anseriform

▪ bird order

Introduction

 any of the 150 species comprising the bird order Anseriformes, which comprises the ducks (duck), geese (goose), and swans (swan) (family Anatidae) and the screamers (the three species of family Anhimidae). Anatidae comprises about 147 species of medium to large birds, usually associated with freshwater or marine habitats. This family is known collectively as waterfowl (in the United States) or wildfowl (in Europe). The three species of screamers are quite different from waterfowl in general appearance. They are moderately long-legged birds about the size of a turkey, with chickenlike beaks and exceptionally large feet.

      Anseriforms are of interest for various reasons. Not only do their mass migrations symbolize the change of seasons to peoples in temperate areas of the world but their annual voyages have long been studied by ornithologists (ornithology) to determine the mechanisms that make these navigational and metabolic feats possible. To the sport hunter many ducks and geese are challenging and worthwhile quarry; domesticated members of the order are reared for meat and feathers, but to the farmer other species are considered pests and crop spoilers. Both domestic and wild anseriforms are occasionally reservoirs of disease. The conservationist views them as biological indicators of the quality of wetlands and, like those habitats, in need of attention.

General features

      Members of the Anatidae vary greatly in size, the largest being some 60 times heavier than the smallest. Most are rather chunky birds, from about 30 cm (1 foot) in length and 250 grams (0.5 pound) in weight in the African pygmy goose (Nettapus auritus) to 1.5 metres (5 feet) in length and weighing more than 17 kg (37 pounds) with a 2-metre (6.6-foot) wingspan in the trumpeter swan (Cygnus buccinator). The neck is medium to long. The bill (beak) is of medium length, typically broad and somewhat flattened, with a rounded tip, often with a pronounced “nail,” giving a slight hook to the tip. The form of the bill, however, varies widely, befitting birds with diets as different as grass, fish, and minute plankton. The bill is often fringed with fine lamellae, short parallel plates at the edges of the mandibles that aid in food handling and, in some species, in straining tiny food organisms from water or mud.

      Anseriform behaviour patterns are an intriguing mixture of the innate and the learned. Some behaviours are so fixed that they can be used as taxonomic characteristics, whereas others are so pliable that completely new traditions can evolve within a few generations.

 Anseriforms breed on every major continent and island except in the Antarctic, and many undertake vast migrations, flying thousands of kilometres. Some species are widespread and number in the millions; others are represented by a few hundred individuals confined to a few islands. In the former category are the pintail (Anas acuta) and the mallard (Anas platyrhynchos), both found throughout the Northern Hemisphere; in the latter are the Hawaiian goose (nene), or nene (Branta sandvicensis), and the Madagascar white-eye (Aythya innotata). extinction has taken at least six species within the last century, with another three likely extinct, having not been seen for a number of years. The Hawaiian goose was once down to fewer than 50 birds, the Laysan race of the mallard to 7 individuals. The creation of wetland refuges has helped to restore these and other declining populations. Nonetheless, even the more common waterfowl, which teem in thousands at certain times and places, are not secure from drastic declines in numbers.

Importance to humans

      The large size of waterfowl has made them from prehistoric times inevitable quarry for humans. Not only do they provide protein but also large amounts of fat and feathers, especially down, much prized in colder regions. Among the unusual uses of waterfowl parts may be mentioned the conversion of swan tracheae into children's whistles (whistle) in Lapland and the eating of the of the king eider's (Somateria spectabilis) billknob as an aphrodisiac in Greenland. Wary and difficult to approach in their watery haunts, waterfowl required ingenuity to take them before the advent of efficient weapons. The period of flightlessness was discovered early and exploited by driving the birds into corrals of stone or netting. From the latter evolved the Dutch method of catching full-winged ducks by enticing them up large net-covered pipes leading from a secluded pond where a decoy duck was placed. Many other ingenious traps were devised, from the clapnets of the ancient Egyptians to the rocket-propelled nets of today's research workers.

