Palaeognathae

Reconstruction of Lithornis, copyright Sheatherius.

Belongs within: Ornithurae.
Contains: Struthio, Apteryx, Aepyornithidae, Casuariiformes, Dinornithiformes, Tinamidae.

The Palaeognathae are a clade of birds found primarily on the southern continents, including the ratites and tinamous. Members of this clade are united by the lack of fusion between the maxillar process of the nasal bone and the maxillary bone, as well as the flat dorsal surface and presence of a pair of furrows on the ventral surface of the mandibular symphysis (Mayr 2009).

Palaeognathae (see below for synonymy)
    |--+--+--Struthio WD17
    |  |  `--Palaeotis Lambrecht 1928 H86, M02 [Palaeotididae M05]
    |  |       `--*P. weigelti Lambrecht 1928 (see below for synonymy) M02
    |  `--+--+--+--Apteryx PB15
    |     |  |  `--Aepyornithidae ML14
    |     |  `--+--Casuariiformes WD17
    |     |     `--+--Dinornithiformes WD17
    |     |        `--Tinamidae WD17
    |     `--Rheidae [Opisthodactylidae, Rheiformes] BKB15
    |          |  i. s.: Opisthodactylus patagonicus Ameghino 1891 AH03
    |          |         Heterorhea dabbeni Rovereto 1914 U93
    |          |--Diogenornis fragilis Alvarenga 1983 ML14, M09
    |          `--+--Pterocnemia pennata L81
    |             `--Rhea Brisson 1760 B94
    |                  |--R. americana JT12
    |                  `--R. pennata JT12
    `--Lithornithidae [Eoaves, Lithornithidae, Lithornithiformes] WD17
         |--Pseudocrypturus Houde 1988 M05
         |    `--P. cercanaxius Houde 1988 M09
         |--Pediorallus Harrison & Walker 1977 M05
         |    `--*P. barbarae Harrison & Walker 1977 M02
         |--Promusophaga Harrison & Walker 1977 M05
         |    `--*P. meini Harrison & Walker 1977 M02
         |--Fissuravis Mayr 2007 M07
         |    `--*F. weigelti Mayr 2007 M07
         `--Lithornis Owen 1840 M05
              |  i. s.: *L. vulturinus Owen 1840 [incl. Promusophaga magnifica Harrison & Walker 1977] M02
              |         L. celetius Houde 1988 M09
              |         L. hookeri (Harrison 1984) [=Pediorallus hookeri] M02
              |         L. nasi (Harrison 1984) M09 [=Pediorallus nasi M02]
              |--L. promiscuus Houde 1988 WD17, M09
              `--+--L. plebius Houde 1988 WD17, M07
                 `--Paracathartes Harrison 1979 WD17, M07
                      `--P. howardae (Harrison 1979) M09

Palaeognathae incertae sedis:
  Remiornis Lemoine 1881 M02 [Remiornithidae M05, Remiornithiformes]
    `--*R. heberti Lemoine 1881 M02
  Eleutherornithidae [Eleutherornithini] U93
    |--Eleutherornis Schaub 1940 M02
    |    `--*E. helveticus Schaub 1940 M02
    `--Proceriavis Harrison & Walker 1979 U93, M02
         `--*P. maritini Harrison & Walker 1979 M02
  Eremopezus Andrews 1904 ML14, RS01 [Eremopezidae, Eremopezithes]
    `--*E. eocaenus Andrews 1904 [incl. Stromeria fajumensis Lambrecht 1929, S. fayumensis] RS01
  Stromeria Lambrecht 1929 U93

Inorganic: Rhea americana minilorientalis Okamura 1987 O87

Palaeognathae [Apteryges, Apterygimorphae, Brevipennes, Palaeognathiformes, Panpalaeognathae, Panratitae, Ratitae, Ratiti, Struthiones, Struthioni, Struthioniformes, Struthionimorphae, Struthionoidea, Timamomorphae]

*Palaeotis weigelti Lambrecht 1928 [incl. Paleogrus geiseltalensis Lambrecht 1935, Ornitocnemus geiseltalensis] M02

*Type species of generic name indicated

REFERENCES

[AH03] Alvarenga, H. M. F., & E. Höfling. 2003. Systematic revision of the Phorusrhacidae (Aves: Ralliformes). Papéis Avulsos de Zoologia 43 (4): 55–91.

[B94] Bock, W. J. 1994. History and nomenclature of avian family-group names. Bulletin of the American Museum of Natural History 222: 1–281.

[BKB15] Burleigh, J. G., R. T. Kimball & E. L. Braun. 2015. Building the avian tree of life using a large-scale, sparse supermatrix. Molecular Phylogenetics and Evolution 84: 53–63.

[H86] Houde, P. 1986. Ostrich ancestors found in the Northern Hemisphere suggest new hypothesis of ratite origins. Nature 324: 563–565.

[JT12] Jetz, W., G. H. Thomas, J. B. Joy, K. Hartmann & A. Ø. Mooers. 2012. The global diversity of birds in space and time. Nature 491: 444–448.

[L81] Long, J. L. 1981. Introduced Birds of the World: The worldwide history, distribution and influence of birds introduced to new environments. Reed: Sydney.

[M05] Mayr, G. 2005. The Paleogene fossil record of birds in Europe. Biological Reviews 80: 515–542.

[M07] Mayr, G. 2007. The birds from the Paleocene fissure filling of Walbeck (Germany). Journal of Vertebrate Paleontology 27 (2): 394-408.

[M09] Mayr, G. 2009. Paleogene Fossil Birds. Springer.

[ML14] Mitchell, K. J., B. Llamas, J. Soubrier, N. J. Rawlence, T. H. Worthy, J. Wood, M. S. Y. Lee & A. Cooper. 2014. Ancient DNA reveals elephant birds and kiwi are sister taxa and clarifies ratite bird evolution. Science 344 (6186): 898–900.

[M02] Mlíkovský, J. 2002. Cenozoic Birds of the World. Part 1: Europe. Ninox Press: Praha.

[O87] Okamura, C. 1987. New facts: Homo and all Vertebrata were born simultaneously in the former Paleozoic in Japan. Original Report of the Okamura Fossil Laboratory 15: 347–573.

[PB15] Prum, R. O., J. S. Berv, A. Dornburg, D. J. Field, J. P. Townsend, E. M. Lemmon & A. R. Lemmon. 2015. A comprehensive phylogeny of birds (Aves) using targeted next-generation DNA sequencing. Nature 526: 569–573.

[RS01] Rasmussen, D. T., E. L. Simons, F. Hertel & A. Judd. 2001. Hindlimb of a giant terrestrial bird from the Upper Eocene, Fayum, Egypt. Palaeontology 44 (2): 325–337.

[U93] Unwin, D. M. 1993. Aves. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 717–737. Chapman & Hall: London.

[WD17] Worthy, T. H., F. J. Degrange, W. D. Handley & M. S. Y. Lee. 2017. The evolution of giant flightless birds and novel phylogenetic relationships for extinct fowl (Aves, Galloanseres). Royal Society Open Science 4: 170975.

Dinornithiformes

Reconstruction of heavy-footed moa Pachyornis elephantopus, copyright Paul Martinson.

Belongs within: Palaeognathae.

The Dinornithiformes, moas, are a group of moderately to extremely large flightless birds known from New Zealand (an historical record of a species from Australia was based on a specimen of mislabelled provenance) that became extinct following human colonisation of the region.

Characters (from Worthy & Holdaway 2002): Skull with small orbits, large olfactory cavities, broad across occipital region. Basitemporal platform with prominent basipterygoid processes. Basiparasphenoidal rostrum long, with triangular expansions extending laterally beneath antorbital plate. Maxilla usually with large dorsal inflation enclosing cavity (maxillary antrum). Beak short, may be pointed or rounded, curved down at tip. 21–23 cervical, six dorsal, 18 sacral, and eleven caudal vertebrae, no pygostyle. Pelvis broad at and posterior to acetabulum. Wing entirely absent, pectoral girdle represented only by vestigial scapulocoracoid bone without glenoid facet. Sternum with well-developed lateral processes. Femur with concave articular surface for antitrochanter on pelvis, tibiotarsus with osseous bridge over tendinal groove, tarsometatarsus with two hypotarsal ridges.

