Anseriformes
Anseriformes is an order of birds also known as waterfowl that comprises about 180 living species of birds in three families: Anhimidae, Anseranatidae, and Anatidae, the largest family, which includes over 170 species of waterfowl, among them the ducks, geese, and swans. Most modern species in the order are highly adapted for an aquatic existence at the water surface. With the exception of screamers, males have penises, a trait that has been lost in the Neoaves, the clade consisting of all other modern birds except the galliformes and paleognaths. Due to their aquatic nature, most species are web-footed.
Evolution
Anseriformes are one of only two types of modern bird to be confirmed present during the Mesozoic alongside the other dinosaurs, and in fact were among the very few birds to survive their extinction, along with their cousins, the Galliformes. These two groups only occupied two ecological niches during the Mesozoic, living in water and on the ground, while the toothed Enantiornithes were the dominant birds that ruled the trees and air. The asteroid that ended the Mesozoic destroyed all trees as well as animals in the open, a condition that took centuries to recover from. The Anseriformes and Galliformes are thought to have survived in the cover of burrows and water, and not to have needed trees for food and reproduction.The earliest known stem anseriform is the presbyornithid Teviornis from the Nemegt Formation of Mongolia. Some members apparently surviving the KT extinction event, including presbyornithids, thought to be the common ancestors of ducks, geese, swans, and screamers, the last group once thought to be Galliformes, but now genetically confirmed to be closely related to geese. The first known duck fossils start to appear about 34 million years ago.
Waterfowl are the best-known examples of sexually antagonistic genital coevolution in vertebrates, causing genital adaptations to coevolve in each sex to advance control over mating and fertilization. Sexually antagonistic coevolution occurs as a consequence of sexual conflict between males and females, resulting in coevolutionary process that reduce fit, or that functions to decrease ease of having sex.
Taxonomy
The Anseriformes and the Galliformes belong to a common group, the Galloanserae. They are the most primitive neognathous birds, and as such they should follow the Palaeognathae in bird classification systems. Several unusual extinct families of birds like the albatross-like pseudotooth birds and the giant flightless gastornithids and mihirungs have been found to be stem-anseriforms based on common features found in the skull region, beak physiology and pelvic region. The genus Vegavis for a while was found to be the earliest member of the anseriform crown group but a recent 2017 paper has found it to be just outside the crown group in the family Vegaviidae. However, the monophyly of Vegaviidae was questioned by Torres et al. who described a nearly complete skull of Vegavis in 2025, supporting its placement within crown group Anseriformes. However, Irazoqui et al. who redescribed this skull suggested that Vegavis can only be confidently placed as a neognath of uncertain affinities.Below is the general consensus of the phylogeny of anseriforms and their stem relatives.
Systematics
systematics, especially regarding placement of some "odd" genera in the dabbling ducks or shelducks, is not fully resolved. See the Anatidae article for more information, and for alternate taxonomic approaches. Anatidae is traditionally divided into subfamilies Anatinae and Anserinae. The Anatinae consists of tribes Anatini, Aythyini, Mergini and Tadornini. The higher-order classification below follows a phylogenetic analysis performed by Mikko's Phylogeny Archive and John Boyd's website.- Order Anseriformes
