Warning: This tree should not be taken as a phylogenetic presentation!
In truth it's a geographic presentation that may or may not have phylogenetic value.
Phylogeny of Paleognathia after Sibley & Ahlquist, 1990
Phylogeny of Paleognathia after Haddrath & Baker, 2001
Phylogeny of Paleognathia after Livezey & Zusi, 2007
<==o Paleognathia Huxley, 1867 (vanhaleukaiset nykylinnut) |?- †Eremopezus eocaenus Andrews, 1904 [Paleognathia incertae sedis] |?- †Stromeria fajumensis Lambrecht, 1929 [Paleognathia incertae sedis] |?-o †Eleutherornithidae [Paleognathia incertae sedis] | |-- †Eleutherornis helveticus | `-- †Proceriavis martini |--o “Oceaniac paleognaths” | |?-o Casuariimorphae Livezey & Zusi, 2007 (“Australian paleognaths”) | | `?-o Casuariformes (Forbes, 1884) (emus and casuars; kasuaarilinnut) | | |?- †“Dromaius” gidju | | |--o Casuariidae Kaup, 1847 (casuars; kasuaarit) | | | `--o Casuarius (casuars; kasuaarit) | | | |?- †C. sp. Rich & van Tets, 1983; L. Plio. Aust. [NG] | | | |--o C. casuarius (Linnaeus, 1758) (kypäräkasuaari) | | | | |-- C. c. casuarius | | | | |-- C. c. bicarunculatus | | | | |-- C. c. tricarunculatus | | | | |-- C. c. lateralis | | | | |-- C. c. sclaterii | | | | |-- C. c. aruensis | | | | |-- C. c. violicollis | | | | `-- C. c. johnsonii | | | |--o C. unappendiculatus Blyth, 1860 (yksipussikasuaari) | | | | |-- C. u. rothschildi | | | | |-- C. u. philipi | | | | |-- C. u. unappendiculatus | | | | |-- C. u. occipitalis | | | | |-- C. u. rufotinctus | | | | `-- C. u. aurentiacus | | | `--o C. bennetti Gould, 1858 (bennettinkasuaari) | | | |-- C. b. papuanus Schlegel, 1871 | | | |-- C. b. goodfellowi | | | |-- C. b. claudii | | | |-- C. b. hecki | | | |-- C. b. picticollis | | | |-- C. b. bennetti | | | `-- C. b. shawmayeri | | `--o Dromaiidae Richmond, 1908 (emus; emut) | | `--o Dromaius Vieillot, 1816 (emut) | | |-- †D. ocypus (AH Miller,, 1963) [Casuarius ocypus AH Miller, 1963] L. Plio. Aust. | | |-- †D. ater Vieillot, 1817 [=? †D. minor] (black emu; mustaemu) | | |-- †D. baudinianus Parker, 1984 (kengurusaarenemu) | | `--o D. novaehollandiae (Latham, 1790) (emu) | | |-- †D. n. diemenensis Le Souef, 1907 | | |-- D. n. woodwardi | | |-- D. n. rothchildi (Mathews, 1912) | | `-- D. n. novaehollandiae (Latham, 1790) | `?-o Apterygimorphae Livezey & Zusi, 2007 (“New Zealandic paleognaths”) | |?-o Apterygiformes (Haeckel, 1866) (kiwis; kiivilinnut) | | `--o Apterygidae Gray, 1840 (kiwis; kiivit) | | `--o Apteryx Shaw, 1813 (kiivit) | | |--o A. australis Shaw, 1813 (ruskokiivi) | | | |-- A. a. australis Shaw, 1813 | | | `-- A. a. lawryi Rothschild, 1893 | | |-- A. mantelli Bartlett, 1852 | | |-- A. owenii Gould, 1847 (pikkukiivi) | | `-- A. haastii Potts, 1872 (kirjokiivi) | `?-o †Dinornithiformes (Gadow, 1893) (moas; moalinnut) | |--o †Dinornithidae (Owen, 1843) (great moas; isomoat) | | `--o †Dinornis Owen, 1843 [taxonomic history] (great moas; isomoat) | | |-- †D. novaezealandiae Owen, 1843 [taxonomic history] (North Island great moa) | | `-- †D. robustus Owen, 1846 [taxonomic history] (South Island great moa) | `--o †Emeidae [Anomalopterygidae (Archey, 1941)] (lesser moas; pikkumoat) | |--o †Emeinae | | |-- †Emeus crassus (Owen, 1846) Reichenbach, 1852 [incl. E. huttonii (Owen, 1846)] [Dinornis crassus Owen, 1846; Dinornis casuarinus Owen, 1846; Dinornis huttonii Owen, 1846; Dinornis major Hutton, 1875; Dinornis rheides Owen, 1851; Emeus casuarinus Oliver, 1930; Emeus huttonii Oliver, 1930; Meionornis Haast, 1874; Mesopteryx Hutton, 1891; Syornis Reichenbach, 1850] | | `--o †Euryapteryx Hutton, 1897(?) | | |-- †E. curtus (Owen, 1846) [incl. †Euryapteryx exilis; Euryapteryx gravis (Owen, 1870)] | | `?