Anseriformes

Anseriformes Temporal range: Eocene-Holocene, 54–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N [1]
Magpie goose, Anseranas semipalmata
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Aves
Clade:	Anserimorphae
Order:	Anseriformes Wagler, 1831
Extant families
Anhimidae Anseranatidae Anatidae
Range of the waterfowl and allies

Anseriformes /ˈænsərɪfɔːrmiːz/is an order of birds also known as waterfowl that comprises about 180 living species of birds in three families: Anhimidae (the 3 screamers), Anseranatidae (the magpie goose), 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. 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.^[2]

The earliest Cretaceous anseriform found so far is Vegavis, a goose-like waterfowl thought to have lived as long as 99 million years ago.^[3] 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 coevolve in each sex to advance control over mating and fertilization. Sexually antagonistic coevolution (or SAC) 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.^[4]

Taxonomy[]

The Anseriformes and the Galliformes (pheasants, etc.) are the most primitive neognathous birds, and should follow ratites and tinamous in bird classification systems. Together they belong to the Galloanserae. 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.^{[5][6][7][8][9][10]} 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.^[11]

Below is the general consensus of the phylogeny of anseriforms and their stem relatives.^{[5][6][7][8][9][11]}

Odontoanserae	†Pelagornithidae (pseudo-tooth birds)   Anserimorphae   †Gastornithidae       †Dromornithidae (mihirungs)       †Vegaviidae     Anseriformes (screamers and waterfowl)

Systematics[]

Anatidae 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.^[12] The Anatinae consists of tribes Anatini, Aythyini, Mergini and . The higher-order classification below follows a phylogenetic analysis performed by Mikko's Phylogeny Archive^[13][14] and John Boyd's website.^[15]

• Order Anseriformes
  • ?†Conflicto Claudia P. Tambussi et al. 2019
  • † Zelenkov 2019
  • Suborder Wetmore & Miller 1926
    • Genus †Chaunoides de Alvarenga 1999
    • Family Anhimidae Stejneger 1885 (screamers)
      • Genus Anhima (Linnaeus 1766) Brisson 1760 (Horned screamer)
      • Genus Chauna Illiger 1811
  • Suborder Anseres (true anseriformes)
    • Superfamily Anseranatoidea
      • Family Anseranatidae Sclater 1880
        • Genus † Mourer-Chauviré, Berthet & Hugueney 2004
        • Genus †Eoanseranas Worthy & Scanlon 2009 (Hand's dawn magpie goose)
        • Genus †Anatalavis Olson & Parris 1987 (Late Cretaceous/Early Paleocene – Early Eocene)
        • Genus Anseranas (Latham 1798) Lesson 1828 (Magpie goose)
    • Superfamily Anatoidea
      • Family †Presbyornithidae Wetmore 1926 (wading-"geese")^
        • Genus †Teviornis Kuročkin, Dyke & Karhu 2002
        • Genus † Howard 1955
        • Genus † (Lydekker 1891) Harrison & Walker 1976
        • Genus †Presbyornis Wetmore 1926
        • Genus †Wilaru Boles et al. 2013
      • Family † Miller & Compton 1939
        • Genus † Miller & Compton 1939 (Rosebud Early Miocene of Bennett County, USA)
          