<==Dinornithiformes [Anomalopterygidae, Anomalopteryginae, Dinornithes, Dinornithoidea, Emeidae]
    |--Megalapteryx Haast 1886 BH05, CC10 [incl. Palaeocasuarius Rothschild 1907 CC10]
    |    |--M. didinus (Owen 1883) (see below for synonymy) CC10
    |    `--M. benhami Archey 1941 BKB15, CC10
    `--+--Dinornis Owen 1843 BH05, CC10 (see below for synonymy)
       |    |--*D. novaezealandiae Owen 1843 CC10 (see below for synonymy)
       |    `--D. robustus Owen 1846 (see below for synonymy) CC10
       `--+--Anomalopteryx Reichenbach 1853 ML14, CC10 [=Anomalornis Hutton 1897; incl. Graya Bonaparte 1856]
          |    `--*A. didiformis (Owen 1843) (see below for synonymy) CC10
          `--+--Pachyornis Lydekker 1891 ML14, CC10 [incl. Mauiornis Oliver 1949 WH02, Pounamua Oliver 1949 WH02]
             |    |--P. geranoides (Owen 1848) ML14, CC10 (see below for synonymy)
             |    `--+--*P. elephantopus (Owen 1856) CC10, BKB15, CC10 (see below for synonymy)
             |       `--P. australis Oliver 1949 BKB15, CC10
             `--Emeinae CC10
                  |--Emeus Reichenbach 1853 BH05, CC10 (see below for synonymy)
                  |    `--*E. crassus (Owen 1846) (see below for synonymy) CC10
                  `--Euryapteryx Haast 1874 (see below for synonymy) CC10
                       |--*E. gravis (Owen 1870) CC10 (see below for synonymy)
                       `--E. curtus (Owen 1846) (see below for synonymy) CC10

*Anomalopteryx didiformis (Owen 1843) [=Dinornis didiformis, *Anomalornis didiformis; incl. Anomalopteryx antiqua Hutton 1893, Anomalopteryx antiquus Hutton 1892, Dinornis dromaeoides Owen 1844, Anomalopteryx dromaeoides, *Graya dromaeoides, D. dromioides Owen 1846, D. (Palapteryx) dromioides, Anomalopteryx fortis Hutton 1893, Anomalornis gracilis Hutton 1897, Dinornis oweni Haast 1886, Anomalornis owenii, Pachyornis owenii, Dinornis parvus Owen 1883, Anomalopteryx parva, Anomalopteryx parvus] CC10

Dinornis Owen 1843 BH05, CC10 [=Megalornis Owen 1843 non Gray 1841 CC10; incl. Moa Reichenbach 1853 CC10, Movia Reichenbach 1853 CC10, Owenia Gray 1855 CC10, Palapteryx Owen 1846 CC10, Tylopteryx Hutton 1891 CC10; Dinornithidae]

*Dinornis novaezealandiae Owen 1843 CC10 [=D. novaezelandiae CC10, *Megalornis novaezealandiae CC10; incl. D. excelsus Hutton 1891 CC10, D. firmus Hutton 1891 CC10, D. gazella Oliver 1949 CC10, D. giganteus Owen 1843 CC10, *Moa gigantea CC10, D. gigas Owen 1846 CC10, D. gracilis Owen 1854 CC10, *Tylopteryx gracilis CC10, D. hercules Oliver 1949 CC10, D. ingens Owen 1844 CC10, *Movia ingens CC10, *Palapteryx ingens CC10, D. struthioides Lydekker 1891 CC10, D. struthoides Owen 1844 CC10, *Owenia struthoides CC10, Tylopteryx struthoides WH02]

Dinornis robustus Owen 1846 [=D. ingens var. robustus, Palapteryx robustus; incl. D. altus Owen 1879, D. maximus Haast 1869, Palapteryx plenus Hutton 1891, D. potens Hutton 1891, D. strenuus Hutton 1893, D. torosus Hutton 1891, D. validus Hutton 1891] CC10

Emeus Reichenbach 1852 BH05, CC10 [incl. Meionornis Haast 1874 CC10, Mesopteryx Hutton 1891 CC10, Syornis Reichenbach 1853 CC10]

*Emeus crassus (Owen 1846) [=Dinornis crassus, Euryapteryx crassa, Eu. crassus, Syornis crassus; incl. Dinornis casuarinus Owen 1846, Anomalopteryx casuarina, Cela casuarinus, Emeus casuarinus, *Meionornis casuarinus, Mesopteryx casuarina, *Syornis casuarinus, Dinornis huttoni Owen 1879, Emeus huttonii, Megalapteryx huttoni, *Mesopteryx huttoni, Dinornis rheides Owen 1870, Cela rheides] CC10

Euryapteryx Haast 1874 [incl. Cela Reichenbach 1853 non Moehring 1758, Celeus Bonaparte 1856 non Boie 1831, Zelornis Oliver 1949] CC10

Euryapteryx curtus (Owen 1846) [=Dinornis curtus, Anomalopteryx curta, A. curtus, *Cela curtus, *Celeus curtus; incl. Euryapteryx exilis Hutton 1897, *Zelornis exilis, E. tane Oliver 1949] CC10

*Euryapteryx gravis (Owen 1870) CC10 [=Dinornis gravis CC10; incl. Emeus boothi Rothschild 1907 CC10, Euryapteryx boothi WH02, Eu. compacta Hutton 1893 CC10, Emeus gravipes Lydekker 1891 CC10, Eu. gravipes CC10, Emeus haasti Rothschild 1907 CC10, Zelornis haasti CC10, Euryapteryx kuranui Oliver 1930 CC10, Emeus parkeri Rothschild 1907 CC10, Euryapteryx pygmaeus Hutton 1891 CC10, Pachyornis pygmaeus CC10]

Megalapteryx didinus (Owen 1883) [=Dinornis didinus, Anomalopteryx didina, Meionornis didinus, Mesopteryx didina, Mes. didinus; incl. Palaeocasuarius elegans Rothschild 1907, *P. haasti Rothschild 1907, Megalapteryx hamiltoni Rothschild 1907, *M. hectori Haast 1886, M. tenuipes Lydekker 1891, Palaeocasuarius velox Rothschild 1907] CC10

*Pachyornis elephantopus (Owen 1856) CC10, BKB15, CC10 [=Dinornis elephantopus CC10, Euryapteryx elephantopus CC10; incl. D. elephantopus var. major Hutton 1875 CC10, Pachyornis immanis Lydekker 1891 CC10, Euryapteryx immanis CC10, Pachyornis inhabilis Hutton 1893 CC10, Dinornis elephantopus var. major Hutton 1875 CC10, Pachyornis (Pounamua) murihiku Oliver 1949 WH02, Euryapteryx ponderosus Hutton 1891 CC10, E. ponderosa CC10, Dinornis queenslandiae De Vis 1884 CC10, Dromiceius queenslandiae CC10, Pachyornis queenslandiae CC10, Pachyornis rothschildi Lydekker 1892 CC10, Pachyornis valgus Hutton 1893 CC10]

Pachyornis geranoides (Owen 1848) ML14, CC10 [=Palapteryx geranoides CC10, Anomalopteryx geranoides CC10, Cela geranoides CC10, Dinornis geranoides CC10, Euryapteryx geranoides CC10, Dinornis expunctus Archey 1927 CC10; incl. Pachyornis mappini Archey 1941 BKB15, CC10, Pac. (Mauiornis) mappini CC10, Pac. (*M.) septentrionalis Oliver 1949 CC10]

*Type species of generic name indicated

REFERENCES

[BH05] Baker, A. J., L. J. Huynen, O. Haddrath, C. D. Millar & D. M. Lambert. 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 USA 102 (23): 8257–8262.

[BKB15] Burleigh, J. G., R. T. Kimball & E. L. Braun. 2015. Building the avian tree of life using a large-scale, sparse supermatrix. Molecular Phylogenetics and Evolution 84: 53–63.

[CC10] Checklist Committee (OSNZ). 2010. Checklist of the Birds of New Zealand, Norfolk and Macquarie Islands, and the Ross Dependency, Antarctica 4^th ed. Ornithological Society of New Zealand and Te Papa Press: Wellington.

