Gruiformes

The **Gruiformes** are an order containing a considerable number of living and extinct bird families, with a widespread geographical diversity. Gruiform means "crane-like".

Traditionally, a number of wading and terrestrial bird families that did not seem to belong to any other order were classified together as Gruiformes. These include 14 species of large cranes, about 145 species of smaller crakes and rails, as well as a variety of families comprising one to three species, such as the Heliornithidae, the limpkin, or the trumpeters. Other birds have been placed in this order more out of necessity to place them *somewhere*; this has caused the expanded Gruiformes to lack distinctive apomorphies. Recent studies indicate that these "odd Gruiformes" are if at all only loosely related to the cranes, rails, and relatives ("core Gruiformes").

## Contents

* 1Systematics
	+ 1.1Phylogeny
* 2See also
* 3References
* 4External links

## Systematics[edit]

There are only two suprafamilial clades (natural groups) among the birds traditionally classified as Gruiformes. Rails (Rallidae), flufftails (Sarothruridae), finfoots and sungrebe (Heliornithidae), adzebills (Aptornithidae), trumpeters (Psophiidae), limpkin (Aramidae), and cranes (Gruidae) compose the suborder Grues and are termed "core-Gruiformes" (Fain et al. 2007). These are the only true Gruiformes. The suborder Eurypygae includes the kagu (Rhynochetidae) and sunbittern (Eurypygidae). These are not even remotely related to Grues. The families of mesites or roatelos (Mesitornithidae), button-quails (Turnicidae), Australian plains-wanderer (Pedionomidae), seriemas (Cariamidae), and bustards (Otididae) each represent distinct and unrelated lineages. Many families known only from fossils have been assigned to the Gruiformes, e.g., Ergilornithidae, Phorusrhacidae, Messelornithidae, Eogruidae, Idiornithidae, Bathornithidae, to name just a few (see below). Though some of these are superficially 'crane-like' and the possibility that some may even be related to extant families traditionally included in the Gruiformes, there are no completely extinct families that can be confidently assigned to core-Gruiformes.

The traditional order Gruiformes was established by the influential German avian comparative anatomist Max Fürbringer (1888). Over the decades, many ornithologists suggested that members of the order were in fact more closely related to other groups (reviewed by Olson 1985, Sibley and Ahlquist 1990). For example, it was thought that sunbittern might be related to herons and that seriemas might be related to cuckoos. Olson and Steadman (1981) were first to correctly disband any of the traditional Gruiformes. They recognized that the Australian plains-wanderer (family Pedionomidae) was actually a member of the shorebirds (order Charadriiformes) based on skeletal characters. This was confirmed by Sibley and Ahlquist (1990) based on DNA–DNA hybridization and subsequently by Paton et al. (2003), Paton and Baker (2006) and Fain and Houde (2004, 2006). Sibley and Ahlquist furthermore removed button-quails (Turnicidae) from the Gruifomes based on large DNA–DNA hybridization distances to other supposed Gruiformes. However, it was not until the work of Paton et al. (2004) and Fain and Houde (2004, 2006) that the correct placement of buttonquails within the shorebirds (order Charadriiformes) was documented on the basis of phylogenetic analysis of multiple genetic loci. Using 12S ribosomal DNA sequences, Houde et al. (1997) were the first to present molecular genetic evidence of gruiform polyphyly, although apparently they were not convinced by it. However, on the basis of numerous additional sequence data, it has been shown decisively that the traditionally recognized Gruiformes consist of five to seven unrelated clades (Fain and Houde 2004, Ericson et al. 2006, Hackett et al. 2008).

Fain and Houde (2004) proposed that Neoaves are divisible into two clades, Metaves and Coronaves, although it has been suggested from the start that Metaves maybe paraphyletic (Fain and Houde 2004, Ericson *et al.* 2006, Hackett *et al.* 2008). Sunbittern, kagu, and mesites all group within Metaves but all the other lineages of "Gruiformes" group either with a collection of waterbirds or landbirds within Coronaves. This division has been upheld by the combined analysis of as many as 30 independent loci (Ericson *et al.* 2006, Hackett *et al.* 2008), but is dependent on the inclusion of one or two specific loci in the analyses. One locus, i.e., mitochondrial DNA, contradicts the strict monophyly of Coronaves (Morgan-Richards *et al.* 2008), but phylogeny reconstruction based on mitochondrial DNA is complicated by the fact that few families have been studied, the sequences are heavily saturated (with back mutations) at deep levels of divergence, and they are plagued by strong base composition bias.