- * ?†Conflicto Claudia P. Tambussi et al. 2019 - tentatively placed here; possibly family Conflictonidae
- * †Anatalavis Olson & Parris 1987 - including Nettapterornis; may belong in Anseranatidae or Conflictonidae
- * †Naranbulagornis Zelenkov 2019
- * †Anachronornis
- * †Paakniwatavis Musser & Clarke 2024
- * Suborder Anhimae Wetmore & Miller 1926
- ** Genus †Chaunoides Alvarenga 1999
- ** Family Anhimidae Stejneger 1885
- *** Genus Anhima Brisson 1760
- *** Genus Chauna Illiger 1811
- * Suborder Anseres
- ** Superfamily Anseranatoidea
- *** Family Anseranatidae Sclater 1880
- **** Genus †Anserpica Mourer-Chauviré, Berthet & Hugueney 2004
- **** Genus †Eoanseranas Worthy & Scanlon 2009
- **** Genus Anseranas Lesson 1828
- ** Superfamily Anatoidea
- *** Family †Presbyornithidae? Wetmore 1926
- **** Genus †Teviornis Kuročkin, Dyke & Karhu 2002
- **** Genus †Telmabates Howard 1955
- **** Genus †Presbyornis Wetmore 1926
- **** Genus †Wilaru Boles et al. 2013
- **** Genus †Bumbalavis Zelenkov 2021
- **** Genus †Murgonornis Worthy et al. 2023
- *** Family †Paranyrocidae Miller & Compton 1939
- **** Genus †Paranyroca Miller & Compton 1939
- *** Family Anatidae Leach 1820
- **** Genus †Garganornis ballmanni Meijer 2014
- **** Subfamily †Romainvilliinae Lambrecht 1933
- ***** Genus †Romainvillia Lebedinský 1927
- ***** Genus †Saintandrea Mayr & De Pietri 2013
- **** Subfamily Dendrocygninae Reichenbach 1849–50
- ***** Genus Dendrocygna Swainson 1837
- ***** Genus Thalassornis Eyton 1838
- **** Subfamily †Dendrocheninae Livezey & Martin 1988
- ***** Genus †Dendrochen Miller 1944
- ***** Genus †Manuherikia Worthy et al. 2007
- ***** Genus †Mionetta Livezey & Martin 1988
- **** Subfamily Stictonettinae
- ***** Genus Stictonetta Reichenbach 1853
- **** Subfamily Anserinae Vigors 1825 sensu Livezey 1996
- ***** Genus †Anserobranta Kuročkin & Ganya 1972
- ***** Genus †Asiavis Nesov 1986
- ***** Genus †"Chenopis" De Vis 1905
- ***** Genus †Cygnavus Lambrecht 1931
- ***** Genus †Cygnopterus Lambrecht 1931
- ***** Genus †Eremochen Brodkorb 1961
- ***** Genus †Megalodytes Howard 1992
- ***** Genus †Annakacygna Matsuoka & Hasegawa 2022
- ***** Genus †Paracygnus Short 1969
- ***** Genus †Presbychen Wetmore 1930
- ***** Genus †Cnemiornis Owen 1866
- ***** Genus †Afrocygnus Louchart et al. 2005
- ***** Genus Coscoroba Reichenbach 1853
- ***** Genus Cereopsis Latham 1801
- ***** Genus Cygnus Garsault 1764
- ***** Genus Branta Scopoli 1769
- ***** Tribe Anserini Vigors 1825
- ****** Genus Anser Brisson 1760
- **** Subfamily Tadorninae Reichenbach 1849–50
- ***** Genus †Australotadorna Worthy 2009
- ***** Genus †Brantadorna Howard 1964
- ***** Genus †Centrornis Andrews 1897
- ***** Genus †Miotadorna Worthy et al. 2007
- ***** Genus †Nannonetta Campbell 1979
- ***** Genus †Pleistoanser Agnolín 2006
- ***** Genus Plectropterus
- ***** Genus Merganetta Gould 1842
- ***** Genus Chloephaga Eyton 1838
- ***** Genus Neochen Oberholser 1918
- ***** Genus Cyanochen Bonaparte 1856
- ***** Genus Tadorna Boie 1822
- ***** Genus †Balcanas Boev 1998
- ***** Genus Radjah Reichenbach, 1853
- ***** Genus Alopochen Stejneger 1885
- ***** Genus Cairina Fleming 1822
- ***** Genus Hymenolaimus Gray 1843
- ***** Genus Sarkidiornis Eyton 1838
- ***** Genus Tachyeres Owen 1875
- ***** Genus †Zqueheanas Agnolin et al 2025
- **** Subfamily Anatinae Vigors 1825 sensu Livezey 1996
- ***** Genus †Dunstanetta Worthy et al. 2007
- ***** Genus †Lavadytis Stidham & Hilton 2015
- ***** Genus †Pinpanetta Worthy 2009
- ***** Genus †Tirarinetta Worthy 2008
- ***** Genus Aix Boie 1828
- ***** Genus Callonetta Delacour 1936
- ***** Genus Chenonetta von Brandt 1836
- ***** Genus Biziura Stephens 1824
- ***** Genus Pteronetta Salvadori 1895
- ***** Genus Marmaronetta Reichenbach 1853
- ***** Genus Asarcornis Salvadori 1895
- ***** Genus Netta Kaup 1829