[ML14] Mitchell, K. J., B. Llamas, J. Soubrier, N. J. Rawlence, T. H. Worthy, J. Wood, M. S. Y. Lee & A. Cooper. 2014. Ancient DNA reveals elephant birds and kiwi are sister taxa and clarifies ratite bird evolution. Science 344 (6186): 898–900.

[WH02] Worthy, T. H., & R. N. Holdaway. 2002. The Lost World of the Moa: Prehistoric life of New Zealand. Indiana University Press: Bloomington (Indiana).

Neognathae

Hoatzin Opisthocomus hoazin, copyright Francesco Veronesi.

Belongs within: Ornithurae.
Contains: Pelagornithidae, Galloanseres, Mirandornithes, Gruiformes, Charadriiformes, Caprimulgiformes, Columbaves, Telluraves, Phaethontiformes, Aequornithia.

The Neognathae are one of the primary clades of living birds, excluding only the ratites and tinamous. Members of the clade are characterised by an intrapterygoidal joint allowing greater mobility of the palate, as well as by a conjoint auditory tube and caudally closed ilioischiadic foramina in the pelvis (Mayr 2009). Within the Neognathae, most recent studies agree on a basal division between the galloanserines (gamefowl and waterfowl) and remaining species in the clade Neoaves. Basal relationships within the Neoaves remain highly contentious, with many large-scale analyses differing significantly in their results; some authors have even suggested that the base of Neoaves may be an insoluble hard polytomy. Particularly contentious has been the position of the hoatzin Opisthocomus hoazin, a medium-sized folivorous bird found in the Amazon basin. The Neotropical sunbittern Eurypyga helias and the New Caledonian kagu Rhynochetos jubatus together form another isolated clade that may be related to the tropicbirds of the Phaethontidae.

Neognathae [Amblypterae, Ciconii, Insessores, Odontoanserae, Pullastrae] HK08
    |--+--Pelagornithidae M11
    |  `--Galloanseres PB15
    `--Neoaves (see below for synonymy) HK08
         |  i. s.: Protopelicanus Reichenbach 1852 M05, M02
         |           `--*P. cuvieri Reichenbach 1852 M02
         |         Onychopteryx [Onychopterygidae] M09
         |           `--O. simpsoni Cracraft 1971 M09
         |--+--Juncitarsus Olson & Feduccia 1980 M05, M02 [Juncitarsinae]
         |  |    |--*J. gracillimus Olson & Feduccia 1980 M02
         |  |    `--J. merkeli Peters 1987 M02
         |  |--Mirandornithes KF-V21
         |  `--Palaelodidae [Palaeolontidae] M09
         |       |--Adelalopus hoogbutseliensis Mayr & Smith 2002 M09
         |       |--Palaelodus Milne-Edwards 1863 BV-R98 [=Paloelodus (l. c.) M02; incl. Probalearica Lambrecht 1933 M02]
         |       |    |--*P. ambiguus Milne-Edwards 1869 BV-R98 (see below for synonymy)
         |       |    |--P. crassipes Milne-Edwards 1863 BV-R98, M02
         |       |    |--P. gracilipes Milne-Edwards 1863 BV-R98, M02 (see below for synonymy)
         |       |    |--P. pledgei Baird & Vickers-Rich 1998 BV-R98
         |       |    `--P. wilsoni Baird & Vickers-Rich 1998 BV-R98
         |       `--Megapaloelodus Miller 1944 M02
         |            |--*M. connectens Miller 1944 M02
         |            |--M. goliath (Milne-Edwards 1868) BV-R98, M02 (see below for synonymy)
         |            `--M. opsigonus Brodkorb 1961 U93
         `--+--+--+--Gruiformes KF-V21
            |  |  `--Charadriiformes KF-V15
            |  `--+--+--Caprimulgiformes KF-V15
            |     |  `--Opisthocomidae [Opisthocomi, Opisthocomiformes] KF-V15
            |     |       |--Hoazinoides magdalenae Miller 1953 U93
            |     |       |--Hoazinavis lacustris JM14
            |     |       `--Opisthocomus Illiger 1811 PB15, B94
            |     |            `--O. hoazin (Müller 1776) ME04
            |     `--Columbaves KF-V15
            `--+--Telluraves KF-V15
               `--+--Phaethontimorphae JM14
                  |    |--Phaethontiformes KF-V15
                  |    `--Eurypygiformes [Eurypygae] JM14
                  |         |--Rhynochetos Verreaux & des Murs 1860 (see below for synonymy) L98
                  |         |    |--R. jubatus L98
                  |         |    `--R. orarius Balouet & Olson 1989 U93
                  |         `--Eurypyga Illiger 1811 [Eurypygidae, Eurypygides] L98
                  |              `--E. helias JM14
                  |                   |--E. h. helias RN72
                  |                   |--E. h. major RN72
                  |                   `--E. h. meridionalis RN72
                  `--Aequornithia KF-V21

Inorganic: Opisthocomidus opisthocomidus minilorientalus Okamura 1987 O87

Megapaloelodus goliath (Milne-Edwards 1868) BV-R98, M02 [=Palaelodus goliath M02, Paloelodus (l. c.) goliath M02]

Neoaves [Aequorlitornithes, Anomalogonates, Charadriimorphae, Ciconiae, Ciconiiformes, Ciconiimorphae, Ciconioidea, Coccyges, Columbea, Coronaves, Cuculimorphae, Cursorimorphae, Dendrornithes, Eurypygoidea, Gaviae, Gavii, Gaviomorphae, Grallae, Impennes, Incessores, Inopinaves, Longipennes, Metaves, Oxypterae, Paludicolae, Passerea, Passeres, Pelecani, Picae, Plethornithes, Psittacimorphae, Pygopodes, Pygopodotubinares, Ralliformes, Simplicirostres, Terrestrornithes, Turniciformes] HK08

*Palaelodus ambiguus Milne-Edwards 1869 BV-R98 [incl. Grus miocenicus Grigorescu & Kessler 1977 M02, G. problematica Milne-Edwards 1871 M02, *Probalearica problematica M02]

Palaelodus gracilipes Milne-Edwards 1863 BV-R98, M02 [incl. incl. Paloelodus (l. c.) minutus Milne-Edwards 1868 BV-R98, M02]

Rhynochetos Verreaux & des Murs 1860 [Rhynocheti, Rhynochetidae, Rhynochetides, Rhynochetoidea] L98

*Type species of generic name indicated

REFERENCES

[BV-R98] Baird, R. F., & P. Vickers-Rich. 1998. Palaelodus (Aves: Palaelodidae) from the Middle to Late Cainozoic of Australia. Alcheringa 22 (2): 135–151.

[B94] Bock, W. J. 1994. History and nomenclature of avian family-group names. Bulletin of the American Museum of Natural History 222: 1–281.

[HK08] Hackett, S. J., R. T. Kimball, S. Reddy, R. C. K. Bowie, E. L. Braun, M. J. Braun, J. L. Chojnowski, W. A. Cox, K.-L. Han, J. Harshman, C. J. Huddleston, B. D. Marks, K. J. Miglia, W. S. Moore, F. H. Sheldon, D. W. Steadman, C. C. Witt & T. Yuri. 2008. A phylogenomic study of birds reveals their evolutionary history. Science 320: 1763–1768.