      In view of their ready adaptation to captivity, it is surprising that few waterfowl have been domesticated. The mallard (Anas platyrhynchos) was exploited 2,000 years ago in China, and 17 varieties have been developed, according to whether meat or egg production is important. The muscovy duck (Cairina moschata) was domesticated in Colombia and Peru before the arrival of the conquistadores. The greylag goose (Anser anser) has been domesticated for at least 4,000 years; Egyptian frescoes of that age already show changes in shape from the natural form, and eight main varieties are now known. The swan goose (Anser cygnoides) of eastern Asia has also been domesticated, with three varieties. Other species, such as the Canada goose (Branta canadensis), the mute swan, and the Egyptian goose (Alopochen aegyptiacus), have been kept in semidomestication for ease of exploitation but without intensive breeding to change their forms. A remarkable form of exploitation has been that of the common eider (Somateria mollissima). Its breeding colonies in the Arctic and subarctic are protected and concentrated by the provision of nest sites and other techniques. The down with which the female lines her nest and covers the eggs is systematically collected during and after incubation without disturbance or loss of productivity, for these birds are very tame.

      Anseriform populations are threatened by the loss of their essential wetland habitats, and they are no longer able to withstand commercial exploitation. This is generally recognized in a great many countries where market hunting is banned and efforts are made to control sport hunting so that annual kill does not exceed annual production. As waterfowl migrations pay no heed to national boundaries, the need for international agreement on their conservation is paramount.

      A great flock of waterfowl is one of the remaining spectacles of nature and one that can be made accessible to crowds of people led through covered approaches to concealing blinds. Such refuges must be numerous and large enough to give adequate feeding grounds. Extra food, however, can be provided as standing crops or distributed from stores. In these ways damage to agricultural interests can be avoided. Complaints of such damage are frequently exaggerated, for in many countries migratory waterfowl arrive after the harvests have been gathered and take only spilled or rejected grain and tubers, thus actually performing a useful cleaning operation. Conflicts can arise, but these can be resolved by changes in husbandry and by techniques of scaring the birds from the fields without injuring them.

      Anseriforms spend much time walking, swimming, or diving, so they are less inconvenienced than most other birds by being pinioned (removal of the tip of one wing) to prevent flight. They can then therefore be kept in open paddocks instead of being confined in cages. Zoos and private aviculturists maintain waterfowl collections of varying size, the largest being that of The Wildfowl & Wetlands Trust (Wildfowl Trust, The), Slimbridge, England. There almost 200 species have been maintained, and more than 100 have bred. Such a remarkably complete comparative collection has been useful for educational and research purposes. Breeding in captivity also enhances the possibility of restoring wild populations that have been diminished. The most notable case has been that of the Hawaiian goose (nene) (Branta sandvicensis). In 1950 about 30 of these birds existed. Twenty years later, thanks to avicultural efforts at Slimbridge and in Hawaii, there were more than a thousand, several hundred of which were released in original habitat, where they have sustained their numbers. Hunting organizations such as Ducks Unlimited conserve wetlands to maintain populations that can replace birds shot. Several species of geese and ducks have responded well to the manipulation of habitat and the provision of nest sites. Artificially encouraged breeding may well become vital as the hitherto-untouched vastness of northern marshes and tundras, where so many waterfowl breed, are opened up, destroyed, or polluted.

Natural history

Life history

      Hatchlings emerge from the egg with a complete covering of down and can take to the water as soon as they leave the nest, within 24 to 48 hours. They can forage for themselves, but at least one parent remains with them, guarding, guiding, and, initially, brooding them at night and during inclement weather.

      The downy plumage is retained for from two weeks (small ducks) to six weeks (large swans) and is then replaced gradually by the feathers of the juvenal plumage. The flight feathers, the last to develop, likewise vary in their rate of growth, taking from five weeks to five months. At fledging, young ducks must make their own way on migration. In species breeding in the far north, this begins in early autumn. Young geese and swans, on the other hand, remain with their parents during their first winter and migrate to and from the wintering grounds in their company. In most large species the juvenal plumage is retained through much of the first year of life. Ducks, however, begin to lose the juvenal body feathers almost at once. Some replace the juvenal plumage with an immature nonbreeding (or “basic”) plumage, acquiring the first nuptial (or “alternate”) plumage in the second autumn. Other species molt directly from juvenal to nuptial and are practically indistinguishable from adults in plumage and size at the age of six months. Swans and geese do not reach full size until the end of their second year, and even at that age swans still retain some immature feathers.

