The Early Eocene Gallinuloides wyomingensis, from Mayr & Weidig (2004).

Belongs within: Neognathae.
Contains: Anseres, Galliformes.

The Galloanserae are a clade of birds uniting the waterfowl (Anseriformes) and gamefowl (Galliformes). The clade is supported by molecular data as well as by morphological characters including presence of a sessile basipterygoid processes, long, blade-like retroarticular processes on the mandible, and a derived morphology of the quadrate (Mayr 2009).

As well as the ducks and geese, the Anseriformes include the screamers (Anhimidae) of South America. Screamers have a short curved bill, soft downy feathers on the relatively small head and short neck, and two large, sharp spurs on the forward edge of each wing. Their toes mostly lack webbing (Austin 1961). Anseriformes have one of the better fossil records among birds, including the giant flightless Gastornithidae of the Palaeogene Holarctic.

The Pangalliformes is the total group for gamebirds, i.e. crown-group Galliformes and all fossil species more closely related to them than to other birds. Pangalliformes probably diverged from other birds during the Cretaceous, and Austinornis has been indicated as a possible Late Cretaceous pangalliform (Clarke 2004).

Stem-group relatives of the Galliformes include the Gallinuloididae, known from well-preserved skeletons from the early Eocene of Europe and North America. The Paraortygidae and Quercymegapodiidae are known from more fragmentary remains from the late Eocene and Oligocene of Europe (Mayr 2009).

<==Galloanseres [Galloanserae, Galloanserimorphae]WD17
    |--+--Vegavis Clarke, Tambussi et al. 2005WD17, CT05 [Vegaviidae, Vegaviiformes]
    |  |    `--*V. iaai Clarke, Tambussi et al. 2005CT05
    |  `--Anseriformes [Anatiformes, Anserimorphae]WD17
    |       |  i. s.: Aminornis exavatus Ameghino 1899M09
    |       |--AnseresCT05
    |       `--Anhimidae [Anhimae, Palamedaeidae, Palamedeidae]CT05
    |            |--Chaunoides antiquus Alvarenga 1999M09
    |            |--Loxornis Ameghino 1894M09, A94
    |            |    `--*L. clivus Ameghino 1895A94, M09
    |            |--Anhima Brisson 1760 [incl. Palamedaea Möhring 1752 (pre-Linnean), Palamedea Linné 1766]B94
    |            |    |--A. cornutaC04 [=Palamedea cornutaSS66]
    |            |    `--‘Palamedea’ cristata [incl. Psophia cariama]G97
    |            `--ChaunaCT05
    |                 |--C. chavariaS66
    |                 |--C. derbiana Gray 1845M66 [incl. C. nigricollis Sclater 1864S66, M66]
    |                 `--C. torquata (Oken 1816)ME04
    `--Pangalliformes [Cracoidea, Galli]C04
         |  i. s.: Austinornis Clarke 2004C04
         |           `--*A. lentus (Marsh 1877) [=Graculavus lentus, Ichthyornis lentus]C04
         |         Amitabha Gulas-Wroblewski & Wroblewski 2003MW04
         |           `--A. urbsinterdictensis Gulas-Wroblewski & Wroblewski 2003MW04
         |         Palaeonossax senectus Wetmore 1956M09
         |    |--Gastornithidae [Diatrymae, Diatrymidae, Diatrymiformes, Gastornithes, Gastornithiformes]RS01
         |    |    |--Zhongyuanus xichuanensis Hou 1980M09
         |    |    `--Gastornis Hébert 1855M05 (see below for synonymy)
         |    |         |--*G. parisiensis Hébert 1855M02 [incl. G. edwardsi Lemoine 1878M09, G. klaasseni Newton 1886M09]
         |    |         |--G. ajax (Shufeldt 1913) [=Diatryma ajax]M02
         |    |         |--G. geiselensis (Fischer 1978) [=Diatryma geiselensis]M09
         |    |         |--G. giganteus (Cope 1876)WD17 [=*Diatryma giganteusM02]
         |    |         |--G. minor Lemoine 1878M09 [=Remiornis minorM02; incl. G. russeli Martin 1992M09]
         |    |         |--G. regens (Marsh 1894)M09 [=*Barornis regensM02, Diatryma regensM09]
         |    |         |--G. sarasini (Schaub 1929) [=Diatryma sarasini]M02
         |    |         |--‘Diatryma’ steini (Matthew & Granger 1917)M07
         |    |         `--*Omorhamphus’ storchii Sinclair 1928M02
         |    `--Dromornithidae [Dromornithes, Dromornithiformes]M11
         |         |  i. s.: Bullockornis planei Rich 1979U93
         |         |--+--Barawertornis tedfordi Rich 1979WD17, M09
         |         |  `--‘Ilbandornis’ lawsoni Rich 1979WD17
         |         `--+--Ilbandornis woodburnei Rich 1979WD17
         |            `--+--‘Dromornis’ murrayi Worthy et al. 2016WD17
         |               `--+--DromornisWD17
         |                  |    |--D. australisFP64
         |                  |    |--D. planei Rich 1979WD17
         |                  |    `--D. stirtoni Rick 1979WD17
         |                  `--Genyornis Stirling & Zietz 1896WD17, D07
         |                       |--G. australisD07
         |                       |--G. newtoni Stirling & Zeitz 1896WD17
         |                       `--G. stirtoniD07
         `--+--Gallinuloididae [Gallinuloidinae]M09
            |    |--Gallinuloides Eastman 1900MW04
            |    |    `--G. wyomingensis Eastman 1900MW04
            |    `--Paraortygoides Mayr 2000MW04
            |         `--*P. messelensis Mayr 2000M02
               |    |--Archaealectrornis sibleyi Crowe & Short 1992M09
               |    |--Pirortyx Brodkorb 1964M09, M02
               |    |    `--*P. major (Gaillard 1939) [=Palaeortyx major]M02
               |    |--Procrax Tordoff & Macdonald 1957M09, MW04
               |    |    `--P. brevipes Tordoff & Macdonald 1957M09
               |    |--Paraortyx Gaillard 1908M02
               |    |    |--*P. lorteti Gaillard 1908 (see below for synonymy)M02
               |    |    `--P. brancoi Gaillard 1908M02
               |    `--Taoperdix Milne-Edwards 1869M09, M02
               |         |--*T. pessieti (Gervais 1862) [=Tetrao pessieti]M02
               |         `--T. miocaena Ballmann 1969M02
                  `--Quercymegapodiidae [Quercymegapodiinae]MW04
                       |--Taubacrex Alvarenga 1988M02
                       |    `--T. granivora Alvarenga 1988M09
                       |--Ludiortyx Brodkorb 1964M05, M02 [incl. Eortyx Brodkorb 1967M02]
                       |    `--L. hoffmanni (Gervais 1852) (see below for synonymy)M02
                       |--Quercymegapodius Mourer-Chauviré 1992M02
                       |    |--*Q. depereti (Gaillard 1908) [=Palaeocryptonyx depereti, Pirortyx depereti]M02
                       |    `--Q. brodkorbi Mourer-Chauviré 1992M09
                       `--Ameripodius Alvarenga 1995M02
                            |--*A. silvasantosi Alvarenga 1995M02
                            `--A. alexis Mourer-Chauviré 2000M02