[JM14] Jarvis, E. D., S. Mirarab, A. J. Aberer, B. Li, P. Houde, C. Li, S. Y. W. Ho, B. C. Faircloth, B. Nabholz, J. T. Howard, A. Suh, C. C. Weber, R. R. de Fonseca, J. Li, F. Zhang, H. Li, L. Zhou, N. Narula, L. Liu, G. Ganapathy, B. Boussau, M. S. Bayzid, V. Zavidovych, S. Subramanian, T. Gabaldón, S. Capella-Gutiérrez, J. Huerta-Cepas, B. Rekepalli, K. Munch, M. Schierup, B. Lindow, W. C. Warren, D. Ray, R. E. Green, M. W. Bruford, X. Zhan, A. Dixon, S. Li, N. Li, Y. Huang, E. P. Derryberry, M. F. Bertelsen, F. H. Sheldon, R. T. Brumfield, C. V. Mello, P. V. Lovell, M. Wirthlin, M. P. Cruz Schneider, F. Prosdocimi, J. A. Samaniego, A. M. Vargas Velazquez, A. Alfaro-Núñez, P. F. Campos, B. Petersen, T. Sitcheritz-Ponten, A. Pas, T. Bailey, P. Scofield, M. Bunce, D. M. Lambert, Q. Zhou, P. Perelman, A. C. Driskell, B. Shapiro, Z. Xiong, Y. Zeng, S. Liu, Z. Li, B. Liu, K. Wu, J. Xiao, X. Yinqi, Q. Zheng, Y. Zhang, H. Yang, J. Wang, L. Smeds, F. E. Rheindt, M. Braun, J. Fjeldså, L. Orlando, F. K. Barker, K. A. Jønsson, W. Johnson, K.-P. Koepfli, S. O'Brien, D. Haussler, O. A. Ryder, C. Rahbek, E. Willerslev, G. R. Graves, T. C. Glenn, J. McCormack, D. Burt, H. Ellegren, P. Alström, S. V. Edwards, A. Stamatakis, D. P. Mindell, J. Cracraft, E. L. Braun, T. Warnow, W. Jun, M. T. P. Gilbert & G. Zhang. 2014. Whole-genome analyses resolve early branches in the tree of life of modern birds. Science 346 (6215): 1320–1331.

[KF-V21] Kuhl, H., C. Frankl-Vilches, A. Bakker, G. Mayr, G. Nikolaus, S. T. Boerno, S. Klages, B. Timmermann & M. Gahr. 2021. An unbiased molecular approach using 3′-UTRs resolves the avian family-level tree of life. Molecular Biology and Evolution 38 (1): 108–127.

[L98] Livezey, B. C. 1998. A phylogenetic analysis of the Gruiformes (Aves) based on morphological characters, with an emphasis on the rails (Rallidae). Philosophical Transactions of the Royal Society of London Series B—Biological Sciences 353: 2077–2151.

[M05] Mayr, G. 2005. The Paleogene fossil record of birds in Europe. Biological Reviews 80: 515–542.

[M09] Mayr, G. 2009. Paleogene Fossil Birds. Springer.

[M11] Mayr, G. 2011. Cenozoic mystery birds—on the phylogenetic affinities of the bony-toothed birds (Pelagornithidae). Zoologica Scripta 40: 448–467.

[ME04] Mayr, G., & P. G. P. Ericson. 2004. Evidence for a sister group relationship between the Madagascan mesites (Mesitornithidae) and the cuckoos (Cuculidae). Senckenbergiana Biologica 84 (1–2): 1–17.

[M02] Mlíkovský, J. 2002. Cenozoic Birds of the World. Part 1: Europe. Ninox Press: Praha.

[O87] Okamura, C. 1987. New facts: Homo and all Vertebrata were born simultaneously in the former Paleozoic in Japan. Original Report of the Okamura Fossil Laboratory 15: 347–573.

[PB15] Prum, R. O., J. S. Berv, A. Dornburg, D. J. Field, J. P. Townsend, E. M. Lemmon & A. R. Lemmon. 2015. A comprehensive phylogeny of birds (Aves) using targeted next-generation DNA sequencing. Nature 526: 569–573.

[RN72] Rutgers, A., & K. A. Norris (eds) 1972. Encyclopaedia of Aviculture vol. 1. London, Blandford Press.

[U93] Unwin, D. M. 1993. Aves. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 717–737. Chapman & Hall: London.

Last updated: 29 January 2022.

Ornithurae

Reconstruction of Enaliornis barretti, copyright Nobu Tamura.

Belongs within: Ornithothoraces.
Contains: Aves.

The Ornithurae are a clade of birds including all modern species together with related fossil forms possessing a very short pygostyle and completely fused tarsometatarsus (Zhou & Zhang 2005). Basal members of this clade include the famous toothed seabirds Ichthyornis and Hesperornis from the Late Cretaceous of North America.

Synapomorphies (from Zhou & Zhang 2005): Pygostyle very short; coracoid strut-like with well-developed procoracoid process; furcula U-shaped; sternum elongate; major digit with first phalanx significantly laterally expanded, tightly abutted by reduced minor digit; tarsometatarsus completely fused.

<==Ornithurae [Ichthyornithiformes, Odontoholomorphae, Odontornithes, Odontotormae, Plataleae] C04
    |--Hongshanornis Zhou & Zhang 2005 ZZ05
    |    `--*H. longicresta Zhou & Zhang 2005 ZZ05
    `--+--+--Liaoningornis Hou 1997 ZZ05, P04
       |  |    `--L. longidigitus Hou 1997 P04
       |  `--+--Yixianornis grabaui ZZ05
       |     `--Yanornis Zhou & Zhang 2001 ZZ05, D07
       |          |--Y. chaoyangensis ZZ05
       |          `--Y. martini D07
       `--+--Apsaravis Norell & Clarke 2001 ZZ05, NC01 [Apsaravidae, Apsaraviformes]
          |    `--*A. ukhaana Norell & Clarke 2001 NC01
          `--+--+--Ambiortus Kurochkin 1982 P04
             |  |    `--A. dementjevi Kurochkin 1982 [incl. Holbotia ponomarenkoi Kurochkin 1982] P04
             |  `--Carinatae P04
             |       |--Ichthyornis Marsh 1872 C04 (see below for synonymy)
             |       |    `--I. anceps (Marsh 1872) (see below for synonymy) C04
             |       `--+--Aves C04
             |          |--Apatornis Marsh 1873 [Apatornithidae] C04
             |          |    `--*A. celer Marsh 1873 C04 [=Ichthyornis celer P04]
             |          |--Iaceornis Clarke 2004 C04
             |          |    `--*I. marshi Clarke 2004 C04
             |          `--Guildavis Clarke 2004 C04
             |               `--*G. tener (Marsh 1880) [=Ichthyornis tener] C04
             `--Hesperornithes [Hesperornithiformes, Odontoholcae, Odontolcae] C04
                  |  i. s.: Asiahesperornis Nessov & Prizemlin 1991 P04
                  |           `--A. bazhanovi Nessov & Prizemlin 1991 P04
                  |         Judinornis Nessov & Borkin 1983 P04
                  |           `--J. nogontsavensis Nessov & Borkin 1983 P04
                  |         Pasquiaornis Tokaryk, Cumbaa & Storer 1997 P04
                  |           |--P. hardiei Tokaryk, Cumbaa & Storer 1998 P04
                  |           `--P. tankei Tokaryk, Cumbaa & Storer 1997 P04
                  |         Potamornis Elżanowski, Paul & Stidham 1999 P04
                  |           `--P. skutchi Elżanowski, Paul & Stidham 1999 P04
                  |--Enaliornis Seeley 1876 P04 (see below for synonymy)
                  |    |--E. barretti Seeley 1876 [=Palaeocolymbus barretti] P04
                  |    |--E. sedgwicki (Seeley 1876) [=Pelagornis sedgwicki] P04
                  |    `--E. seeleyi D07
                  `--+--Baptornis Marsh 1877 P04 [Baptornithidae FP64]
                     |    `--B. advenus Marsh 1877 C04
                     `--Hesperornithidae C04
                          |--Coniornis Marsh 1893 FP64, P04
                          |    `--C. altus Marsh 1893 [incl. Hesperornis montanus Shufeldt 1915] P04
                          |--Lestornis Marsh 1876 C04, P04
                          |    `--L. crassipes Marsh 1876 C04 [=Hesperornis crassipes P04]
                          |--Parahesperornis Martin 1983 C04, P04
                          |    `--P. alexi Martin 1983 P04
                          `--Hesperornis Marsh 1872 [incl. Hargeria Lucas 1903] P04
                               |--H. gracilis Marsh 1876 C04 [=Hargeria gracilis P04]
                               `--H. regalis Marsh 1872 P04