          Anatidae
      • Family Anatidae Leach 1820 (almost 150 species)
        • Subfamily † Lambrecht 1933
          • Genus † Lebedinský 1927 (Late Eocene/Early Oligocene)
          • Genus † Mayr & De Pietri 2013
        • Subfamily Dendrocygninae Reichenbach 1849–50
          • Genus Dendrocygna Swainson 1837 (whistling ducks)
          • Genus Thalassornis Eyton 1838 (White-backed duck)
        • Subfamily † Livezey & Martin 1988
          • Genus † Miller 1944
          • Genus †Manuherikia Worthy et al. 2007
          • Genus † Livezey & Martin 1988
        • Subfamily Stictonettinae
          • Genus Stictonetta (Gould 1841) Reichenbach 1853 (Freckled duck)
        • Subfamily Anserinae Vigors 1825 sensu Livezey 1996 (swans and geese)
          • Genus † Kuročkin & Ganya 1972
          • Genus † Nesov 1986
          • Genus † De Vis 1905
          • Genus † Lambrecht 1931
          • Genus † Lambrecht 1931
          • Genus † Brodkorb 1961
          • Genus † Howard 1992
          • Genus † Short 1969
          • Genus † Wetmore 1930
          • Genus †Cnemiornis Owen 1866 (New Zealand Geese)
          • Genus Coscoroba (Molina 1782) Reichenbach 1853 (Coscoroba swan)
          • Genus Cereopsis Latham 1801 (Cape Barren goose)
          • Genus Cygnus Garsault 1764
          • Genus † Louchart et al. 2005
          • Genus Branta Scopoli 1769
          • Tribe Anserini Vigors 1825
            • Genus Anser Brisson 1760
        • Subfamily Tadorninae Reichenbach 1849–50 (shelducks and sheldgeese)
          • Genus †Australotadorna Worthy 2009
          • Genus † Howard 1964
          • Genus †Centrornis Andrews 1897 (Malagasy sheldgoose)
          • Genus †Miotadorna Worthy et al. 2007 (St. Bathans shelduck)
          • Genus † Campbell 1979
          • Genus † Agnolín 2006
          • Genus Plectropterus (Linnaeus 1766) (Spur-winged goose)
          • Genus Merganetta Gould 1842 (Torrent duck)
          • Genus Chloephaga Eyton 1838
          • Genus Neochen Oberholser 1918
          • Genus Cyanochen (Rüppell 1845) Bonaparte 1856 (Blue-winged goose)
          • Genus Tadorna Boie 1822
          • Genus Radjah Reichenbach, 1853
          • Genus Alopochen Stejneger 1885
          • Genus Cairina (Linnaeus 1758) Fleming 1822 (Muscovy duck)
          • Genus Hymenolaimus (Gmelin 1789) Gray 1843 (Blue duck)
          • Genus Sarkidiornis Eyton 1838
          • Genus Tachyeres Owen 1875 (Steamer ducks)
        • Subfamily Anatinae Vigors 1825 sensu Livezey 1996
          • Genus Aix Boie 1828
          • Genus Callonetta Delacour 1936 (Ringed teal)
          • Genus Chenonetta von Brandt 1836 (Australian wood duck)
          • Genus Biziura Stephens 1824 (musk ducks)
          • Genus Pteronetta (Cassin 1860) Salvadori 1895 (Hartlaub's duck)
          • Genus Marmaronetta (Ménétries 1832) Reichenbach 1853 (Marbled duck)
          • Genus Asarcornis (Müller 1842) Salvadori 1895 (White-winged duck)
          • Genus Netta Kaup 1829
          • Genus Lophonetta (King 1828) Riley 1914 (Crested duck)
          • Genus Amazonetta (Gmelin 1789) von Boetticher 1929 (Brazilian teal)
          • Genus †Dunstanetta Worthy et al. 2007 (Johnstone's duck)
          • Genus † Stidham & Hilton 2015
          • Genus †Pinpanetta Worthy 2009
          • Genus † Worthy 2008
          • Tribe Oxyurini Swainson 1831 (stiff-tailed ducks and allies)
            • Genus Heteronetta (Merrem 1841) Salvadori 1865 (Black-headed duck)
            • Genus Nomonyx (Linnaeus 1766) Ridgway 1880 (Masked duck)
            • Genus Oxyura Bonaparte 1828
          • Genus Nettapus von Brandt 1836 (Pygmy geese)
          • Genus †Anabernicula Ross 1935
          • Genus Malacorhynchus Swainson 1831 (Pink-eared duck)
          • Genus Salvadorina Rothschild & Hartert 1894 (Salvadori's teal)
          • Genus Speculanas (King 1828) von Boetticher 1929 (Bronze-winged duck)
          • Tribe Mergini Rafinesque 1815 (eiders, scoters, mergansers and other sea-ducks)
            • Genus †Chendytes Miller 1925
            • Genus †Shiriyanetta Watanabe & Matsuoka 2015
            • Genus Histrionicus Lesson 1828 (Harlequin duck)
            • Genus †Camptorhynchus (Gmelin 1789) Bonaparte 1838 (Labrador duck)
            • Genus Clangula Leach 1819 (Long-tailed duck)
            • Genus Polysticta stelleri (Pallas 1769) Eyton 1836 (Steller's eider)
            • Genus Somateria Leach 1819 (Eiders)
            • Genus Melanitta Boie 1822 (Scoters)
            • Genus Bucephala Baird 1858
            • Genus Mergellus Selby 1840 (Smew)
            • Genus Lophodytes (Linnaeus 1758) Reichenbach 1853 (Hooded merganser)
            • Genus Mergus Linnaeus 1758 non Brisson 1760
          • Tribe Anatini Vigors 1825 sensu Livezey 1996 (dabbling ducks and moa-nalos)
            • Genus †Matanas Worthy et al. 2007 (Enright's duck)
            • Genus Anas Linnaeus 1758
            • Genus Sibirionetta (Georgi 1775) (Baikal teal)
            • Genus Mareca (Stephens 1824)
            • Genus Spatula Boie 1822
          • Tribe Aythyini Delacour and Mayr, 1945 (diving ducks)
            • Genus Aythya Boie 1822