      In migratory species social flocking and pair formation occur on the wintering grounds, followed by the return migration to the breeding grounds. In the high Arctic regions the birds arrive ready to nest as soon as the snow cover melts and the water opens; in lower latitudes the process is more leisurely. When the female duck begins to incubate (incubation), the male generally deserts her and joins forces with other males, often after making a molt-migration to another area some distance from the breeding site. The nuptial plumage is lost, and a dull “eclipse” plumage, rather femalelike, is assumed before the simultaneous molt of the flight feathers. The resulting flightless condition lasts three or four weeks, during which the birds skulk in thick cover or remain on large bodies of water. In female ducks this wing molt is delayed until after the young have fledged. The eclipse plumage is succeeded in three to six months by the next nuptial plumage. In the case of swans and geese the male remains and molts with the female and family; immature birds and unsuccessful breeders may, however, make a molt-migration to a separate area.

      The foregoing life cycles are typical of waterfowl breeding in north temperate or Arctic regions. In the tropics and south temperate regions, migration of waterfowl may not be necessary. When it does occur, migration may be in response to ecological factors such as the onset of the rainy season. Males and females tend to be similar in plumage and to lack eclipse plumages. The birds are ready to breed at any time that conditions are favourable. Undoubtedly there is a relationship between hormonal balance and the appearance or absence of eclipse or brightly coloured plumages, but the exact mechanism is not yet known.

      The mature anseriform has a potentially long life, though this is seldom achieved in the wild. In captivity ducks have lived for 20 years, geese and swans for more than 30; there are reports of geese exceeding 40 years of age and swans, 50. Wild populations suffer heavy losses (up to 70 percent) among first-year birds. The adults then experience annual mortalities of 10 to 50 percent, depending on the extent of hunting and on natural factors. At the higher end of this scale the population is almost completely replaced within three years. With luck and cunning an occasional bird may survive for 15 to 20 years or more.

Behaviour

      Keeping the plumage waterproof occupies much of the time not spent feeding or sleeping. The bill (beak) is used both to stimulate the oil gland (preen gland) (situated above the tail) and to spread the oil. Rubbing the chin and throat on oiled areas also helps the process. Preening occurs at the same time, the fine structure of the feathers being nibbled into the interlocking position necessary to prevent the entry of water. Rearrangement of the feathers involves preening, scratching with the feet, and a general body shake produced by a muscular contraction sweeping from tail to neck. Various wing-stretching movements settle the flight feathers. Bathing movements include dipping the head, beating the wings on the surface and, at high intensity, actual diving or somersaulting through the water. Sleep often follows such maintenance activities, the bill being turned and placed under the scapular (shoulder) feathers. Bathing is often a communal activity, and mutual preening is seen in several species.

      Social (Social behaviour in animals) displays (display behaviour) are one of the most interesting aspects of waterfowl behaviour, and many of the signal movements involved in the displays are clearly derived from maintenance activities. Thus, preening dorsally, on the breast, and especially behind the wing can be seen in ritualized form in social situations. Likewise, the wing stretch and the general body shake occur in threat or sexual displays.

      Most waterfowl are gregarious and have well-developed social integration (animal communication) signals. Many indicate the intention to fly by head-shaking or chin-lifting, so that the group takes off together. In swans and geese vocalization plays a part in this situation and also in group flight. Formal aggressive displays are important in gregarious species to prevent actual fighting and injury and instead to establish a stable dominance order. In threat, the behaviour often tends to increase the apparent body size, as when swans raise their wings and ruffle the scapular feathers. Wing flaps and flicking of the folded wings are common in geese. The chin may be lifted during aggressive display, but more frequently the head is thrust forward, often with the bill open. The striking nature of these gestures is often emphasized by patches of colour, which may be concealed before the movement is made. Since these gestures may occur without associated feather patterns, this suggests that the behaviour patterns evolved first.

