
1552:
1548:
1545:
1542:
1538:
1535:
1532:
1528:
1525:
1522:
1518:
1515:
1512:
1511:Grus cubensis
1508:
1505:
1504:
1501:Takahē stride
1499:
1493:
1489:
1482:
1479:
1474:
1470:
1467:
1464:
1460:
1457:
1454:
1450:
1447:
1444:
1440:
1437:
1434:
1430:
1427:
1426:
1423:
1419:
1412:
1411:Columbiformes
1409:
1404:
1401:
1398:
1397:
1393:
1390:
1385:
1381:
1378:
1377:
1373:
1370:
1365:
1364:C. calcitrans
1361:
1357:
1354:
1351:
1347:
1344:
1341:
1340:
1335:
1332:
1328:
1325:
1322:
1318:
1315:
1310:
1306:
1303:
1300:
1296:
1293:
1290:
1289:Ptaiochen pau
1286:
1283:
1280:
1276:
1273:
1272:
1270:
1267:
1263:
1259:
1256:
1254:
1250:
1247:
1245:
1241:
1238:
1237:
1236:
1235:Steamer ducks
1233:
1230:
1226:
1225:Finsch's duck
1223:
1221:
1217:
1214:
1212:
1211:Anas nesiotis
1208:
1207:Campbell teal
1205:
1203:
1199:
1196:
1193:
1189:
1186:
1183:
1182:Anas marecula
1179:
1176:
1175:
1172:
1171:Campbell teal
1168:
1161:
1158:
1155:
1154:
1150:
1147:
1142:
1138:
1135:
1132:
1128:
1125:
1122:
1118:
1115:
1114:
1110:
1107:
1103:
1100:
1094:
1090:
1087:
1085:
1081:
1078:
1076:
1072:
1069:
1068:
1067:
1063:
1060:
1056:
1052:
1049:
1047:
1043:
1040:
1038:
1034:
1031:
1029:
1025:
1022:
1020:
1016:
1015:American rhea
1013:
1012:
1011:
1007:
1004:
1003:
999:
996:
992:
988:
985:
981:
977:
974:
971:
967:
964:
963:
962:
958:
955:
953:
949:
946:
944:
940:
937:
933:
929:
926:
924:
920:
917:
916:
915:
911:
908:
907:
903:
900:
895:
891:
888:
885:
881:
878:
875:
871:
868:
867:
863:
860:
855:
851:
848:
845:
841:
838:
835:
831:
830:Mantell's moa
828:
825:
821:
818:
815:
811:
808:
805:
804:Emeus crassus
801:
798:
795:
791:
788:
785:
781:
778:
775:
771:
768:
767:
763:
760:
756:
752:
749:
747:
743:
740:
736:
732:
729:
727:
723:
720:
719:
718:
714:
711:
708:
704:
701:
696:
692:
691:Tasmanian emu
689:
686:
682:
679:
676:
672:
669:
667:
663:
660:
659:
658:
654:
651:
650:
646:
643:
638:
634:
631:
628:
624:
623:Asian ostrich
621:
619:
615:
612:
607:
603:
600:
598:
594:
593:Masai ostrich
591:
589:
585:
582:
580:
576:
573:
572:
571:
567:
564:
563:
560:
556:
550:
546:
539:
536:
531:
530:Palaeognathae
528:
526:
524:
520:
519:dromornithids
516:
512:
508:
504:
503:bathornithids
500:
499:phorusrhacids
496:
492:
487:
485:
481:
473:
471:
469:
465:
461:
457:
452:
450:
445:
443:
438:
433:
425:
423:
421:
417:
413:
409:
405:
401:
396:
392:
390:
385:
382:
378:
374:
370:
366:
362:
358:
357:peramorphosis
353:
351:
343:
341:
339:
335:
332:
328:
327:Palaeognathae
324:
317:
315:
313:
309:
305:
301:
297:
293:
289:
287:
282:
278:
274:
270:
268:
264:
260:
256:
252:
248:
245:
240:
236:
228:
223:
221:
219:
218:dromornithids
215:
211:
207:
203:
202:bathornithids
199:
195:
190:
188:
184:
180:
176:
172:
171:domestic duck
168:
163:
161:
157:
153:
149:
145:
141:
137:
133:
129:
125:
121:
117:
109:
105:
101:
97:
93:
89:
84:
77:
72:
68:
64:
57:
53:
48:
41:
36:
32:
31:King penguins
28:
22:
3912:
3907:
3868:
3864:
3858:
3833:
3829:
3823:
3814:
3769:
3765:
3755:
3710:
3706:
3696:
3669:
3663:
3646:
3642:
3636:
3611:
3603:
3570:
3566:
3560:
3525:
3521:
3511:
3478:
3474:
3468:
3423:
3419:
3409:
3366:
3362:
3320:
3316:
3292:. Retrieved
3288:the original
3278:
3261:
3257:
3251:
3227:(1): 35–49.