Some fossil anseriform taxa not assignable with certainty to a family are:

• † (London Clay Early Eocene of London, England) – Presbyornithidae?
• †Garganornis ballmanni Meijer 2014

Unassigned Anatidae:

• † Jánossy 1979 [?Mergus]
• † Campbell 1979
• † Lambrecht 1933
• †
• † Campbell 1979
• † Lambrecht 1933
• † Mlíkovský 2002
• † ["Anas" scaldii]
• †Ankonetta larriestrai Cenizo & Agnolín 2010
• † Tonni 1979
• † nomen dubium
• †
• † Harrison & Walker 1978
• †Chenoanas deserta Zelenkov 2012
• † Cheneval 1984 [non Mlíkovský 2002]
• † Emslie 1992
• † Ameghino 1894
• †Mioquerquedula minutissima Zelenkov & Kuročkin 2012 [ Milne-Edwards 1867]
• † Harrison & Walker 1979
• † Umanskaya 1979
• † Ameghino 1899
• † Zelenkov 2011
• † Zelenkov 2011
• † Zelenkov 2011
• Bonaparte 1856 [ Heine & Reichenow 1890; Delacour 1937; Delacour 1937]
• †Bambolinetta (Portis 1884) Mayr & Pavia 2014 [Anas lignitifila Portis 1884]
• † (Kuročkin 1976) Zelenkov 2012 [ Kuročkin 1976; (Kuročkin 1976) Mlíkovský & Švec 1986]
• †Sinanas Yeh 1980
• †Talpanas Olson & James 2009 (Kaua'i mole duck)
• † Howard 1966
• †Chelychelynechen Olson & James 1991 (turtle-jawed moa-nalo)
• †Ptaiochen Olson & James 1991 (small-billed moa-nalo)
• †Thambetochen Olson & Wetmore 1976

In addition, a considerable number of mainly Late Cretaceous and Paleogene fossils have been described where it is uncertain whether or not they are anseriforms. This is because almost all orders of aquatic birds living today either originated or underwent a major radiation during that time, making it hard to decide whether some waterbird-like bone belongs into this family or is the product of parallel evolution in a different lineage due to adaptive pressures.