      Pair-forming displays are well developed and characteristic of each species. This is necessary if mating with closely related and coexisting species is to be avoided. Swans and geese cement the pair bond by a “triumph ceremony,” with mutual head waving and calling, typically when the male has driven off an intruder. Male sheldgeese have a puffing, strutting display. Their females incite them to attack other birds by sideways jabbing movements of the bill. Female incitement behaviour is found throughout the rest of the family, but the male pairing behaviour is more varied. It is particularly striking in the dabbling ducks (dabbling duck), such as the mallard, teal, and pintail, where it is often social, a group of males displaying around a solitary female, who appears to do the mate selection. Ethologists have endeavoured to give objective names to the various postures and sequences. Thus “head-up-tail-up” involves a simultaneous upward jerk of head and tail and a lifting of the folded wings to display the speculum, a set of metallic-coloured secondary flight feathers of the upper wing. The “grunt-whistle” involves throwing an arc of water at the female by a sideways flick of the bill, followed by a rearing up of the body, shaking of the head and tail, and, during the whole sequence, giving the call indicated by the behavioral term. The mergansers (merganser) also have elaborate male displays: bowing while producing dovelike coos; flagging the head from side to side; jerking the head back on the tail and kicking up a spurt of water (the “head-throw-kick” of the goldeneye, Bucephala clangula). Perhaps the most bizarre display is found in the stifftails (stifftail), such as the ruddy duck (Oxyura jamaicensis) of North America. The males cock their tails over their backs, inflate their lower necks, and beat on them with their bright blue bills, producing a chittering sound that terminates in a burp.

      Displays that maintain the pair bond are less varied and tend to be more marked in the long-lived geese and swans, which pair for life, than in the short-lived ducks. Mutual preening and drinking displays are of this category, as are precopulatory displays, often ritualized feeding movements such as mutual head dipping, bill dipping, or head pumping. Following copulation, other mutual displays occur. The geese and swans indulge in calling with upstretched necks and lifted wings. More extraordinary is the “step-dance” of the whistling ducks (whistling duck), in which the partners rear up side by side, treading water and each raising the outside wing.

      The display patterns, whatever their origin, appear to be ready-formed and complete. Although much waterfowl behaviour can thus be said to be innate, other activities are learned during the life of the individual. One special type of learning, which is called imprinting, has been much studied in this group. The newly hatched, downy young exhibit a strong but unspecific following response to a large moving object, especially if it emits rhythmical sounds. While following such an object, they become “imprinted” with the knowledge of its characteristics and thereafter treat it as the parent, even though it may be something wildly unsuitable—a balloon, for instance. The class of object is not necessarily treated as the sexual partner when the duck matures, but it does appear that male dabbling ducks learn the plumage patterns of the female from the ducks with which they are reared. Females, however, apparently have the ability to ignore such early experience and to select the appropriate male when they mature.

      Many other aspects of waterfowl behaviour show a similar interweaving of innate and learned responses. Thus the young bird hatches with distinct preferences for the colour, size, and shape of objects at which it will peck. Within these limits, and those imposed by its bill structure and the habitat to which it is taken by its parent, the young bird can be quite adventurous in its feeding, gradually excluding unpalatable or unsuitable objects.

      Three main lines of feeding behaviour have evolved in the waterfowl—diving, dabbling, and grazing. Those that dive for food fall into two groups: inland species ( pochards and the scaup) that favour relatively shallow lakes up to 6 metres (20 feet) deep and feed predominantly on plants such as pondweeds, and mergansers that feed in deeper marine waters on invertebrates and fish. Dabbling ducks feed on the surface or by upending themselves to reach food on the bottom, largely plant material. The grazing geese and swans take grasses and sedges on dry land or in marshes, where they seek roots both on land and under the surface. Many geese have adapted to various farm crops, with new feeding traditions spreading rapidly as the birds of a group learn from one another.

      Migratory behaviour is likewise partly innate, partly learned. Young ducks of migratory populations set off in the autumn in approximately the right direction, even if there are no old birds to guide them. Others show a fixed orientation after capture and release that has no apparent function and so has been termed “nonsense” orientation. In both cases the direction is innate, and so is the ability to determine it with reference to the Sun's position (making allowance for its diurnal movement) or to the patterns of the stars. It appears, however, that the details of the Sun movement and star patterns have to be learned, as do the characteristics of the home from which the bird starts and the wintering site at which it ends. Like other migratory birds, waterfowl have the ability to return to the nesting area when they are displaced from it, but it is not clear how far they can be truly said to navigate—that is, to pinpoint their position in two coordinates, again perhaps with reference to the Sun or stars. In geese and swans, the young travel with their parents, so the possibility exists of their learning the chains of lakes, rivers, and other landmarks over which they fly and “map reading” on subsequent migrations. Certainly the migratory behaviour is highly modifiable in these longer-lived birds. New wintering resorts are adopted if their food supply or lack of disturbance makes them favourable. The migratory urge can be overcome completely in geese that are reared artificially. If the climate is not too severe and food and mates are available, such birds will remain where they grow up, even though they are free to come and go.