3224:
3220:
3162:
3158:
3147:
3112:
3108:
3098:
3087:. Retrieved
3083:the original
3073:
3056:
3052:
3011:
3007:
2939:
2935:
2896:
2890:
2881:
2877:
2830:
2826:
2781:(1): 35–49.
2778:
2774:
2752:
2742:
2697:
2693:
2626:
2622:
2612:
2592:
2585:
2574:. Retrieved
2570:
2561:
2545:
2533:
2524:
2515:
2508:
2498:
2495:Lyall's wren
2479:
2449:
2429:
2410:Tyto pollens
2409:
2399:
2389:
2378:Strigiformes
2369:
2365:
2355:
2345:
2326:
2296:
2287:
2278:
2269:
2260:
2251:
2242:
2233:
2223:
2214:
2205:
2196:
2187:
2178:
2169:
2160:
2151:
2142:
2133:
2130:King penguin
2124:
2094:
2074:
2065:
2056:
2025:
2021:
2011:
2001:
1991:
1982:
1972:
1962:
1953:
1944:
1934:
1925:
1916:
1907:
1897:
1887:
1875:
1868:
1865:Tahiti crake
1858:
1850:Porzana atra
1849:
1839:
1836:Kosrae crake
1829:
1819:
1809:
1799:
1789:
1779:
1769:
1759:
1749:
1739:
1730:
1720:
1710:
1707:Réunion rail
1701:
1692:
1683:
1674:
1671:Snoring rail
1664:
1661:Chatham rail
1652:
1645:
1635:
1625:
1615:
1612:Roviana rail
1606:
1597:
1588:
1585:Calayan rail
1579:
1569:
1560:
1550:
1540:
1530:
1520:
1510:
1472:
1462:
1452:
1442:
1432:
1402:
1400:Brown mesite
1383:
1363:
1359:
1349:
1337:
1331:Branta rhuax
1330:
1320:
1308:
1298:
1288:
1278:
1261:
1252:
1243:
1228:
1219:
1210:
1201:
1191:
1181:
1159:
1149:Anseriformes
1140:
1130:
1120:
1092:
1083:
1074:
1066:Rhea pennata
1065:
1054:
1045:
1036:
1027:
1018:
1009:
1006:Greater rhea
991:Apteryx rowi
990:
987:Okarito kiwi
979:
969:
960:
951:
942:
931:
922:
913:
893:
883:
873:
853:
843:
833:
823:
813:
803:
793:
783:
773:
754:
745:
734:
725:
716:
706:
694:
684:
674:
665:
662:Mainland emu
656:
636:
626:
617:
605:
596:
587:
578:
569:
523:plotopterids
488:
477:
453:
446:
429:
408:Okinawa rail
397:
393:
386:
381:novel niches
364:
354:
347:
321:
290:
271:
249:
244:cursoriality
232:
198:terror birds
191:
164:
113:
70:
56:Apteryx rowi
55:
52:Okarito kiwi
34:
3264:: 115–122.
2571:TerraNature
2053:Junín grebe
2026:A. defossor
1859:Porzana rua
1766:Laysan rail
1654:Gallirallus
1622:Tahiti rail
1220:Anas eatoni
1151:(waterfowl)
1109:Galliformes
1062:Lesser rhea
840:Crested moa
800:Eastern moa
540:(ostriches)
511:geranoidids
460:New Zealand
416:Laysan duck
400:Zapata rail
292:New Zealand
210:geranoidids
140:cassowaries
114:Flightless
92:New Zealand
86:An extinct
3865:The Condor
3649:: 77–104.
3294:2007-08-27
3089:2007-08-27
2965:2328/35953
2576:2007-08-27
2553:References
2535:Scytalopus
2396:Cretan owl
2305:Suliformes
2105:(penguins)
1979:Snipe-rail
1959:Giant coot
1572:(likely †)
1481:Gruiformes
1111:(landfowl)
1102:Neognathae
998:Rheiformes
850:Upland moa
653:Common emu
414:, and the
263:allopatric
3772:(1): 75.
3595:225249314
3544:0362-2525
3379:CiteSeerX
2982:206555952
2884:: 85–118.