- ***** Genus Lophonetta Riley 1914
- ***** Genus Amazonetta von Boetticher 1929
- ***** Tribe Oxyurini Swainson 1831
- ***** Genus †Anabernicula Ross 1935
- ****** Genus Heteronetta Salvadori 1865
- ****** Genus Nomonyx Ridgway 1880
- ****** Genus Oxyura Bonaparte 1828
- ***** Genus Nettapus von Brandt 1836
- ***** Genus Malacorhynchus Swainson 1831
- ***** Genus Salvadorina Rothschild & Hartert 1894
- ***** Genus Speculanas von Boetticher 1929
- ***** Tribe Mergini Rafinesque 1815
- ****** Genus †Chendytes Miller 1925
- ****** Genus †Shiriyanetta Watanabe & Matsuoka 2015
- ****** Genus †Camptorhynchus Bonaparte 1838
- ****** Genus Histrionicus Lesson 1828
- ****** Genus Clangula Leach 1819
- ****** Genus Polysticta stelleri Eyton 1836
- ****** Genus Somateria Leach 1819
- ****** Genus Melanitta Boie 1822
- ****** Genus Bucephala Baird 1858
- ****** Genus Mergellus Selby 1840
- ****** Genus Lophodytes Reichenbach 1853
- ****** Genus Mergus Linnaeus 1758 non Brisson 1760
- ***** Tribe Anatini Vigors 1825 sensu Livezey 1996
- ****** Genus †Matanas Worthy et al. 2007
- ****** Genus Anas Linnaeus 1758
- ****** Genus Sibirionetta
- ****** Genus Mareca
- ****** Genus Spatula Boie 1822
- ***** Tribe Aythyini Delacour and Mayr, 1945
- ****** Genus Aythya Boie 1822
- †Allgoviachen Mayr et al. 2022
- †"Anas" albae Jánossy 1979
- †"Anas" amotape Campbell 1979
- †"Anas" isarensis Lambrecht 1933
- †"Anas" luederitzensis Lambrecht 1929
- †"Anas" sanctaehelenae Campbell 1979
- †"Anas" eppelsheimensis Lambrecht 1933
- †Aldabranas cabri Harrison & Walker 1978
- †Ankonetta larriestrai Cenizo & Agnolín 2010
- †Bambolinetta Mayr & Pavia 2014
- †Cayaoa bruneti Tonni 1979
- †Chelychelynechen Olson & James 1991
- †Chenoanas deserta Zelenkov 2012
- †Cygnopterus alphonsi Cheneval 1984
- †Eoneornis Ameghino 1891
- †Eutelornis Ameghino 1891
- †Gracanicanetta Bochenski et al 2025
- †Helonetta brodkorbi Emslie 1992
- †Heteroanser vicinus Zelenkov 2012
- †Kustokazanser
- †Lavanttalornis Bochenski et al. 2023
- †Loxornis clivus Ameghino 1894
- Metopiana Bonaparte 1856
- †Mioquerquedula minutissima Zelenkov & Kuročkin 2012
- †Nogusunna conflictoides Zelenkov 2011
- †"Oxyura" doksana Mlíkovský 2002
- †Paracygnopterus scotti Harrison & Walker 1979
- †Proanser major Umanskaya 1979
- †Protomelanitta Zelenkov 2011
- †Ptaiochen Olson & James 1991
- †Sharganetta mongolica Zelenkov 2011
- †Sinanas Yeh 1980
- †Talpanas Olson & James 2009
- †Teleornis Ameghino 1899
- †Thambetochen Olson & Wetmore 1976
- †Uyrekura Zelenkov 2023
- †Wasonaka Howard 1966
- "Presbyornithidae" gen. et sp. indet. – Presbyornithidae?
- UCMP 117599
- Petropluvialis – may be same as Palaeopapia
- Agnopterus – includes Cygnopterus lambrechti
- "Headonornis hantoniensis" BMNH PAL 4989 – formerly "Ptenornis"
- Palaeopapia
- "Anas" creccoides
- "Anas" skalicensis
- "Anas" risgoviensis
- †"Anas" meyerii Milne-Edwards 1867
- †Eonessa anaticula Wetmore 1938
Phylogeny
| Anseriformes classification |
Molecular studies
Studies of the mitochondrial DNA suggest the existence of four branches – Anseranatidae, Dendrocygninae, Anserinae and Anatinae – with Dendrocygninae being a subfamily within the family Anatidae and Anseranatidae representing anindependent family. The clade Somaterini has a single genus Somateria.
Cited texts
- Murray, P. F. & Vickers-Rich, P. Magnificent Mihirungs: The Colossal Flightless Birds of the Australian Dreamtime. Indiana University Press.
Category:Bird orders
Category:Extant Maastrichtian first appearances
Category:Late Cretaceous taxonomic orders
Category:Paleocene taxonomic orders
Category:Eocene taxonomic orders
Category:Oligocene taxonomic orders
Category:Miocene taxonomic orders
Category:Pliocene taxonomic orders
Category:Pleistocene taxonomic orders
Category:Holocene taxonomic orders
Category:Taxa named by [Johann Georg Wagler]