The Anatidae are the biological family of water birds that includes ducks, geese, and swans. The family has a cosmopolitan distribution, occurring on all the world's continents except Antarctica. These birds are adapted for swimming, floating on the water surface, and, in some cases, diving in at least shallow water. The family contains around 174 species in 43 genera.
They are generally herbivorous and are monogamous breeders. A number of species undertake annual migrations. A few species have been domesticated for agriculture, and many others are hunted for food and recreation. Five species have become extinct since 1600, and many more are threatened with extinction.
Description and ecology 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.
Anatidae is a large proportion of the 3% of bird species to possess a penis, 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
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. The name of the author is not specified in the document, Leach was the Keeper of Zoology at the time. 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 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 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.
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 has resulted in the inclusion of 10 extant genera with about two-dozen living species [one probably extinct] in this subfamily, mostly from the Southern Hemisphere but a few in the Northern Hemisphere; the affiliations of several presumed tadornine genera has later been questioned and the group in the traditional lineup is likely to be paraphyletic.) Tadorna, shelducks (6 species, 1 probably extinct) – possibly paraphyletic Radjah, Radjah shelduck Salvadorina, Salvadori's teal Centrornis, Madagascar sheldgoose (prehistoric, tentatively placed here) Alopochen, Egyptian goose and Mascarene shelducks (1 living species, 2 extinct) Neochen, (2 species) Chloephaga, sheldgeese (4 species) Hymenolaimus, blue duck Merganetta, torrent duck
Subfamily: Aythyinae, diving ducks (Some 15 species of diving ducks, of worldwide distribution, in two to four genera; The 1986 morphological analysis suggested the probably extinct pink-headed duck of India, previously treated separately in Rhodonessa, should be placed in Netta, but this has been questioned. Furthermore, while morphologically close to dabbling ducks, the mtDNA data indicate a treatment as distinct subfamily is indeed correct, with the Tadorninae being actually closer to dabbling ducks than the diving ducks) Netta, red-crested pochard and allies (4 species, 1 probably extinct) Aythya, pochards, scaups, etc. (12 species) Subfamily: Anatinae, dabbling ducks and moa-nalos (The dabbling duck group, of worldwide distribution, were previously restricted to just one or two genera, but had been extended to include eight extant genera and about 55 living species, including several genera formerly known as the "perching ducks"; mtDNA on the other hand confirms that the genus Anas is over-lumped and casts doubt on the diving duck affiliations of several genera [see below]. The moa-nalos, of which four species in three genera are known to date, are a peculiar group of flightless, extinct anatids from the Hawaiian Islands. Gigantic in size and with massive bills, they were believed to be geese, but have been shown to be actually very closely related to mallards. They evolved filling the ecological niche of turtles, ungulates, and other megaherbivores. Anas: pintails, mallards, etc. (40–50 living species, 3 extinct) Chendytes, diving-geese (extinct c. 450–250 BCE, A basal member of the dabbling duck clade) 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)