Gastornis Hébert 1855M05 [incl. Barornis Marsh 1894M02, Diatryma Cope 1876M09, M05, Omorhamphus Sinclair 1928M02]

Ludiortyx hoffmanni (Gervais 1852) [=Tringa hoffmanni, *Eortyx hoffmanni, Palaeortyx hoffmanni; incl. P. blanchardi Milne-Edwards 1869, *Ludiortyx blanchardi]M02

*Paraortyx lorteti Gaillard 1908 [incl. Palaeortyx cayluxensis Milne-Edwards 1892 non Lydekker 1891, Par. cayluxensis, Pal. gaillardi Lambrecht 1933, Ludiortyx gaillardi, Paraortyx gaillardi]M02

*Type species of generic name indicated


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

[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.

[CT05] Clarke, J. A., C. P. Tambussi, J. I. Noriega, G. M. Erickson & R. A. Ketcham. 2005. Definitive fossil evidence for the extant avian radiation in the Cretaceous. Nature 433: 305–308.

[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.

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[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.

[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.

[MW04] Mayr, G., & I. Weidig. 2004. The Early Eocene bird Gallinuloides wyomingensis—a stem group representative of Galliformes. Acta Palaeontologica Polonica 49 (2): 211–217.

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

[M66] Moore, T. J. 1866. On the habitat of the Derbyan crested screamer (Chauna derbiana, G. R. Gray). Proceedings of the Zoological Society of London 1866: 368–369.

[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.

[S66] Sclater, P. L. 1866. Note on Chauna nigricollis. Proceedings of the Zoological Society of London 1866: 369–370.

[SS66] Sclater, P. L., & O. Salvin. 1866. Catalogue of birds collected by Mr. E. Bartlett on the River Uyacali, Eastern Peru, with notes and descriptions of new species. Proceedings of the Zoological Society of London 1866: 175–201.

[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.

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