Anatidae

The ducks, geese, and swans are small- to large-sized birds with a broad and elongated general body plan. Diving species vary from this in being rounder. Extant species range in size from the cotton pygmy goose, at as little as 26.5 cm (10.5 in) and 164 g (5.8 oz), to the trumpeter swan, at as much as 183 cm (6 ft) and 17.2 kg (38 lb). The largest anatid ever known is the extinct flightless Garganornis ballmanni at 22 kg (49 lb). The wings are short and pointed, and supported by strong wing muscles that generate rapid beats in flight. They typically have long necks, although this varies in degree between species. The legs are short, strong, and set far to the back of the body (more so in the more aquatic species), and have a leathery feel with a scaly texture. Combined with their body shape, this can make some species awkward on land, but they are stronger walkers than other marine and water birds such as grebes or petrels. They typically have webbed feet, though a few species such as the Nene have secondarily lost their webbing. The bills are made of soft keratin with a thin and sensitive layer of skin on top (which has a leathery feel when touched). For most species, the shape of the bill tends to be more flattened to a greater or lesser extent. These contain serrated lamellae which are particularly well defined in the filter-feeding species. Their feathers are excellent at shedding water due to special oils. Many of the ducks display sexual dimorphism, with the males being more brightly coloured than the females (although the situation is reversed in species such as the paradise shelduck). The swans, geese, and whistling-ducks lack sexually dimorphic plumage. Anatids are vocal birds, producing a range of quacks, honks, squeaks, and trumpeting sounds, depending on species; the female often has a deeper voice than the male. Anatids are generally herbivorous as adults, feeding on various water-plants, although some species also eat fish, molluscs, or aquatic arthropods. One group, the mergansers, are primarily piscivorous, and have serrated bills to help them catch fish. In a number of species, the young include a high proportion of invertebrates in their diets, but become purely herbivorous as adults. ==Breeding==

The anatids are generally seasonal and monogamous breeders. The level of monogamy varies within the family; many of the smaller ducks only maintain the bond for a single season and find a new partner the following year, whereas the larger swans, geese and some of the more territorial ducks maintain pair bonds over a number of years, and even for life in some species. However, forced extrapair copulation among anatids is common, occurring in 55 species in 17 genera. though they vary significantly in size, shape, and surface elaboration. Most species are adapted for copulation on the water only. They construct simple nests from whatever material is close at hand, often lining them with a layer of down plucked from the mother's breast. In most species, only the female incubates the eggs. The young are precocial, and are able to feed themselves from birth. One aberrant species, the black-headed duck, is an obligate brood parasite, laying its eggs in the nests of gulls and coots. While this species never raises its own young, a number of other ducks occasionally lay eggs in the nests of conspecifics (members of the same species) in addition to raising their own broods. ==Relationship with humans==

Duck, eider, and goose feathers and down have long been popular for bedspreads, pillows, sleeping bags, and coats. The members of this family also have long been used for food. Humans have had a long relationship with ducks, geese, and swans; they are important economically and culturally to humans, and several duck species have benefited from an association with people. However, some anatids are agricultural pests, and have acted as vectors for zoonoses such as avian influenza. Since 1600, five species of ducks have become extinct due to the activities of humans, and subfossil remains have shown that humans caused numerous extinctions in prehistory. Today, many more are considered threatened. Most of the historic and prehistoric extinctions were insular species, vulnerable due to small populations (often endemic to a single island), and island tameness. Evolving on islands that lacked predators, these species lost antipredator behaviours, as well as the ability to fly, and were vulnerable to human hunting pressure and introduced species. Other extinctions and declines are attributable to overhunting, habitat loss and modification, and hybridisation with introduced ducks (for example the introduced ruddy duck swamping the white-headed duck in Europe). Numerous governments and conservation and hunting organisations have made considerable progress in protecting ducks and duck populations through habitat protection and creation, laws and protection, and captive-breeding programmes. ==Systematics==