- †E. geranoides (Owen, 1848) [Anomalopteryx geranoides Lydekker, 1891; Dinornis expunctus Archey, 1927; Euryapteryx compacta Hutton, 1893; Euryapteryx Kuranui Oliver, 1930; Emeus boothi Rothschild, 1907; Emeus haasti Rothschild, 1907; Emeus parkeri Rothschild, 1907; Palapteryx geranoides Owen, 1848; Euryapteryx geranoides Archey, 1941; Pachyornis mappini Archey, 1941; Pachyornis geranoides] (Mantell's Moa) | `--o †Anomalopteryginae (Archey, 1941) | |-- †Megalapteryx didinus (Owen, 1883) Haast, 1886 [incl. M. benhami Archey, 1941] (Upland Moa) | |-- †Anomalopteryx didiformis [incl. A. oweni] | `--o †Pachyornis Lydekker, 1891 | |-- †P. elephantopus (Owen, 1856) [Dinornis elephantopus Owen, 1856; Dinornis queenslandiae De Vis, 1884; Euryapteryx ponderosus Hutton, 1891; Pachyornis immanus Lydekker, 1891; Pachyornis inhabilis Hutton, 1893; Pachyornis major Hutton, 1875; Pachyornis rothschildi Lydekker, 1892; Pachyornis valgus Hutton, 1893] (Heavy-footed Moa) | |?- †P. australis Oliver, 1949 (Crested Moa) | `-- †P. mappini Archey, 1941 [incl. P. septentrionalis] [=? Pachyornis geranoides (Owen, 1848)] `--+?-o “South American paleognaths” | |?-o Tinamiformes (Huxley, 1872) [Crypturi Goodchild, 1891: Dromaeomorphae (Huxley, 1867):] (tinamous; tinamilinnut) | | `-- Tinamidae Gray, 1840 (tinamous; tinamit) | `?-o Rheiformes (nandus; nandulinnut) | |--o †Opisthodactylidae Ameghino, 1895 [possibly synonymous to Rheidae] | | |?- †Diogenornis fragilis Alvarenga, 1983: U. Pal. SSA. | | `-- †Opisthodactylus patagonicus Ameghino, 1895; U. Mioc. SSA. | `--o Rheidae (Bonaparte, 1853) (rheas; nandut) | |-- †Heterorhea dabbenei Rovereto, 1914; U. Plio. SSA. | |-- †Hinasuri nehuensis Ameghino, 1891; U. Plio. SSA. | `--o Rhea Brisson, 1760 [incl. Pterocnemia GR Gray, 1870] | |--o R. americana (Linnaeus, 1758) (Greater Rhea; amerikannandu) | | |-- R. a. albescens Lynch & Holmberg, 1878 | | |-- R. a. americana (Linnaeus, 1758) | | |-- R. a. araneipes Brodkorb, 1938 | | |-- R. a. intermedia Rothschild & Chubb, 1914 | | `-- R. a. nobilis Brodkorb, 1939 | `--o R. pennata d'Orbigny, 1834 [Rhea darwinii Gould, 1837] (Lesser Rhea; patagoniannandu) | |-- R. p. garleppi (Chubb, 1913) | |-- R. p. pennata d'Orbigny, 1834 (Darwin's Rhea; darwininnandu) | `-- R. p. tarapacensis (Chubb, 1913) (Puna Rhea; punannandu) `?-o Struthionimorphae Livezey & Zusi, 2007 (“Laurasian & African paleognaths”) |?- †Remiornis minor [Remiornithiformes: Remiornithidae] [Palaeognathia incertae sedis] |?-o †Lithornithiformes | `--o †Lithornithidae | |-- †Pseudocrypturus cercanaxius | |-- †Paracathartes howardae | |-- †Promusophaga magnifica Harrison & Walker, 1977 | `--o †Lithornis | |-- †L. celetius | `--+-- †L. plebius | |-- †L. promiscuus | |-- †L. nasi | |-- †L. hookeri | `-- †L. vulturinus |?