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Anas

Female Mallard (Anas platyrhynchos) with brood of young, a typical member of this genus.

Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Subclass: Neornithes
Infraclass: Neognathae
Superorder: Galloanserae
Order: Anseriformes
Family: Anatidae
Subfamily: Anatinae
Genus: Anas
Linnaeus, 1758
Species

Some 40-50, but see text.

Anas is a genus of dabbling ducks.It includes mallards, wigeons, teals, pintails and shovelers in a number of subgenera. Some authorities prefer to elevate the subgenera to genus rankCarboneras (1992). Indeed, as the moa-nalos are very close to this clade and may have evolved later than some of these lineages, it is rather the absence of a thorough review than lack of necessity that this genus is rather over-lumped.

Contents

Systematics

The phylogeny of this genus is one of the most confounded ones of all living birds. Research is hampered by the fact the radiation of the two major groups of Anas - the teals and mallard groups -; took place in a very short time and fairly recently, roughly in the mid-late Pleistocene. Furthermore, hybridization probably has long played a major role in Anas evolution, with within-subgenus hybrids regularly and between-subgenus hybrids not infrequently being fully fertileCarboneras (1992); see also Mariana Mallard.. The relationships between species are much obscured by this fact, and mtDNA sequence data is of dubious value in resolving their relationshipse.g. Kulikova et al. (2005); on the other hand, nuclear DNA sequences evolve too slowly to resolve the phylogeny of the subgenus Anas for example.

Some major clades can be discerned. For example, that the traditional subgenus Anas, the mallard group, forms a monophyletic (in the loose sense, i.e. non-holophyletic) group has never been seriously questioned by modern science and is as good as confirmed (but see below). On the other hand, the phylogeny of the teals is very confusing.

It is fairly clear by now that the dabbling duck lineages more distantly related to mallard group (which includes the type species of Anas) than the wigeons are should arguably be separated in their own genera. These would include the Baikal Teal, the Garganey, the spotted black-capped Punanetta group, and the shovelers and other blue-winged species. Whether the widgeons, which are very distinct in morphologyLivezey (1991) and behaviorJohnson et al. (2000), but much less so in mtDNA cytochrome b and NADH dehydrogenase subunit 2 sequencesJohnson & Sorenson (1999), should also be considered a distinct genus Mareca (including the Gadwall and Falcated Duck) is essentially the one remaining point of dispute as regards the question which taxa should remain in this genus and which ones should not.

Species

The following arrangement is based on morphologicalLivezey (1991), molecularJohnson & Sorenson (1999), McCracken et al. (2001) and behavioralJohnson et al. (2000) characters and presents apparent major evolutionary groupings compared to the subgenera the species were placed in at one time or another.

The puzzling Baikal Teal (Anas formosa, male)

Probable genus N.N.

Probable genus Querquedula - Garganey (may include Punanetta)

Probable genus Punanetta

Red Shoveler, Anas platalea

Probable genus Spatula - blue-winged ducks/shovelers and allies (polyphyletic?)

Male American Wigeon, Anas americana

Possible genus Mareca - wigeons (may include Chaulelasmus and Eunetta)

Subgenus Chaulelasmus - Gadwall

Subgenus Eunetta - Falcated Duck

Subgenus Dafila - pintails

Subgenus Nettion - teals (paraphyletic)

Green-winged Teal, Anas carolinensis

  • Green-winged/Red-and-green head clade

Subgenus Melananas - African Black Duck

Yellow-billed Duck, Anas undulata

Subgenus Anas - mallard and relatives (may include Melananas)

The last male Mariana Mallard, c.1980

  • American clade
    • Mottled Duck, Anas fulvigula - sometimes included in Anas platyrhynchos
      • Florida Duck, Anas fulvigula fulvigula - sometimes included in Anas platyrhynchos
    • American Black Duck, Anas rubripes - sometimes included in Anas platyrhynchos
    • Mexican Duck, Anas diazi - sometimes included in Anas platyrhynchos
  • Ambiguous
    • Mallard, Anas platyrhynchos
    • Spotbill, Anas poecilorhyncha
      • Chinese Spotbill, Anas (poecilorhyncha) zonorhyncha - sometimes considered a subspecies of Anas superciliosa

Formerly placed in Anas:

A number of fossil species of Anas have been described. Their relationships are often undetermined:

  • Anas sp. (Late Miocene of Rudabánya, Hungary)A mid-sized species: Bernor et al. (2002)
  • Anas greeni (Ash Hollow Late Miocene?/Early Pliocene of South Dakota, USA) - Nettion red-and-green head clade (doubtful)?
  • Anas ogallalae (Ogalalla Late Miocene?/Early Pliocene of Kansas, USA) - Nettion red-and-green head clade (doubtful)?
  • Anas pullulans (Juntura Late Miocene?/Early Pliocene of Juntura, USA) - Punanetta?
  • Anas cheuen (Early-Middle Pleistocene of Argentina) - Dafila?
  • Anas bunkeri (Early -? Middle Pliocene - Early Pleistocene of WC USA) - Nettion red-and-green head clade?
  • Anas pachyscelus (Shore Hills Late Pleistocene of Bermuda, W Atlantic)
  • Anas schneideri (Late Pleistocene of Little Box Elder Cave, USA)

Several prehistoric waterfowl supposedly part of the Anas assemblage are now placed elsewhere:

  • "Anas" basaltica (Late Oligocene of "Warnsdorf", Czechia) is apparently an indeterminate heron.
  • "Anas" blanchardi, "A." consobrina, "A." natator are now in Mionetta
  • "Anas" creccoides (Early-mid Oligocene of Belgium), "A." risgoviensis (Late Miocene of Bavaria, Germany) and "A." skalicensis (Early Miocene of "Skalitz", Czechia), though possibly anseriform, cannot be placed with any certainty among modern birds at all.
  • "Anas" eppelsheimensis (Early Pliocene of Eppelsheim, Germany), "A." isarensis (Late Miocene of Aumeister, Germany) and "A." luederitzensis (Kalahari Early Miocene of Lüderitzbucht, Namibia) are Anatidae of unclear affiliations.
  • "Anas" integra, "A." oligocaena are now in Dendrochen
  • "Anas" robusta is now tentatively placed in Anserobranta
  • "Anas" velox (Middle - Late? Miocene of C Europe) and "A." meyerii (Middle Miocene of Öhningen, Germany; possibly the same species) do not seem to belong into the present genus either; they may still turn out to be ancestral dabbling ducks.

Highly problematic, albeit in a theoretical sense, is the placement of the moa-nalos. These are in all probability derived from a common ancestor of the Pacific Black Duck, the Laysan Duck, and the Mallard, and an unknown amount of other lineages. Phylogenetically, they may even form a clade within the traditional genus AnasSorenson (1999). However, as opposed to these species - which are well representative of dabbling ducks in general - the moa-nalos are the most radical departure from the anseriform bauplan known to science. This illustrates that in a truly evolutionary sense, a strictly phylogenetic taxonomy may be difficult to apply.

Footnotes

References

  • Bernor, R.L.; Kordos, L. & Rook, L. (eds): Recent Advances on Multidisciplinary Research at Rudabánya, Late Miocene (MN9), Hungary: A compendium. Paleontographica Italiana 89: 3-36. PDF fulltext
  • Carboneras, Carles (1992): Family Anatidae (Ducks, Geese and Swans). In: del Hoyo, Josep; Elliott, Andrew & Sargatal, Jordi (editors): Handbook of Birds of the World, Volume 1: Ostrich to Ducks: 536-629. Lynx Edicions, Barcelona. ISBN 84-87334-10-5
  • Johnson, Kevin P. & Sorenson, Michael D. (1999): Phylogeny and biogeography of dabbling ducks (genus Anas): a comparison of molecular and morphological evidence. Auk 116(3): 792–805. PDF fulltext
  • Johnson, Kevin P. McKinney, Frank; Wilson, Robert & Sorenson, Michael D. (2000): The evolution of postcopulatory displays in dabbling ducks (Anatini): a phylogenetic perspective. Animal Behaviour 59(5): 953–963 PDF fulltext
  • Kulikova, Irina V.; Drovetski, S. V.; Gibson, D. D.; Harrigan, R. J.; Rohwer, S.; Sorenson, Michael D.; Winker, K.; Zhuravlev, Yury N. & McCracken, Kevin G. (2005): Phylogeography of the Mallard (Anas platyrhynchos): Hybridization, dispersal, and lineage sorting contribute to complex geographic structure. Auk 122(3): 949-965. [English with Russian abstract] DOI: 10.1642/0004-8038(2005)122[0949:POTMAP]2.0.CO;2 PDF fulltext Erratum: Auk 122(4): 1309. DOI: 10.1642/0004-8038(2005)122[0949:POTMAP]2.0.CO;2
  • Livezey, B. C. (1991): A phylogenetic analysis and classification of recent dabbling ducks (Tribe Anatini) based on comparative morphology. Auk 108(3): 471–507. PDF fulltext
  • McCracken, Kevin G.; Johnson, William P. & Sheldon, Frederick H. (2001): Molecular population genetics, phylogeography, and conservation biology of the mottled duck (Anas fulvigula). Conservation Genetics 2(2): 87–102. doi:10.1023/A:1011858312115 PDF fulltext
  • Sorenson et al (1999): Relationships of the extinct moa-nalos, flightless Hawaiian waterfowl, based on ancient DNA. Proceedings of the Royal Society.

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