The kagu and sunbittern are one another's closest relatives. It had been proposed (Cracraft 2001) that they and the recently extinct adzebills (family Aptornithidae) from New Zealand constitute a distinct Gondwanan lineage. However, sunbittern and kagu are believed to have diverged from one another long after the break-up of Gondwanaland and the adzebills are in fact members of the Grues (Houde *et al.* 1997, Houde 2009). The seriemas and bustards represent distinct lineages within neoavian waterbirds.
Rails are one of the most widespread Gruiformes

### Phylogeny[edit]

Rallidae based on the work by John Boyd.

*Ergilornis*, a member of the extinct clade Eogruidae.
**ORDER GRUIFORMES**

* Family †Songziidae Hou, 1990
	+ Genus †*Songzia* Hou, 1990
* **Suborder Grui**
	+ Superfamily Gruoidea Vigors, 1825
		- Family †Geranoididae Wetmore, 1933
		- Family †Parvigruidae Mayr, 2005
			* Genus †*Parvigrus* Mayr, 2005
			* Genus †*Rupelrallus* Fischer, 1997
		- Family Aramidae Bonaparte, 1854 (limpkin)
			* Genus †*Badistornis* Wetmore, 1940
			* Genus *Aramus* Vieillot, 1816 [*Courlili* Buffon, 1781; *Notherodius* Wagler, 1827] (limpkins)
		- Family Psophiidae Bonaparte, 1831 (trumpeters)
			* Genus *Psophia* Linnaeus, 1758
		- Family †Eogruidae Wetmore, 1934
			* Genus †*Sonogrus* Kuročkin, 1981
			* Genus †*Eogrus* Wetmore, 1932 [*Progrus* Bendukidze, 1971]
			* Subfamily †Ergilornithinae
				+ Genus †*Ergilornis* Kozlova, 1960
				+ Genus †*Amphipelargus* Lydekker, 1891
				+ Genus †*Urmiornis* Mecquenem, 1908
		- Family Gruidae (cranes)
			* Genus †*Camusia* Seguí, 2002
			* Subfamily Balearicinae Brasil, 1913
				+ Genus †*Aramornis* Wetmore, 1926
				+ Genus †*Geranopsis* Lydekker, 1871
				+ Genus †*Eobalearica* Gureev, 1949
				+ Genus *Balearica* Brisson, 1760 [*Geranarchus* Gloger, 1842] (crowned cranes)
			* Subfamily Gruinae Vigors, 1825
				+ Genus †*"Grus" conferta* Miller & Sibley, 1942 [Olson & Rasmussen, 2001]
				+ Genus †*"Probalearica" mongolica* Kurochkin, 1985
				+ Genus †*Palaeogrus* Portis, 1885 [*Palaeogrus* Salvadori, 1884 *nomen nudum*]
				+ Genus *Antigone* (Linnaeus, 1758)
				+ Genus *Leucogeranus* (Pallas, 1773)
				+ Genus *Grus* Brisson, 1760 non Moehring, 1758 [*Anthropoides* Vieillot, 1816; *Bugeranus* Gloger, 1841; *Megalornis* Gray, 1841; *Leucogeranus* Bonaparte, 1855; *Mathewsena* Iredale, 1914; *Mathewsia* Iredale, 1911; *Limnogeranus* Sharpe, 1893; *Laomedontia* Reichenbach, 1852; *Philorchemon* Gloger, 1842; *Scops* Gray, 1840 non Moehring, 1758 non Bruennich, 1772 npn Savigny, 1809] (cranes)
* **Suborder Ralli**
	+ Family †Aptornithidae (adzebills)
	+ Family Sarothruridae (flufftails)
		- Genus *Mentocrex* Peters, 1933 (wood rails)
		- Genus *Sarothrura* Heine, 1890 non Hasselt, 1823 [*Corethrura* Reichenbach, 1849 non Hope, 1843 non Gray, 1846; Daseioura *Penhallurick, 2003] (flufftails)*
	+ Family Heliornithidae Gray, 1841 (finfoots and sungrebe)