Ornithurae incertae sedis:
  Chaoyangia Hou & Zhang 1993 P04
    `--C. beishanensis Hou & Zhang 1993 P04
  Gansus Hou & Liu 1984 P04
    `--G. yumenensis Hou & Liu 1984 P04
  Gargantuavis Buffetaut & Le Loeuff 1998 P04
    `--G. philoinos Buffetaut & Le Loeuff 1998 P04
  Horezmavis Nessov & Borkin 1983 P04
    `--H. eocretacea Nessov & Borkin 1983 P04
  Limenavis Clarke & Chiappe 2001 P04
    `--L. patagonica Clarke & Chiappe 2001 P04
  Qinornis paleocenica Xue 1995 M09
  Cimolopteryx retusus C04

Enaliornis Seeley 1876 P04 [incl. Palaeocolymbus Seeley 1876 P04, Pelagornis Seeley 1876 non Lartet 1857 P04; Enaliornithidae FP64]

Ichthyornis Marsh 1872 C04 [incl. Angelinornis Kashin 1972 P04, Colonosaurus Marsh 1872 P04; Ichthyornidae, Ichthyornithes, Ichthyornithidae]

Ichthyornis anceps (Marsh 1872) [=Graculavus anceps; incl. G. agilis Marsh 1873, Ichthyornis agilis, Plegadornis antecessor Wetmore 1962, Angelinornis antecessor, Ichthyornis antecessor, *I. dispar Marsh 1872, Colonosaurus mudgei Marsh 1872, I. validus Marsh 1880, I. victor Marsh 1876] C04

*Type species of generic name indicated

REFERENCES

[C04] Clarke, J. A. 2004. Morphology, phylogenetic taxonomy, and systematics of Ichthyornis and Apatornis (Avialae: Ornithurae). Bulletin of the American Museum of Natural History 286: 1–179.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[FP64] Fisher, J., & R. T. Peterson. 1964. The World of Birds: A comprehensive guide to general ornithology. Macdonald: London.

[M09] Mayr, G. 2009. Paleogene Fossil Birds. Springer.

[NC01] Norell, M. A., & J. A. Clarke. 2001. Fossil that fills a critical gap in avian evolution. Nature 409: 181–184.

[P04] Padian, K. 2004. Basal Avialae. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 210–231. University of California Press: Berkeley.

[ZZ05] Zhou, Z., & F. Zhang. 2005. Discovery of an ornithurine bird and its implications for Early Cretaceous avian radiation. Proceedings of the National Academy of Sciences of the USA 102: 18998–19002.

Last updated: 6 August 2017.

Troodontidae

Skeletal reconstruction of Saurornithoides junior, copyright Jaime A. Headden.

Belongs within: Maniraptora.

The Troodontidae are a group of small, cursorial theropods known from the Cretaceous of Asia and North America with a single possible representative, Koparion douglassi, also known from the Upper Jurassic (Makovicky & Norell 2004). Certain fragmentary taxa from the late Cretaceous of Europe, such as Bradycneme draculae and Elopteryx nopcsai, have also been suggested to be troodontids but are not universally accepted as such.

Characters (from Makovicky & Norell 2004, possibly excluding Jinfengopteryx): Dentary bearing groove on at least part of labial face containing exits for neurovascular foramina; quadrate with pneumatic foramen present caudally; basicranium with basisphenoid recess absent; internarial bar dorsoventrally flattened; large number of teeth present; mesial dentary teeth closely packed; laterosphenoid with depression present on ventral surface of postorbital process; reduced basal tubera usually lying directly ventral to occipital condyle; transverse processes elongate; distal caudals with sulcus on dorsal midline in place of neural spine; metatarsus asymmetrical with slender metatarsal II markedly shorter than metatarsals III and IV, metatarsal IV robust.

<==Troodontidae [Saurornithoididae]
    |--Jinfengopteryx GC13
    `--+--Mei Xu & Norell 2004 GC13, XN04
       |    `--*M. long Xu & Norell 2004 XN04
       `--+--Sinovenator Xu, Norell et al. 2002 MN04
          |    `--*S. changii Xu, Norell et al. 2002 XN02
          `--+--Sinusonasus Xu & Wang 2004 GC13, XW04
             |    `--*S. magnodens Xu & Wang 2004 XW04
             `--+--Byronosaurus Norell, Makovicky & Clark 2000 GC13, MN04
                |    `--B. jaffei Norell, Makovicky & Clark 2000 MN04
                `--+--Sinornithoides Russell & Dong 1993 GC13, MN04
                   |    `--S. youngi Russell & Dong 1993 MN04
                   `--+--Zanabazar GC13
                      |--Troodon Leidy 1856 (see below for synonymy) MN04
                      |    |--*T. formosus Leidy 1856 N85 (see below for synonymy)
                      |    |--‘Pectinodon’ asiamericanus Nessov 1995 (n. d.) MN04
                      |    `--T. isfarensis Nessov 1995 (n. d.) MN04
                      `--Saurornithoides Osborn 1924 MN04
                           |--S. asiamericanus D07
                           |--S. isfarensis D07
                           |--S. junior Barsbold 1974 MN04
                           `--S. mongoliensis Osborn 1924 MN04 [=Troodon mongoliensis P88]

Troodontidae incertae sedis:
  Borogovia Osmólska 1987 MN04
    `--B. gracilicrus Osmólska 1987 MN04
  Bradycneme draculae Harrison & Walker 1975 (n. d.) MN04
  Heptasteornis Harrison & Walker 1975 (n. d.) MN04
    `--H. andrewsi Harrison & Walker 1975 [=Troodon andrewsi] P88
  Koparion Chure 1994 H98
    `--K. douglassi Chure 1994 MN04
  Tochisaurus Kurzanov & Osmólska 1991 MN04
    `--T. nemegtensis Kurzanov & Osmólska 1991 MN04
  Elopteryx [Elopterygidae] MN04
    `--E. nopcsai Andrews 1913 (n. d.) MN04
  ‘Laelaps’ cristatus Cope 1876 (n. d.) MN04 [=Dromaeosaurus cristatus O69]
  ‘Laelaps’ explanatus Cope 1876 (n. d.) MN04 [=Dromaeosaurus explanatus O69]
  ‘Laelaps’ laevifrons Cope 1876 (n. d.) MN04 [=Dromaeosaurus laevifrons O69]
  Paronychodon lacustris Cope 1876 (n. d.) MN04
  Zapsalis abradens Cope 1876 (n. d.) MN04

Troodon Leidy 1856 [incl. Pectinodon Carpenter 1982, Polyodontosaurus Gilmore 1932, Stenonychosaurus Sternberg 1932] MN04

*Troodon formosus Leidy 1856 N85 [incl. Pectinodon bakkeri Carpenter 1982 MN04, Polyodontosaurus grandis Gilmore 1932 MN04, Stenonychosaurus inequalis Sternberg 1932 MN04]

*Type species of generic name indicated

REFERENCES

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[GC13] Godefroit, P., A. Cau, Hu D.-Y., F. Escuillié, Wu W. & G. Dyke. 2013. A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds. Nature 498: 359–362.

[H98] Holtz, T. R., Jr. 1998. A new phylogeny of the carnivorous dinosaurs. Gaia 15: 5–61.

[MN04] Makovicky, P. J., & M. A. Norell. 2004. Troodontidae. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 184–195. University of California Press: Berkeley.

[N85] Norman, D. 1985 (reprinted 2000). The Illustrated Encyclopedia of Dinosaurs. Salamander Books: London.

[O69] Ostrom, J. H. 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Bulletin of the Peabody Museum of Natural History 30: 1–165.

[P88] Paul, G. S. 1988. Predatory Dinosaurs of the World: A Complete Illustrated Guide. Simon & Schuster: New York.

[XN04] Xu, X., & M. A. Norell. 2004. A new troodontid dinosaur from China with avian-like sleeping posture. Nature 431: 838–841.

[XN02] Xu, X., M. A. Norell, X.-L. Wang, P. J. Makovicky & X.-C. Wu. 2002. A basal troodontid from the Early Cretaceous of China. Nature 415: 780–784.

[XW04] Xu X. & Wang X. 2004. A new troodontid (Theropoda: Troodontidae) from the Lower Cretaceous Yixian Formation of western Liaoning, China. Acta Geologica Sinica (English Edition) 78: 22–26.