-o †Aepyornithiformes (Newton, 1884) (elephantbirds; norsulinnut) | `--o †Aepyornithidae Bonaparte, 1853 (elephantbirds; norsulinnut) | |?- †Paraphysornis brasiliensis | |?- †Mullerornis betsilei | `--o †Aepyornis (elephantbirds; norsulinnut) | |-- †A. sp.1 | |-- †A. sp.2 | |-- †A. sp.3 | |-- †A. sp.4 | |-- †A. medius | |-- †A. hildebrandti | `-- †A. maximus `?-o Struthioniformes (Latham, 1790) (ostrichies; strutsilinnut) `--o Struthionidae Vigors, 1825 (ostriches; strutsit) |?- ?†Palaeotis weigelti [Palaeogrus geiseltalensis Lambrecht, 1935] M. Eoc. CEu. `--o Struthio Linnaeus, 1758 (ostriches; strutsit) |-- †S. orlovi |-- †S. brachydactylus |-- †S. wimani |-- †S. chersonensis |-- †S. asiaticus |-- S. molybdophanes Reichenow, 1883 (Masai ostrich; masaistrutsi) `--o S. camelus Linnaeus, 1758 (ostrich; strutsi) |-- S. c. australis Gurney, 1868 (South African ostrich; eteläafrikanstrutsi) |-- †S. c. syriacus Rothschild, 1919 (arabianstrutsi) |-- S. c. camelus Linnaeus, 1758 (afrikanstrutsi) `-- S. c. massaicus Neumann, 1898 (masaistrutsi 2)
After Mitchell, Llamas, Soubrier, Rawlence, Worthy, Wood, Lee & Cooper, 2014<==o Paleognathia |-- Struthioniformes `--+-- Rheiformes `--+--+-- Tinamiformes | `-- †Dinornithiformes `--+-- Casuariformes `--+-- Apterygiformes `-- †Aepyornithiformes Reference(s):
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After Baker, Haddrath, McPherson & Cloutier, 2014<==o Paleognathia |-- Struthioniformes `--+--+-- Tinamiformes | `-- †Dinornithiformes `--+-- Rheiformes `--+-- Casuariformes `-- Apterygiformes Reference(s):
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After Sibley & Ahlquist, 1990<==o Eoaves |-- Galloanserae `--o Ratitae |--o Tinamiformes | `-- Tinamidae `--o Struthioniformes |--o Struthioni | |--o Struthionides | | `-- Struthionidae | `--o Rheides | `-- Rheidae `--o Casuarii |-- Apterygidae `--o Casuariidae |-- Casuariini `-- Dromaiini |
After Haddrath & Baker, 2001<==o Paleognathia |?- †Lithornithiformes |?- †Aepyornithiformes |-- Tinamiformes `--+-- †Dinornithiformes `--+-- Rheiformes `--+-- Struthioniformes `--+-- Apterygiformes `-- Casuariformes |
After Livezey & Zusi, 2007<==o Paleognathae |--o Crypturi | `--o Dromaeomorphae | `--o Tinamiformes | `-- Tinamidae `--o Ratitae `==o Apterygimorphae |--o Apterygiformes | `-- Apterygidae `--+--o †Dinornithiformes | |-- †Anomalopterygidae | `-- †Dinornithidae `--+--o Casuariimorphae | `--o Casuariformes | |-- Casuariidae | `-- Dromaiidae `--o Struthionimorphae `--+--o †Aepyornithiformes | `-- †Aepyornithidae `--o Struthioniformes |-- Struthionidae `-- Rheidae |
Note:
Species of moa, and particulary genus Dinornis seems to have unusually large sexual dimorphism, which has caused different morphs to be classified as separate species (Worthy, 1994). Recent articles of Bunce et al (2003) and Huynen et al (2003) has demostrated this by using molecular methods of taxonomy. Because only the genus Dinornis was revisited, I have not chanced the status of other species of moas. [MKH, 2005]
Reference(s):
- Baker, A. J., Huynen, L. J., Haddrath, O., Millar, C. D. & Lambert, D. M., 2005: Reconstructing the tempo and mode of evolution in an extinct clade of birds with ancient DNA: The giant moas of New Zealand.