      One form of waterfowl behaviour that seems to be largely innate is sound production (birdsong). Vocalizations are rather simple and apparently do not offer the opportunity for individual elaboration as in songbirds. A range of honks, whistles, grunts, coos, and quacks are produced, mainly in the context of social contact and flock cohesion or in courtship display. The voices of the sexes are generally quite dissimilar. Some nonvocal sounds are also produced by the wings of some species and by the inflatable air sacs of others.

locomotion

      Only the screamers (screamer), with their broad wings and light bodies, are able freely to indulge in soaring flight. The magpie goose (Anseranas semipalmata) has somewhat similar wings, but typically the waterfowl have strong, rapid flight, their heavy body and relatively small wings giving a very high wing loading. The trumpeter swan (Cygnus buccinator) is probably the heaviest flying bird, sometimes weighing more than 17 kg (37 pounds). Trumpeter swans beat their wings about three times per second, the smaller ducks twice as fast. Geese tend to fly in long extended lines, often adopting a V-formation. Cruising air speeds of most waterfowl appear to be about 65 km (40 miles) per hour; when pressed they can certainly reach 100 km (62 miles) per hour. On migration most flocks fly at altitudes of between 300 and 600 metres (1,000 and 2,000 feet). Uncommonly they may be seen around 3,000 metres (10,000 feet), and the barheaded goose (Anser indicus), breeding in Tibet and wintering in India, must fly at 6,000 metres (20,000 feet) to get through the Himalayan passes.

      Swans and the heavier geese require a running start into the wind when taking off from water or land. Pochards (pochard) and mergansers also skitter along the surface before becoming airborne. Three species of steamer ducks (steamer duck) (genus Tachyeres) have lost the power of flight and, even when in a hurry, can only churn along the surface like paddle steamers. Small geese and most ducks can launch themselves directly into the air, pushing with the legs and giving a tremendous thrust with the wings, which may strike the water. In landing, the wings and spread feet are used for braking and guidance. Altitude is often lost rapidly by alternate sideslips, a process carried to extremes in the downward “whiffling” of geese, in which the birds may actually roll over on their backs, still keeping the head straight and level. Many ducks are quite maneuverable near the ground, back-flapping their way down vertically for the last few feet. The pochards land faster and normally only on water.

      Walking on land is well-developed in the longer-legged geese and in gooselike species. The “goose-step,” with exaggeratedly lifted feet, is exemplified by the spur-winged goose (Plectropterus gambensis). Others walk more straightforwardly and can outrun a pursuing human. In the ducks, whose short legs are situated rearward and farther apart, the gait is at best a waddle. The legs of most pochards and mergansers are so far back that the body is carried almost upright when out of the water. This development reaches its extreme in the stifftails (stifftail), which rarely leave the water on foot.

 The magpie goose, with long, nearly web-free front toes and a long rear toe, is able to perch on treetop twigs, but this is very much an exception. A number of other ducks, especially the hole-nesting perching ducks (perching duck) such as the wood duck and the mandarin duck (Aix sponsa and A. galericulata), can perch on branches or scramble up rough surfaces.

      Most waterfowl are especially adapted for swimming, with their waterproof plumage, fat-insulated body, and powerful legs with webbed feet. The feet paddle alternately in slow swimming, but the whole leg is used when the bird is moving fast. All waterfowl are able to dive if pressed, and about 40 percent use diving as their normal feeding procedure. They submerge by arching the body and propelling themselves forward with both feet so that entry is in a smooth arc. The whistling ducks, however, take a jumping “header,” in which they clear the water like coots (coot). Stifftails are consummate divers, simply slipping underwater. Diving in fast-running streams requires both great power and adaptations, as seen in the streamlined form and the long, stiff tail of the torrent ducks (torrent duck) (genus Merganetta). Once a diving duck is underwater, the legs are sculled together. In some species the wings are opened and used as steering rather than as propelling devices.

      Long-tailed, or old squaw, ducks (Clangula hyemalis) have been caught in fishing nets more than 50 metres (160 feet) deep, but this is exceptional; most species do not dive much below 6 metres (20 feet). They normally remain below for less than 30 seconds, occasionally up to 90 seconds, but they are physiologically capable of much longer dives.