Last updated: 3 July 2018.

Oviraptorosauria

Reconstruction of Caudipteryx zoui, by Matt Martyniuk.

Belongs within: Maniraptora.

The Oviraptorosauria are a group of cursorial theropods known from the Cretaceous of Eurasia and North America. Mostly less than two metres in length, they are characterised by deep, short skulls with the dentition reduced or absent. The ecology of oviraptorosaurs has been subject to much debate; they may have included herbivorous or omnivorous forms.

The majority of oviraptorosaurs belong to a clade Caenagnathoidea, which has been defined as the least inclusive clade including Oviraptor philoceratops and Caenagnathus collinsi. This clade is supported by several synapomorphies, including the presence of two longitudinal ridges and a toothlike ventral process on the palatal shelf of the maxilla, and a jaw joint close to the midline of the skull (Osmólska et al. 2004). Non-caenagnathoid oviraptorosaurs include the early Cretaceous Caudipteryx, the type species of which, C. zoui, is known from fossils in which a fan of feathers is preserved at the end of the tail.

Synapomorphies (from Osmólska et al. 2004): Premaxilla with crenulated ventral margin; parietal at least as long as frontal; ascending process of quadratojugal bordering more than three-quarters of infratemporal fenestra; U-shaped mandibular symphysis present; dentary edentulous; pubic foot with cranial process longer than caudal process.

<==Oviraptorosauria
    |  i. s.: Thecocoelurus Huene 1923 OCB04
    |           `--*T. daviesi (Seeley 1888) OCB04 [=Thecospondylus daviesi OCB04, Coelurus daviesi N02]
    |--+--Protarchaeopteryx Ji & Ji 1997 GC13, P04
    |  |    `--P. robusta Ji & Ji 1997 P04
    |  `--Incisivosaurus Xu, Cheng et al. 2002 XT07, OCB04
    |       `--*I. gauthieri Xu, Cheng et al. 2002 OCB04
    `--+--Similicaudipteryx GC13
       `--+--Avimimus Kurzanov 1981 [Avimimidae] OCB04
          |    `--A. portentosus Kurzanov 1981 OCB04
          `--+--Caudipteryx Ji, Currie et al. 1998 XT07, OCB04 [Caudipterygidae]
             |    |--C. dongi Zhou & Wang 2000 OCB04
             |    `--C. zoui Ji, Currie et al. 1998 OCB04
             |--Microvenator Ostrom 1970 GC13, OCB04
             |    `--M. celer Ostrom 1970 OCB04
             `--Caenagnathoidea [Caenagnathinae, Oviraptoroidea] OCB04
                  |  i. s.: Shixinggia oblita N10
                  |--Caenagnathidae [Elmisauridae] OCB04
                  |    |--Hagryphus giganteus N10
                  |    |--Elmisaurus Osmólska 1981 MOW02, OCB04
                  |    |    `--E. rarus Osmólska 1981 OCB04 [=Chirostenotes rarus P88]
                  |    |--Caenagnathasia Currie, Godfrey & Nessov 1993 OCB04
                  |    |    `--C. martinsoni Currie, Godfrey & Nessov 1993 1994 OCB04
                  |    |--Nomingia Barsbold, Osmólska et al. 2000 OCB04
                  |    |    `--N. gobiensis Barsbold, Osmólska et al. 2000 OCB04
                  |    `--Chirostenotes Gilmore 1924 OCB04 (see below for synonymy)
                  |         |--C. elegans (Parks 1933) OCB04 (see below for synonymy)
                  |         `--C. pergracilis Gilmore 1924 (see below for synonymy) OCB04
                  `--Oviraptoridae XT07
                       |  i. s.: Nemegtomaia barsboldi (Lü et al. 2004) [=Nemegtia barsboldi] N10
                       |--Gigantoraptor Xu, Tan et al. 2007 XT07
                       |    `--*G. erlianensis Xu, Tan et al. 2007 XT07
                       `--+--Oviraptorinae OCB04
                          |    |--Oviraptor Osborn 1924 OCB04
                          |    |    `--O. philoceratops Osborn 1924 OCB04
                          |    |--Rinchenia Barsbold 1997 OCB04
                          |    |    `--R. mongoliensis (Barsbold 1986) [=Oviraptor mongoliensis] OCB04
                          |    `--Citipati Clark, Norell & Barsbold 2001 OCB04
                          |         `--C. osmolskae Clark, Norell & Barsbold 2001 OCB04
                          `--Ingeniinae OCB04
                               |--+--Conchoraptor Barsbold 1986 OCB04
                               |  |    `--C. gracilis Barsbold 1986 OCB04
                               |  `--Khaan Clark, Norell & Barsbold 2001 OCB04
                               |       `--K. mckennai Clark, Norell & Barsbold 2001 OCB04
                               `--+--‘Ingenia’ Barsbold 1981 non Gerlach 1957 OCB04
                                  |    `--I. yanshini Barsbold 1981 OCB04 [=Oviraptor (Ingenia) yanshini P88]
                                  `--Heyuannia Lü 2003 ZZ08, OCB04
                                       `--H. huangi Lü 2003 OCB04

Chirostenotes Gilmore 1924 OCB04 [incl. Caenagnathus Sternberg 1940 OCB04, Macrophalangia Sternberg 1932 N85]

Chirostenotes elegans (Parks 1933) OCB04 [=Ornithomimus elegans OCB04; incl. Caenagnathus sternbergi Cracraft 1971 OCB04, Chirostenotes sternbergi D07]

Chirostenotes pergracilis Gilmore 1924 [incl. Caenagnathus collinsi Sternberg 1940, Macrophalangia canadensis Sternberg 1932] OCB04

*Type species of generic name indicated

REFERENCES

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[GC13] Godefroit, P., A. Cau, Hu D.-Y., F. Escuillié, Wu W. & G. Dyke. 2013. A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds. Nature 498: 359–362.

[MOW02] Maryańska, T., H. Osmólska & M. Wolsan. 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica 47 (1): 97–116.

[N02] Naish, D. 2002. The historical taxonomy of the Lower Cretaceous theropods (Dinosauria) Calamospondylus and Aristosuchus from the Isle of Wight. Proceedings of the Geologists' Association 113: 153–163.

[N10] Naish, D. 2010. Tetrapod Zoology: Book One. CFZ Press: Bideford (UK).

[N85] Norman, D. 1985 (reprinted 2000). The Illustrated Encyclopedia of Dinosaurs. Salamander Books: London.

[OCB04] Osmólska, H., P. J. Currie & R. Barsbold. 2004. Oviraptorosauria. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 165–183. University of California Press: Berkeley.

[P04] Padian, K. 2004. Basal Avialae. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 210–231. University of California Press: Berkeley.

[P88] Paul, G. S. 1988. Predatory Dinosaurs of the World: A Complete Illustrated Guide. Simon & Schuster: New York.

[XT07] Xu, X., Q. Tan, J. Wang, X. Zhao & L. Tan. 2007. A gigantic bird-like dinosaur from the Late Cretaceous of China. Nature 447: 844–847.

[ZZ08] Zhang, F., Z. Zhou, X. Xu, X. Wang & C. Sullivan. 2008. A bizarre Jurassic maniraptoran from China with elongate ribbon-like feathers. Nature 455: 1105–1108.

Last updated: 11 July 2018.

Casuariiformes

Emus Dromaius novaehollandiae, photographed by Bill & Mark Bell.

Belongs within: Palaeognathae.

The Casuariiformes are an Australasian clade of large flightless birds including the emus (Dromaius) and cassowaries (Casuarius). The emu Dromaius novaehollandiae is an inhabitant of open habitats in mainland Australia, while the less cursorial cassowaries are found in thick forest in New Guinea and northern Queensland. Cassowaries have mostly black plumage with bare, bright blue head and neck, often with one or two bright, contrastingly coloured wattles hanging from the neck, and a horny casque on the top of the head.