2671:227261010
2460:(parrots)
2370:A. brevis
2091:Great auk
2045:Great auk
2018:Adzebills
1603:Guam rail
1394:(mesites)
1089:Puna Rhea
532:(ratites)
430:Although
373:pygostyle
273:Gigantism
259:reptilian
255:mammalian
251:Gigantism
132:ostriches
120:evolution
76:Ostriches
3922:Category
3806:24885927
3747:23690614
3587:33211563
3552:37183492
3503:37855732
3460:23690614
3345:86511951
3243:22831877
3199:18765814
3139:22977150
3030:24825849
2974:24855267
2849:20525622
2797:22831877
2734:18765814
2663:33268368
2036:(grebes)
1517:Red rail
1317:Nēnē-nui
1269:Moa-nalo
1157:Mihirung
790:Bush moa
507:eogruids
484:Holocene
449:penguins
384:flight.
300:penguins
277:tinamous
267:Gondwana
206:eogruids
152:penguins
40:Penguins
3885:1368847
3850:4088088
3830:The Auk
3797:4101845
3774:Bibcode
3738:3677478
3715:Bibcode
3483:Bibcode
3451:3677478
3428:Bibcode
3371:Bibcode
3337:2462941
3190:2533212
3167:Bibcode
3130:3479725
2944:Bibcode
2936:Science
2725:2533212
2702:Bibcode
2654:7710364
2631:Bibcode
1877:Porzana
1000:(rheas)
764:(moa) †
480:extinct
334:tinamou
323:Ratites
224:History
179:mallard
128:ratites
3883:
3848:
3804:
3794:
3745:
3735:
3684:
3624:
3593:
3585:
3550:
3542:
3501:
3458:
3448:
3401:951896
3399:
3381:
3343:
3335:
3241:
3197:
3187:
3137:
3127:
3028:
2980:
2972:
2911:
2847:
2795:
2732:
2722:
2669:
2661:
2651:
2629:(49).
2600:
2476:Kākāpō
2469:Kākāpō
2380:(owls)
1904:Takahē
904:(kiwi)
517:, and
420:Hawaii
406:, the
331:volant
306:, the
304:takahē
302:, the
216:, and
150:) and
146:, and
108:takahē
106:, and
100:kākāpō
3881:JSTOR
3846:JSTOR
3620:–37.
3591:S2CID
3499:S2CID
3397:S2CID
3341:S2CID
3333:JSTOR
2978:S2CID
2874:(PDF)
2667:S2CID
2532:some
412:Japan
377:alula
365:ratis
148:kiwis
144:rheas
116:birds
3908:Kiwi
3802:PMID
3743:PMID
3682:ISBN
3622:ISBN
3583:PMID
3548:PMID
3540:ISSN
3456:PMID
3239:PMID
3195:PMID
3135:PMID
3053:Ibis
3026:PMID
2970:PMID
2909:ISBN
2845:PMID
2793:PMID
2730:PMID
2659:PMID
2598:ISBN
2368:and
2024:and
1557:Weka
1492:Weka
1429:Dodo
1422:Dodo
1362:and
404:Cuba
369:keel
350:keel
308:weka
296:kiwi
177:and
169:and
136:emus
104:weka
96:kiwi
3911:in
3873:doi
3838:doi
3834:108
3792:PMC
3782:doi
3733:PMC
3723:doi
3711:110
3674:doi
3651:doi
3618:136
3575:doi
3571:196
3530:doi
3526:284
3491:doi
3446:PMC
3436:doi
3424:110
3389:doi
3325:doi
3321:144
3266:doi
3229:doi
3185:PMC
3175:doi
3163:105
3125:PMC
3117:doi
3113:279
3061:doi
3057:116
3016:doi
2960:hdl
2952:doi
2940:344
2901:doi
2835:doi
2783:doi
2720:PMC
2710:doi
2698:105
2649:PMC
2639:doi
458:of
456:moa
447:In
418:of
410:of
402:of
257:or
204:),
124:fly
88:moa
50:An
3924::
3879:.
3869:90
3867:.
3844:.
3832:.
3800:.
3790:.
3780:.
3770:14
3768:.
3764:.
3741:.
3731:.
3721:.
3709:.
3705:.
3680:.
3647:25
3645:.
3589:.
3581:.
3569:.
3546:.
3538:.
3524:.
3520:.
3497:.
3489:.
3479:12
3477:.
3454:.
3444:.
3434:.
3422:.
3418:.
3395:.
3387:.
3377:.
3367:14
3365:.
3353:^
3339:.
3331:.
3319:.
3303:^
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