• "Presbyornithidae" gen. et sp. indet. (Barun Goyot Late Cretaceous of Udan Sayr, Mongolia) – Presbyornithidae?
• UCMP 117599 (Hell Creek Late Cretaceous of Bug Creek West, USA)
• (Late Eocene of England) – may be same as Palaeopapia
• (Late Eocene – Late Oligocene of Europe) – includes Cygnopterus lambrechti
• "Headonornis hantoniensis" BMNH PAL 4989 (Hampstead Early Oligocene of Isle of Wight, England) – formerly "Ptenornis"
• (Hampstead Early Oligocene of Isle of Wight, England)
• "Anas" creccoides (Early/Middle Oligocene of Belgium)
• "Anas" skalicensis (Early Miocene of "Skalitz", Czech Republic)
• "Anas" risgoviensis (Late Miocene of Bavaria, Germany)
• † Milne-Edwards 1867 [ (Milne-Edwards 1867) Brodkorb 1964]
• †Eonessa anaticula Wetmore 1938 {Eonessinae Wetmore 1938}

Phylogeny[]

Living Anseriformes based on the work by John Boyd.^[15]

showAnseriformes classification

Anhimidae   Anhima     Chauna       Anseres Anseranatidae   Anseranas     Anatidae Dendrocygninae   Dendrocygna     Thalassornis       ?Plectropterinae   Plectropterus     Stictonettinae   Stictonetta       Anserinae ?   Nettapus       Biziura       Oxyurini   Heteronetta       Nomonyx     Oxyura       Anserini   Malacorhynchus         Coscoroba     Cereopsis         Sthenelides     Cygnus       Branta     Anser               Anatinae   Merganetta         Chloephaga           Neochen         Radjah       Tadorna     Alopochen           Mergini   Histrionicus       †Camptorhynchus       Clangula       Polysticta     Somateria       Melanitta       Bucephala       Mergellus       Lophodytes     Mergus                     Cairina     Aix       Callonetta         ?Salvadorina       Aythyini     Chenonetta       Hymenolaimus     Sarkidiornis             Pteronetta     Cyanochen         Marmaronetta       Asarcornis       Netta           Aythya               Anatini   Anas         Sibirionetta     Spatula         Tachyeres       Lophonetta       Amazonetta     Speculanas

• 

  Crested screamer (Chauna torquata)

• 

  Magpie goose (Anseranas semipalmata), sole surviving member of a Mesozoic lineage

• 

  Cast of Dromornis stirtoni, a mihirung, from Australia.

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 an independent family.^[16] The clade Somaterini has a single genus Somateria.

See also[]

• List of Anseriformes by population
• List of Anseriformes

References[]

Cited texts[]

Wikimedia Commons has media related to Anseriformes.

The Wikibook Dichotomous Key has a page on the topic of: Anseriformes

• Agnolin, F (2007). "Brontornis burmeisteri Moreno & Mercerat, un Anseriformes (Aves) gigante del Mioceno Medio de Patagonia, Argentina". Revista del Museo Argentino de Ciencias Naturales. 9: 15–25. doi:10.22179/revmacn.9.361.
• Clarke, J. A.; Tambussi, C. P.; Noriega, J. I.; Erickson, G. M.; Ketcham, R. A. (2005). "Definitive fossil evidence for the extant avian radiation in the Cretaceous". Nature. 433 (7023): 305–308. Bibcode:2005Natur.433..305C. doi:10.1038/nature03150. PMID 15662422. S2CID 4354309.
• Livezey, B. C.; Zusi, R. L. (2007). "Higher-order phylogeny of modern birds (Theropoda, Aves: Neornithes) based on comparative anatomy. II. Analysis and discussion". Zoological Journal of the Linnean Society. 149 (1): 1–95. doi:10.1111/j.1096-3642.2006.00293.x. PMC 2517308. PMID 18784798.
• Murray, P. F. & Vickers-Rich, P. (2004) Magnificent Mihirungs: The Colossal Flightless Birds of the Australian Dreamtime. Indiana University Press.