Characters (from here): Large graviportal flightless birds with 3 toes (hind toe lost). Sexes alike in coloration; plumage sombre brown or brownish black, naked blue skin on the head and neck. Plumage loose and hairlike due to the lack of barbules; no differentiated tail feathers. Wings and keel on sternum reduced, legs enlarged. Vestigial clavicles present in shoulder girdle; humerus much reduced, shorter than combined forewing and manus. Manus with a single digit (the third), bearing a long claw. Palate palaeognathous or dromaeognathous, with large vomers extending back to separate short palatines.

<==Casuariiformes [Casuariimorphae]
    |--Casuarius Brisson 1760 (see below for synonymy) B94
    |    |--C. australis Wall in Gould 1865 [incl. C. johnsoni Krefft 1867] N13
    |    |--C. bennettii S66
    |    |--C. bicarunculatus S66
    |    |--C. casuarius HK08
    |    |--C. galeatus N13
    |    |--C. lydekkeri F71 [=C. bennetti lydekkeri FP64]
    |    `--C. uniappendiculatus [incl. C. kaupi Rosenberg 1861] S66
    `--Dromaiidae [Dromaiinae, Dromiceiidae] B94
         |--Metapteryx bifrons WH02
         |--Emuarius U93
         |    |--E. gidju (Patterson & Rich 1987) [=Dromaius gidju] M09
         |    `--E. guljaruba Boles 2001 M09
         `--Dromaius Vieillot 1816 [incl. Dromiceius Vieillot 1816] B94
              |--D. ater FS01
              |--D. baudinianus FS01
              |--‘Dromaeus’ minor A14
              |--D. novaehollandiae (Latham 1790) WS48 (see below for synonymy)
              |    |--D. n. novaehollandiae WS48
              |    |--D. n. rothschildi (Mathews 1912) [=Dromiceius novaehollandiae rothschildi] WS48
              |    `--D. n. woodwardi (Mathews 1912) [=Dromiceius novaehollandiae woodwardi] WS48
              `--D. patricius F71

Casuarius Brisson 1760 [incl. Cela Möhring 1752 (pre-Linnean) non Cela Reichenbach 1852, Hippalectryo Gloger 1842; Casuariidae, Casuariinae, Celidae, Hippalectryonidae] B94

Dromaius novaehollandiae (Latham 1790) WS48 [=Casuarius novaehollandiae WS48, Dromiceius novaehollandiae FP64]

*Type species of generic name indicated

REFERENCES

[A14] Anderson, W. 1914. Notes on the occurrence of the sand-rock containing bones of extinct species of marsupials (emu, kangaroo, wombat, etc.) on King Island, Bass Strait, Tasmania. Records of the Australian Museum 10 (9): 275–283.

[B94] Bock, W. J. 1994. History and nomenclature of avian family-group names. Bulletin of the American Museum of Natural History 222: 1–281.

[FP64] Fisher, J., & R. T. Peterson. 1964. The World of Birds: A comprehensive guide to general ornithology. Macdonald: London.

[FS01] Flannery, T., & P. Schouten. 2001. A Gap in Nature: Discovering the World's Extinct Animals. Text Publishing: Melbourne.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[HK08] Hackett, S. J., R. T. Kimball, S. Reddy, R. C. K. Bowie, E. L. Braun, M. J. Braun, J. L. Chojnowski, W. A. Cox, K.-L. Han, J. Harshman, C. J. Huddleston, B. D. Marks, K. J. Miglia, W. S. Moore, F. H. Sheldon, D. W. Steadman, C. C. Witt & T. Yuri. 2008. A phylogenomic study of birds reveals their evolutionary history. Science 320: 1763–1768.

[M09] Mayr, G. 2009. Paleogene Fossil Birds. Springer.

[N13] North, A. J. 1913. On the early history of the Australian cassowary (Casuarius australis, Wall). Records of the Australian Museum 10 (4): 39–48, pls 8–9.

[S66] Schlegel, H. 1866. Extract from a letter addressed to Mr. P. L. Sclater respecting Casuarius uniappendiculatus. Proceedings of the Zoological Society of London 1866: 168.

[U93] Unwin, D. M. 1993. Aves. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 717–737. Chapman & Hall: London.

[WS48] Whittell, H. M., & D. L. Serventy. 1948. A systematic list of the birds of Western Australia. Public Library, Museum and Art Gallery of Western Australia, Special Publication 1: 1–126.

[WH02] Worthy, T. H., & R. N. Holdaway. 2002. The Lost World of the Moa: Prehistoric life of New Zealand. Indiana University Press: Bloomington (Indiana).

Last updated: 29 August 2019.

Aves

Greater rheas Rhea americana, copyright Nino Barbieri.

Belongs within: Ornithurae.
Contains: Struthio, Casuariiformes, Apteryx, Aepyornithidae, Dinornithiformes, Lithornithidae, Tinamidae, Neoaves, Anseriformes, Pangalliformes.

The name 'Aves' is used here to refer to the crown group of birds. In recent years, a consensus has developed between most phylogenetic studies (both morphological and molecular) that the Palaeognathae (ratites and tinamous) are the basalmost lineage within modern birds, with their sister group divided between the Galloanserae (including gamefowl and waterfowl) and the Neoaves (including all other birds). Living ratites are flightless birds of South America, Africa and Australasia, characterised by the absence of a keel on the sternum. They include the Rheidae, rheas, of South America, three-toed ostrich-like birds with greyish plumage. Recent molecular analyses (e.g. Burleigh et al. 2015) have suggested that the flighted Tinamidae (tinamous), which do have a keeled sternum, are nested within the living ratites, suggesting convergent evolution of the unkeeled sternum in various ratite lineages.

Several fossil birds remain doubtfully placed within this framework. Remiornis heberti is known from limb bones in the late Palaeocene of France, and may be a member of the palaeognaths (Mayr 2005). Eremopezus eocaenus is another very large bird from the upper Eocene of Egypt; though known only from leg bones, these indicate that it had very mobile toes, probably related to raptorial habits, rather than the cursorial or graviportal feet of modern ratites (Rasmussen et al. 2001).

See also: A frustrating giant bird; How irritable is this bird?; Bird evolution—problems with Science; More from the "They don't write papers like they used to" files.

<==Aves [Cursores, Gallinae, Neornithes]
    |--Palaeognathae (see below for synonymy) HK08
    |    |  i. s.: Remiornis Lemoine 1881 M02 [Remiornithidae M05, Remiornithiformes]
    |    |           |--*R. heberti Lemoine 1881 M02
    |    |           `--R. minor (Lemoine 1878) [=Gastornis minor] M02
    |    |         Palaeotis Lambrecht 1928 M02 [Palaeotididae M05]
    |    |           `--*P. weigelti Lambrecht 1928 (see below for synonymy) M02
    |    |         Eleutherornis Schaub 1940 M02 [Eleutherornithidae FP64, Eleutherornithini]
    |    |           `--*E. helveticus Schaub 1940 M02
    |    |         Eremopezus Andrews 1904 ML14, RS01 [Eremopezidae, Eremopezithes]
    |    |           `--*E. eocaenus Andrews 1904 [incl. Stromeria fajumensis Lambrecht 1929, S. fayumensis] RS01
    |    |--Struthio BKB15
    |    `--+--Rheidae [Rheiformes] BKB15
    |       |    |  i. s.: Opisthodactylus patagonicus Ameghino 1891 AH03
    |       |    |         Heterorhea dabbeni FP64
    |       |    |--Diogenornis fragilis ML14
    |       |    `--+--Pterocnemia pennata L81
    |       |       `--Rhea Brisson 1760 HK08, B94
    |       |            |--R. americana HK08
    |       |            `--R. pennata JT12
    |       `--+--+--Casuariiformes BKB15
    |          |  `--+--Apteryx BKB15
    |          |     `--Aepyornithidae ML14
    |          `--+--Dinornithiformes BKB15
    |             `--+--Lithornithidae ML14
    |                `--Tinamidae BKB15
    `--Neognathae [Amblypterae, Ciconii, Insessores, Pullastrae] HK08
         |  i. s.: Walbeckornis Mayr 2007 M07
         |           `--*W. creber Mayr 2007 M07
         |--Neoaves HK08
         `--Galloanseres [Galloanserae, Galloanserimorphae] JM14
              |--Anseriformes HK08
              `--Pangalliformes HK08