		- Genus *Heliopais* Sharpe, 1893 (Asian/masked finfoots)
		- Genus *Podica* Lesson, 1831 [*Rhigelura* Wagler, 1832; *Podoa* Bonaparte, 1857 non Illiger, 1811] (African finfoots)
		- Genus *Heliornis* Bonnaterre, 1791 [*Podoa* Illiger, 1811 non Bonaparte, 1857; *Plotoides* Brookes, 1830; *Podia* Swainson, 1837] (American finfoot)
	+ Family Rallidae (crakes, moorhens, gallinules, and rails)
		- Genus †*Aletornis* Marsh, 1872 [*Protogrus*]
		- Genus †*Australlus* Worthy & Boles, 2011
		- Genus †*Baselrallus* De Pietri & Mayr, 2014
		- Genus †*Belgirallus* Mayr & Smith, 2001
		- Genus †*Capellirallus* Falla, 1954 (snipe-billed rail)
		- Genus †*Creccoides* Shufeldt, 1892
		- Genus †*Eocrex* Wetmore, 1931
		- Genus †*Euryonotus* Mercerat, 1897
		- Genus †*Fulicaletornis* Lambrecht, 1933
		- Genus †*Hovacrex* Brodkorb, 1965 (Hova gallinule)
		- Genus †*Ibidopsis* Lydekker, 1891
		- Genus †*Latipons* Harrison & Walker, 1979
		- Genus †*Miofulica* Lambrecht, 1933
		- Genus †*Miorallus* Lambrecht, 1933
		- Genus †*Nesophalaris* Brodkorb & Dawson, 1962
		- Genus †*Nesotrochis* Wetmore, 1918 (West Indian cave-rails)
		- Genus †*Palaeoaramides* Lambrecht, 1933
		- Genus †*Palaeorallus* Wetmore, 1931
		- Genus †*Paraortygometra* Lambrecht, 1933
		- Genus †*Parvirallus* Harrison & Walker, 1979
		- Genus †*Pastushkinia* Zelenkov, 2013
		- Genus †*Quercyrallus* Lambrecht, 1933
		- Genus †*Rallicrex* Lambrecht, 1933
		- Genus †*Rhenanorallus* Mayr, 2010
		- Genus †*Vitirallus* Worthy, 2004 (Viti Levu rails)
		- Genus †*Wanshuina* Hou, 1994
		- Genus †*Youngornis* Yeh, 1981
		- Genus †Rallidae gen. et sp. indet. [*Fulica podagrica* (partim)] (Barbados rail)
		- Genus †Rallidae gen. et sp. indet. (Easter Island rail)
		- Genus †Rallidae gen. et sp. indet. (Fernando de Noronha rail)
		- Genus †Rallidae gen. et sp. indet. (Tahitian "goose”)
		- Genus †Rallidae gen. et sp. indet. (Bokaak "bustard")
		- Genus †Rallidae gen. et sp. indet. ('Amsterdam Island' rail)
		- Genus *Rougetius* Bonaparte, 1856 (Rouget's Rails)
		- Subfamily Rallinae Rafinesque, 1815
			* Genus †*Pleistorallus* Worthy, 1997 (Fleming's rails)
			* Genus *Anurolimnas* Sharpe, 1893 (Chestnut-headed Crakes)
			* Genus *Biensis* (Madagascan Rails)
			* Genus *Rallicula* Schlegel, 1871 [*Corethruropsis* Salvadori, 1876] (forest-rails)
			* Genus *Rallus* Linnaeus, 1758 [†*Epirallus* Miller, 1942]
			* Genus †*Aphanapteryx* von Frauenfeld, 1868 [*Pezocrex* Hachisuka, 1953] (Mauritius/Red rails)
			* Genus †*Erythromachus* Milne-Edwards, 1873 (Rodriquez rails)
			* Genus *Dryolimnas* Sharpe, 1893
			* Genus *Crex* Bechstein, 1803 [*Crecopsis* Sharpe, 1893] (greater crakes)
			* Genus *Lewinia* Gray, 1855 [*Aramidopsis* Sharpe, 1893; *Donacias* Heine & Reichenow, 1890; *Hyporallus* Iredale & Mathews, 1926]
			* Genus *Canirallus* Bonaparte, 1856 (grey-throated rail)
			* Genus *Gymnocrex* Salvadori, 1875 (bare-faced rails)