(Anas crecca), gadwall (Anas strepera), northern pintail (Anas acuta), mallard (Anas platyrhynchos), greater scaup (Aythya marila), long-tailed duck (Clangula hyemalis), northern shoveler (Anas clypeata), garganey (Anas querquedula), Eurasian wigeon (Anas penelope), ferruginous duck (Aythya nyroca), common goldeneye (Bucephala clangula), common merganser (Mergus merganser), smew (Mergellus albellus), tufted duck (Aythya fuligula), red-breasted merganser (Mergus serrator), common pochard (Aythya ferina). Post of Belarus, 1996. History of classification The name Anatidae for the family was introduced by the English zoologist William Elford Leach in a guide to the contents of the British Museum published in 1819. While the status of the Anatidae as a family is straightforward, and which species properly belong to it is little debated, the relationships of the different tribes and subfamilies within it are poorly understood. The listing in the box at right should be regarded as simply one of several possible ways of organising the many species within the Anatidae; see discussion in the next section. The systematics of the Anatidae are in a state of flux. Previously divided into six subfamilies, a study of anatomical characters by Livezey suggests the Anatidae are better treated in nine subfamilies. This classification was popular in the late 1980s to 1990s. But mtDNA sequence analyses indicate, for example, the dabbling and diving ducks do not belong in the same subfamily. While shortcomings certainly occur in Livezey's analysis, mtDNA is an unreliable source for phylogenetic information in many waterfowl (especially dabbling ducks) due to their ability to produce fertile hybrids, in rare cases possibly even beyond the level of genus (see for example the "Barbary duck"). Because the sample size of many molecular studies available to date is small, mtDNA results must be considered with caution. While a comprehensive review of the Anatidae which unites all evidence into a robust phylogeny is still lacking, the reasons for the confusing data are at least clear: As demonstrated by the Late Cretaceous fossil Vegavis iaai—an early modern waterbird which belonged to an extinct lineage—the Anatidae are an ancient group among the modern birds. Their earliest direct ancestors, though not documented by fossils yet, likewise can be assumed to have been contemporaries with the non-avian dinosaurs. The long period of evolution and shifts from one kind of waterbird lifestyle to another have obscured many plesiomorphies, while apparent apomorphies are quite often the result of parallel evolution, for example the "non-diving duck" type displayed by such unrelated genera as Dendrocygna, Amazonetta, and Cairina. For the fossil record, see below. Alternatively, the Anatidae may be considered to consist of three subfamilies (ducks, geese, and swans, essentially) which contain the groups as presented here as tribes, with the swans separated as subfamily Cygninae, the goose subfamily Anserinae also containing the whistling ducks, and the Anatinae containing all other clades. ==Genera==