Aves incertae sedis:
  ‘Dimorpha’ Hodgson 1841 nec Panzer 1806 nec Gray 1840 C-S93
  Malacias S89
    |--M. castanopterus Salvadori 1889 S89
    `--M. melanoleucus S89
  Poliopsar S89
    |--P. fuscogularis Salvadori 1889 S89
    `--P. nemoricola [=Sturnia nemoricola] S89
  Trichostoma rubiginosa Walden 1875 S89
  Actinura ramsayi Walden 1875 S89
  Digenea cerviniventris Sharpe 1879 S89
  Lophospizias trivirgatus [=Astur trivirgatus] S89
  Scelospizias poliopsis S89
  Xantholaema haemacephala S89
  Chrysophlegma S89
    |--C. chlorolophum [=C. chlorolophus, Geclaus chlorolophus] S89
    `--C. flavinucha S89
  Lepocestes pyrrhotis [=Venilia pyrrhotis] S89
  Rhopodytes S89
    |--R. sumatranus [=Poliococcyx sumatranus] S89
    `--R. tristis S89
  Centrococcyx bengalensis S89
  Entomobia smyrnensis S89
  Dichoceros bicornis S89
  Hydrornis oatesi S89
  Cryptolopha S89
    |--C. bicolor Styan 1892 L03
    |--C. schwaneri S89
    |--C. superciliaris S89
    `--C. tephrocephala S89
  Muscicapula maculata S89
  Stoparola melanops S89
  Volvocivora melanoptera [=Campophaga melanoptera] S89
  Chaptia aenea S89
  Bhringa remifer S89
  Chibia splendens S89
  Dendrophila frontalis S89
  Machlolophus spilonotus S89
  Allotrius S89
    |--A. intermedius S89
    `--A. melanotis [=Ptererythrius melanotis] S89
  Mesia argentauris S89
  Siva S89
    |--S. cyanuroptera S89
    `--S. sordida S89
  Staphidia striata S89
  Timelia bengalensis S89
  Dryonastes chinensis S89
  Trochalopteron melanostigma S89
  Lioptila saturata S89
  Suya superciliaris S89
  Notodela leucura [=Myiomela leucura] S89
  Oreocincla dauma S89
  Oreicola ferrea S89
  Euplocomus lineatus S89
  Arboricola S89
    |--A. brunneipectus S89
    `--A. rufogularis S89
  Ardeiralla flavicollis S89
  Iyngipicus pumilus [=Picus pumilus] S89
  Nectarophila hasselti [=Cinnyris hasselti] S89
  Cyrtostomus flammaxillaris [=Cinnyris flammaxillaris] S89
  Cyanops ramsayi P89
  Erythrosterna acornaus Be66
  Petrocossyphus cyaneus Be66
  Chrysonotus intermedius Be66
  Cyanotis Scl66
    |--C. azarae Scl66
    `--C. omnicolor Scl66
  Noctua Sch66
    |--N. polleni Sch66
    `--N. sonneratii Sch66
  Gervaisia albispecularis Sch66
  Oxynotus Sch66
    |--O. ferrugineus Sch66
    `--O. newtoni Sch66
  Hartlaubia madagascariensis Sch66
  Taccocua leschenaultii Bu66
  ‘Heteromorpha’ Hodgson 1843 nec Hübner 1822 nec Kirby 1823 NO08
  Fluviatilavis Harrison 1983 M02
    `--*F. antunesi Harrison 1983 M02
  Actiornis Lydekker 1891 M02
    `--*A. anglicus Lydekker 1891 M02
  Ardeacites Haushalter 1855 M02
    `--*A. molassicus Haushalter 1855 M02
  Chenornis Portis 1884 M02
    `--*C. graculoides Portis 1884 M02
  Coturnipes Harrison & Walker 1977 M02
    `--*C. cooperi Harrison & Walker 1977 M02
  Miofulica Lambrecht 1933 M02
    `--*M. dejardini (Beneden 1871) [=Fulica dejardinii] M02
  Gigantibis Harrison & Walker 1976 M02
    `--*G. incognita Harrison & Walker 1976 M02
  Homalopus Milne-Edwards 1871 M02
    `--*H. picoides Milne-Edwards 1871 M02
  Palaeopapia Harrison & Walker 1979 [=Howardia Harrison & Walker 1976 non Berlese & Leonardi 1896] M02
    `--*P. eous (Harrison & Walker 1976) [=*Howardia eous] M02
  Ibidopodia Milne-Edwards 1871 M02
    `--*I. palustris Milne-Edwards 1871 M02
  Kievornis Aver’janov, Potapova & Nesov 1990 M02
    `--*K. rogovitshi Aver’janov, Potapova & Nesov 1990 M02
  Litoripes Harrison & Walker 1979 M02
    `--*L. medius Harrison & Walker 1979 M02
  Oligocathartes Harrison & Walker 1979 M02
    `--*O. olsoni Harrison & Walker 1979 M02
  Ornitholithes M02
    |--O. bosniaskii Portis 1887 M02
    |--O. faujasi Omboni 1885 M02
    |--O. gabbrensis Portis 1887 M02
    |--O. procaccinii Portis 1887 M02
    `--O. tenuipennis Omboni 1885 M02
  Ornitholithus Dughi & Sirugue 1962 M02
    |--*O. biroi Dughi & Sirugue 1962 M02
    `--O. arcuatus Dughi & Sirugue 1962 (see below for synonymy) M02
  Parvicuculus Harrison & Walker 1977 [Parvicuculidae] M02
    `--*P. minor Harrison & Walker 1977 M02
  Parvigyps Harrison & Walker 1977 M02
    `--*P. praevox Harrison & Walker 1977 M02
  Petropluvialis Harrison & Walker 1976 M02
    `--*P. simplex Harrison & Walker 1976 M02
  Piscator Harrison & Walker 1976 M02
    `--*P. tenuirostris Harrison & Walker 1976 M02
  Pliogrus Lambrecht 1933 M02
    `--*P. germanicus Lambrecht 1933 M02
  Proceriavis Harrison & Walker 1979 M02
    `--*P. maritini Harrison & Walker 1979 M02
  Procuculus Harrison & Walker 1977 M02
    `--*P. minutus Harrison & Walker 1977 M02
  Proplegadis Harrison & Walker 1971 M02
    `--*P. fisheri Harrison & Walker 1971 M02
  Pumiliornis Mayr 1999 M02
    `--*P. tessellatus Mayr 1999 M02
  Teracus Milne-Edwards 1871 [=Ieracus Aymard 1856 (n. n.)] M02
    `--*T. littoralis Milne-Edwards 1871 [=Ieracus littoralis Aymard 1856 (n. n.)] M02
  ‘Macropus’ Nuttall 1824 nec Shaw 1790 nec Latreille 1802 nec Thunberg 1805 S61
  Pengana robertbolesi N10
  Nyctalops Wagler 1832 H11
  Corvultur albicollis K08
  Cranellus Tobias 1844 KA-Z11
  Dysmorodrepanis munroi HSS13
  Graculavidae M07
    |--Graculavus Marsh 1872 M02
    |    `--*G. velox C04
    |--Telmatornis M07
    |--Laornis M07
    `--Palaeotringa M07
  Tshulia Nessov 1988 M07
  Palintropus Brodkorb 1970 M07
  Plectophora Gray 1834 LT61
  Chettusia gregaria C01

Nomen nudum: Ornitholithes ralli Anonymous in Karg 1805 M02

Inorganic: Rhea americana minilorientalis Okamura 1987 O87

Ornitholithus arcuatus Dughi & Sirugue 1962 [incl. O. mammeatus Dughi & Sirugue 1962, O. mammillatus Dughi & Sirugue 1962, O. mammosus Dughi & Sirugue 1962] M02

Palaeognathae [Apteryges, Apterygimorphae, Brevipennes, Palaeognathiformes, Panpalaeognathae, Panratitae, Ratitae, Ratiti, Struthiones, Struthioni, Struthioniformes, Struthionimorphae, Struthionoidea, Timamomorphae] HK08

*Palaeotis weigelti Lambrecht 1928 [incl. Paleogrus geiseltalensis Lambrecht 1935, Ornitocnemus geiseltalensis] M02

*Type species of generic name indicated

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