			* Genus *Gallirallus* Lafresnaye, 1841 [*Tricholimnas* Sharpe, 1893; *Nesoclopeus* Peters, 1932; *Cabalus* Hutton, 1874; *Habropteryx* Stresemann, 1932; *Eulabeornis* Gould, 1844; †*Diaphorapteryx* Forbes, 1893; *Hypotaenidia* Reichenbach, 1853; *Sylvestrornis* Mathews, 1928]
		- Subfamily Gallinulinae Gray, 1840
			* Tribe Pardirallini Livezey, 1998 [Aramidinae] (Wood-rails & allies)
				+ Genus *Pardirallus* Bonaparte, 1856 [*Ortygonax* Heine, 1890]
				+ Genus *Mustelirallus* Bonaparte, 1858 [*Neocrex* Sclater & Salvin, 1869; *Cyanolimnas* Barbour & Peters, 1927]
				+ Genus *Amaurolimnas* Sharpe 1893 (Rufous rails; Uniform crakes)
				+ Genus *Aramides* Pucheran, 1845
			* Tribe Gallinulini Gray, 1840 [Fulicarinae (Nitzsch, 1820) sensu Livezey, 1998]
				+ Genus *Tribonyx* Du Bus de Gisignies, 1840 [*Brachyptrallus* Lafresnaye, 1840; *Microtribonyx* Sharpe, 1893] (native-hens)
				+ Genus *Porzana* Vieillot, 1816 [*Limnobaenus* Sundevall, 1872; *Phalaridion* Kaup, 1829; *Porzanoidea* Mathews, 1912; *Porzanoides* Condon, 1975; *Rallites* Pucheran, 1845; *Schoenocrex* Roberts, 1922; *Porphyriops* Pucheran, 1845]
				+ Genus *Paragallinula* Sangster, García-R & Trewick, 2015 (Lesser Moorhen)
				+ Genus *Gallinula* Brisson, 1760 [*Hydrogallina* Lacépède, 1799; *Stagnicola* Brehm, 1831; *Porphyriornis* Allen, 1892 *Pareudiastes* Hartlaub & Finsch, 1871 *Edithornis*]
				+ Genus *Fulica* Linnaeus, 1758 [†*Palaeolimnas* Forbes, 1893]
		- Subfamily Porphyrioninae Reichenbach, 1849
			* Tribe Porphyrionini Reichenbach, 1849 (Purple gallinules & swamphens)
				+ Genus †*Aphanocrex* Wetmore, 1963 (St. Helena swamphens)
				+ Genus *Porphyrio* Brisson, 1760 [*Notornis* Owen, 1848]
			* Tribe Himantornithini Bonaparte, 1856 (Bush-hens & Waterhens)
				+ Genus *Himantornis* Hartlaub, 1855 (Nkulenga rails)
				+ Genus *Megacrex* D'Albertis & Salvadori, 1879 (New Guinea Flightless Rails)
				+ Genus *Aenigmatolimnas* (Striped Crakes)
				+ Genus *Gallicrex* Blyth, 1852 [*Gallinulopha* Bonaparte, 1854; *Hypnodes* Reichenbach, 1853] (Watercocks)
				+ Genus *Amaurornis* Reichenbach, 1853 [*Erythra* Reichenbach, 1853; *Pisynolimnas* Heine & Reichenow, 1890; *Poliolimnas* Sharpe, 1893] (Bush-hen)
			* Tribe Zaporniini Des Murs, 1860 (Old world crakes)
				+ Genus *Rallina* Gray, 1846 [*Euryzona* Gray, 1855; *Tomirdus* Mathews, 1912] (chestnut-rails)
				+ Genus *Zapornia* Stephens, 1824 [*Limnocorax* Peters, 1854; *Limnobaenus*; *Corethrura* Grey, 1846]
			* Tribe Laterallini Tif, 2014 (New world crakes)
				+ Genus *Micropygia* Bonaparte, 1856 (Ocellated Crakes)
				+ Genus *Rufirallus* (russet-crowned crake)
				+ Genus *Laterallus* Gray, 1855 (ruddy crakes)
				+ Genus *Coturnicops* Gray, 1855 (barred-backed crakes)
				+ Genus *Hapalocrex* (Yellow-breasted Crakes)
				+ Genus *Limnocrex*
				+ Genus *Mundia* Bourne, Ashmole & Simmons, 2003 (Ascension Island Crakes)
				+ Genus *Creciscus* Cabanis, 1857 [*Atlantisia* Lowe, 1923] (blackish crakes)