For the living and recently extinct members of each genus, see the article List of Anatidae species. • Subfamily: Dendrocygninae (one pantropical genus, of distinctive long-legged goose-like birds) • Dendrocygna, whistling ducks (8 living species) • Thalassornis, white-backed duck • Subfamily: Anserinae, swans and geese (3–7 extant genera with 25–30 living species, mainly cool temperate Northern Hemisphere, but also some Southern Hemisphere species, with the swans in one genus [two genera in some treatments], and the geese in three genera [two genera in some treatments]. Some other species are sometimes placed herein, but seem somewhat more distinct [see below]) • Cygnus, true swans (6 species, 4 sometimes separated in Olor) • Anser, grey geese and white geese (11 species) • Branta, black geese (6 living species) • Subfamily: Stictonettinae (one genus in Australia, formerly included in the Oxyurinae, but with anatomy suggesting a distinct ancient lineage perhaps closest to the Anserinae, especially the Cape Barren goose) • Stictonetta, freckled duck • Subfamily: Plectropterinae (one genus in Africa, formerly included in the "perching ducks", but closer to the Tadorninae) • Plectropterus, spur-winged goose • Subfamily: Tadorninae – shelducks and sheldgeese (This group of larger, often semiterrestrial waterfowl can be seen as intermediate between Anserinae and Anatinae. The 1986 revision) • Spatula, shovelers • Mareca, wigeons and gadwalls • Lophonetta, crested duck • Speculanas, bronze-winged duck • Amazonetta, Brazilian teal • Sibirionetta, Baikal teal • Chelychelynechen, turtle-jawed moa-nalo (prehistoric) • Thambetochen, large-billed moa-nalos (2 species, prehistoric) • Ptaiochen, small-billed moa-nalo (prehistoric) • Tribe: Mergini, eiders, scoters, sawbills and other sea-ducks couple, male on the right (There are 9 extant genera and some 20 living species; most of this group occur in the Northern Hemisphere, but a few [mostly extinct] mergansers in the Southern Hemisphere) • Shiriyanetta (prehistoric) • Polysticta, Steller's eider • Somateria, eiders (3 species) • Histrionicus, harlequin duck (includes Ocyplonessa) • Camptorhynchus, Labrador duck (extinct) • Melanitta, scoters (6 species) • Clangula, long-tailed duck (1 species) • Bucephala, goldeneyes (3 species) • Mergellus, smew • Lophodytes, hooded merganser • Mergus, mergansers (4 living species, 1 extinct). • Tribe: Oxyurini, stiff-tail ducks (a small group of 3–4 genera, 2–3 of them monotypic, with 7–8 living species) • Oxyura, stiff-tailed ducks (5 living species) • Nomonyx, masked duck • Heteronetta, black-headed duck • Unresolved: The largest degree of uncertainty concerns whether a number of genera are closer to the shelducks or to the dabbling ducks., a species of unclear affiliation Aix sponsa See also the monotypic subfamilies above, and the "perching ducks" • Coscoroba, coscoroba swan – Anserinae or same subfamily as Cereopsis? • Cereopsis, Cape Barren goose – Anserinae, Tadorninae, or own subfamily? • Biziura, musk ducks (1 living species) - Oxyurini? • Cnemiornis, New Zealand geese (prehistoric) – as Cereopsis • Malacorhynchus, pink-eared ducks (1 living species) – Tadorninae, Oxyurinae or Dendrocheninae? • Sarkidiornis, comb duck – Tadorninae or closer to dabbling ducks? • Tachyeres, steamer ducks (4 species) – Tadorninae or closer to dabbling ducks? • Cyanochen, blue-winged goose – Tadorninae or more distant clade? • Nettapus, pygmy geese (3 species) – Anatinae or part of Southern Hemisphere radiation? • Pteronetta, Hartlaub's duck – traditionally dabbling ducks, but may be closer to Cyanochen • Cairina and Asarcornis, Muscovy duck and white-winged duck, respectively (2 species) – traditionally dabbling ducks, but may be paraphyletic, with one species in Tadorninae and the other closer to diving ducks • Aix, Mandarin duck and wood duck (2 species) – dabbling ducks or Tadorninae? • Callonetta, ringed teal – dabbling ducks or Tadorninae? • Chenonetta, maned duck (1 living species) – dabbling ducks or Tadorninae? Includes Euryanas. • Marmaronetta, marbled duck – formerly dabbling ducks; actually a diving duck or a distinct subfamily Prehistoric species is the only living member of the genus Chenonetta. From subfossil bones found on Kauai (Hawaiian Islands), two enigmatic waterfowl are known. • Ankonetta larriestrai (Miocene of Argentina) • "Anser" scaldii (Late Miocene of Antwerp, Belgium) – anserine or tadornine* Anatidae gen. et sp. indet. (Waite Late Miocene of Alcoota, Australia) – anatine, oxyurine? • "Anas" velox (Middle–Late? Miocene of C Europe) – anatine? May include "A." meyerii • Anatidae gen. et sp. indet. (Waite Late Miocene of Alcoota, Australia) – tadornine? • Anatidae gen. et sp. indet. MNZ S42797 (Bathans Early/Middle Miocene of Otago, New Zealand) • Anatidae gen. et sp. indet. (Middle Miocene of Nördlinger Ries, Germany) – tadornine? • Anatidae gen. et sp. indet. (Sajóvölgyi Middle Miocene of Mátraszõlõs, Hungary) The middle Eocene Eonessa was formerly thought to belong to Anatidae, however reexamination of the holotype in 1978 resulted in the genus being placed as Aves incertae sedis. ==See also==