–Proceedings of the National Academy of Sciences of the United States of America: Vol. 102, #23, pp. 8257-8262 [doi: 10.1073/pnas.0409435102] - Bunce, M., Worthy, T. H., Ford, T., Hoppitt, W., Willerslev, E., Drummond, A. & Cooper, A., 2003: Extreme reversed sexual size dimorphism in the extinct New Zealand moa Dinornis.
–Nature: Vol. 425, pp. 172-175 - Cracraft, J., 2001: Avian evolution, Gondwana biogeography and the Cretaceous-Tertiary mass extinction event.
–Proceedings of the Royal Society of London: Biological series: Vol. 268, pp. 459-469 - Feduccia, A., 1980: The Age of Birds.
–Harvard University Press, Cambridge, Massachusetts & London, England, 1980, ix-196 - Feduccia, A., 1996: The origin and evolution of birds.
–Yale University Press, New Haven and London, 1996, xii-420 - Holdaway, R. N., Worthy, T. H. & Tennyson, A. J. D., 2001: A working list of breeding bird species of the New Zealand region at first human contact.
–New Zealand Journal of Zoology: Vol. 28, pp. 119-187 - Huynen, L., Millar, C. D., Scofield, R. P. & Lambert, D. M., 2003: Nuclear DNA sequences detect species limits in ancient moa.
–Nature: Vol. 425, pp. 175-178 - Livezey, B. C. & Zusi, R. L., 2007: Higher-order phylogeny of modern birds (Theropoda, Aves: Neornithes) based on comperative anatomy. II. Analysis and discussion.
–Zoological Journal of the Linnean Society: Vol. 149, #1, pp. 1-95 - Mitchell, K. J., Llamas, B., Soubrier, J., Rawlence, N. J., Worthy, T. H., Wood, J., Lee, M. S. Y. & Cooper, A., 2014: Ancient DNA reveals elephant birds and kiwi are sister taxa and clarifies ratite bird evolution.
–Science: Vol. 344, #6186, pp. 898-900 [doi: 10.1126/science.1251981] - Olson, S. L., 1985: The fossil record of birds. 79-239.
in Farner, D., King, J. & Parkes, K. (eds.), 1985: Avian Biology. vol VIII.
–Academic Press, New York. 1985. - Rasmussen, D. T., Olson, S. L. & Simons, E. L., 1987: Fossil birds from the Oligocene Jebel Quatrani Formation, Fayum Province, Egypt.
–Smithsonian Contributions to Paleobiology: No. 62, pp. iv-20 - Sibley, C. G. & Monroe, B. L., 1990: Distribution and taxonomy of birds of the world.
–Yale University Press, New Haven & London, 1990. xxiv-1111. - Worthy, T. H., 1994: Reappraisal of Dinornis (Aves: Dinornithiformes) species – a morphometric analysis.
–New Zealand Journal of Zoology: Vol. 21, pp. 113-134