When considered to be monophyletic, it was assumed that Gruiformes was among the more ancient of avian lineages. The divergence of "gruiforms" among "Metaves" and "Coronaves" is proposed to be the first divergence among Neoaves, far predating the Cretaceous–Paleogene extinction event c. 66 mya (Houde 2009). No unequivocal basal gruiforms are known from the fossil record. However, there are several genera that are not unequivocally assignable to the known families and that may occupy a more basal position:

* *Propelargus* (Late Eocene/Early Oligocene of Quercy, France) – cariamid or idornithid
* *Rupelrallus* (Early Oligocene of Germany) – rallid? parvigruid?
* *Badistornis* (Brule Middle Oligocene of Shannon County, Missouri) – aramid?
* *Probalearica* (Late Oligocene? – Middle Pliocene of Florida, France?, Moldavia and Mongolia) – gruid? A *nomen dubium*?
* "Gruiformes" gen. et sp. indet. MNZ S42623 (Bathans Early/Middle Miocene of Otago, New Zealand) – Aptornithidae?
* *Aramornis* (Sheep Creek Middle Miocene of Snake Creek Quarries, U.S.) – gruid? aramid?
* *Euryonotus* (Pleistocene of Argentina) – rallid?

Other even more enigmatic fossil birds and five living families are occasionally suggested to belong into this order, such as the proposed Late Cretaceous family Laornithidae and the following taxa:

* Family †Gastornithidae (diatrymas) (fossil)
* Family †Messelornithidae (Messel-birds)
* Family †Salmilidae (fossil) – distinct order (Cariamiformes)
* Family †Geranoididae (fossil) – distinct order (Cariamiformes)[*citation needed*]; however, Mayr (2016) argued they might be members of Gruiformes, specifically stem group representatives of the Gruoidea.
* Family †Bathornithidae (fossil) – distinct order (Cariamiformes)
* Family †Idiornithidae (fossil) – distinct order (Cariamiformes)
* Family †Phorusrhacidae (terror birds) (fossil) – distinct order (Cariamiformes)
* Family Cariamidae (seriemas) – Neoavian landbirds – distinct order (Cariamiformes)
* Family Otididae (bustards) – Neoavian waterbirds – distinct order
* Family Eurypygidae (sunbittern) – prospective "Metaves" – new order Eurypygiformes together with kagu
* Family Rhynochetidae (kagu) – prospective "Metaves" – new order Eurypygiformes together with sunbittern
* Family Mesitornithidae (mesites, roatelos, monias) prospective "Metaves" – distinct order
* *Horezmavis* (Bissekty Late Cretaceous of Kyzyl Kum, Uzbekistan)
* *Telmatornis* (Navesink Late Cretaceous?)
* *Amitabha* (Bridger middle Eocene of Forbidden City, Wyoming) – rallid?
* *Eobalearica* (Ferghana Late? Eocene of Ferghana, Uzbekistan) – gruid?
* *"Phasianus" alfhildae* (Washakie B Late Eocene of Haystack Butte, U.S.)
* *Talantatos* (Late Eocene of Paris Bain, France)
* *Telecrex* (Irdin Manha Late Eocene of Chimney Butte, China) – rallid?
* Neornithes incerta sedis (Late Paleocene/Early Eocene of Ouled Abdoun Basin, Morocco)
* *Aminornis* (Deseado Early Oligocene of Rio Deseado, Argentina) – aramid?
* *Loncornis* (Deseado Early Oligocene of Rio Deseado, Argentina) – aramid?
* *Riacama* (Deseado Early Oligocene of Argentina)
* *Smiliornis* (Deseado Early Oligocene of Argentina)
* *Pseudolarus* (Deseado Early Oligocene – Miocene of Argentina) – gruiform?
* *Gnotornis* (Brule Late Oligocene of Shannon County, Missouri) – aramid?
* *Anisolornis* (Santa Cruz Middle Miocene of Karaihen, Argentina) – aramid?
* *Occitaniavis* – cariamid or idiornithid, includes *Geranopsis elatus*