Reproductive behaviour

      Copulation in waterfowl typically occurs in the water, with the pair in isolation. Indeed, the avoidance of disturbance at this time may be a major function of territorial behaviour. Colonial nesting is not normal, but high densities may occur when certain areas are unusually safe and suitable—islands, for instance. More than 200 gadwall (Anas strepera) nests per acre (494 per hectare) have been reported in such circumstances. More usually the nests are scattered, the distance between them being determined in part by the strength of the territorial behaviour. Some swans, shelducks, and sheldgeese are highly aggressive and defend a territory around the nest. In most species, however, the situation is less clear-cut. The pairs of ducks remain within a “home range,” incorporating water, a loafing spot, nesting cover, and food, but they do not vigorously defend its borders. Rather, the territory proper seems to be an area about the female, the territory moving as she does. A male intruding on a pair gives rise to the “three-bird chase,” which is a characteristic of prenesting behaviour. Geese likewise defend an area around the female as well as the immediate vicinity of the nest.

      Nest site selection is done mainly by the female, although the male may accompany her in her search. Nests built over water among marsh vegetation are found in the primitive species, and the protection thus afforded from predators may have tipped evolution toward a more wholly aquatic existence. Many species nest on dry land, with some ducks settling up to 1.6 km (1 mile) from water. Some species, especially the shelducks (shelduck), use crevices and holes for nest sites. They do not construct their own but take over what is available, such as rabbit burrows. Most hole nesters, however, utilize preexisting treeholes and will quite readily adopt nesting boxes; old tree nests of other species are sometimes used. A successful species such as the mallard is adaptable in its choice of nest site and may be found in any of the above situations.

      Nest building is generally done by the female, though the male assists in some species. It is a general rule that waterfowl are behaviorally incapable of carrying material in the bill, and even in the rare instance of courtship feeding shown by the red-crested pochard (Netta rufina), the female has to swim to the male to take his “gift.” In most waterfowl, therefore, the nest is constructed of materials on the site within reach of the beak. They are pulled in and dropped sideways over the shoulder, forming a pile as the bird rotates. With the exception of species that share incubation, such as the magpie goose, the whistling ducks, and the white-backed duck (Thalassornis leuconotus), the nest is progressively lined with down preened from the female's breast. This down is pulled over the eggs when the female leaves, keeping them warm and, in open sites, protected from the sight of predators.

      Egg laying usually occurs early in the morning and takes only a few minutes, though the female remains on the nest for up to an hour. Generally one egg per day is laid, with occasional gaps. The greatest spread recorded is in the velvet or white-winged scoter (Melanitta fusca), which takes 15 days to lay its usual clutch of nine eggs. Sperm remains viable in a female mallard for a similar period, so it is probable that copulation prior to egg laying is sufficient to fertilize the whole clutch. Clutch size is variable within a species, but averages range from 3 to 12 eggs. There do not appear to be definite relations between clutch size and taxonomic, ecological, or geographical groupings, and its significance in anseriforms is still controversial. There is, however, a tendency for larger clutches to be laid by species that devote little parental care to their offspring. Egg size varies from 3 to 15 percent of the body weight in different species, the relatively largest being produced by the stifftails. In all cases the eggshells are immaculate white, green, or brownish. Surface finish ranges from very smooth to dull and large-pored.

      The laying of eggs in nests other than their own is widespread among waterfowl, and very large clutches (up to 22 in the mallard) are usually the work of more than one female. In the pochards and stifftails such nest- parasitism is so common that it influences breeding biology. Only one species, the black-headed duck (Heteronetta atricapilla) of South America, is an obligate nest-parasite, always laying in the nests of other species.

      With few exceptions, only the female incubates, the average period ranging from 22 to 39 days. Usually she leaves the nest periodically to feed, covering the eggs with down. In some species, such as the common eider (Somateria mollissima), she apparently fasts during the four-week period. Her resulting constipation may be the cause of the extremely evil-smelling feces that she splashes over the clutch if suddenly flushed from it. This was once thought to deter predators, but experiments have not confirmed this. As incubation proceeds, aided by both the high temperature (about 38 °C [101 °F]) and high humidity maintained in the nest, the female becomes increasingly reluctant to leave the nest when a predator approaches. If flushed late in incubation or with hatched young, she will feign injury, flapping or dragging her wings in an uncoordinated fashion that serves to distract predators.

 The length of the incubation period shows little relation to taxonomic position or body size, but both it and the subsequent fledging period are correlated with the latitude at which breeding takes place. The continuous daylight of the Arctic summer enables the young to feed around the clock, but this does not sufficiently offset the fact that there is only a short interval between the melting of the snows and the onset of autumnal bad weather. Selection in the course of evolution has reduced the breeding sequence as far as possible. Thus the small high-Arctic races of the Canada goose (Branta canadensis) have so reduced the period from nest initiation to fledging that they can breed in areas having but two snow-free months. Larger races nesting in California take twice as long. In the wild, waterfowl do not breed more than once per year, even when there is enough time. Most will renest, however, if the first clutch is destroyed. The second clutch is generally smaller but may have a higher fledging success in that hatching occurs in the less uncertain weather of late summer. Aviculturists regularly take advantage of this capacity, incubating the first clutch under bantam hens.

      Since incubation does not start until the clutch is complete, hatching is generally synchronized and may be complete within six hours. Another six hours is required for the down to dry, and young birds usually remain in the nest overnight. While in the nest they are brooded by the female and become imprinted (imprinting) with her visual and auditory characteristics. The downy young have sufficient internal food reserves to last two or three days if inclement weather prevents them from feeding as soon as they leave the nest.

      The nesting behaviour of screamers (screamer) is somewhat different from that of waterfowl. A substantial nest is built on marshy ground or in shallow water, sometimes reaching a height of 30 cm (1 foot) or more. The clutch varies from two to six eggs, depending on the species. Young screamers are covered with yellow down and leave the nest when a few days old, following their parents in the manner of young waterfowl.

      Knowledge of waterfowl breeding, although far from complete, has come a long way from medieval times when barnacle geese (barnacle goose) (Branta leucopsis) were thought to hatch from barnacles on rocks because no one had yet discovered their high-Arctic nests.

Ecology

      Most waterfowl are associated with fresh water as opposed to salt water, at least during the breeding season. Relatively few inhabit deep lakes and fast streams of low productivity. More prefer glaciated areas rich in shallow, productive lakes and marshes: the “pothole” country of North America, the silted deltas of rivers, and coastal marshes. An area with many small bodies of water will support more ducks than a single large lake, because shoreline is important. Such environments produce diverse food and provide isolation for breeding pairs, cover from predators, and shelter from winds. Paradoxically, periodic droughts (drought) are advantageous to ducks. Small lakes dry out and their bottoms become colonized with plants. When reflooding occurs, a rich diet of seeds and of invertebrate animals that feed on the decaying vegetation becomes available. Long-term or permanent droughts, however, can be disastrous. The increasing acreage of artificial water reservoirs only partly offsets the loss, for they are normally large, deep waters, of little use to waterfowl except for roosting.

      The ability to migrate (migration) greatly increases the ecological flexibility of waterfowl. They can exploit summer resources of the northern tundra without having to evolve the encumbering adaptations needed to survive the winter there; they can move great distances to take advantage of local rainfalls in arid countries; they can retire to secluded areas when flightless and then seek the rich harvests of human-altered environments when fully equipped to face the dangers. There are many patterns of migration, some regular and in fixed, reciprocal directions, others varying with changes in weather. The distances covered also vary widely. The longest waterfowl migrations are probably those of the blue-winged teal (Anas discors), which nests up to 60° N in North America and winters beyond 30° S, a distance of over 9,600 km (6,000 miles). In the Old World the northern shoveler (Anas clypeata) has a similar distance of up to about 11,000 km (6,800 miles). The northern pintail and the garganey (Anas querquedula), which breed in the Siberian tundra and taiga and winter in the tropical swamps of Senegal and Chad, are even more remarkable in their adaptability.

      The general timing of regular long-distance migrations has been built into an internal hormonal cycle over the course of evolution. These “annual clocks” are kept in synchrony by changes in day length, but the immediate factors initiating the stages of migration are climatic. A mass southward movement occurs when air pressure systems produce a favourable wind flow; overcast conditions or a warm spell may temporarily halt the birds; bright, cold conditions move them on.

      Sudden adverse weather (climate) may cause considerable mortality, especially where a long sea crossing is involved, as from Greenland to Britain. But the effects of weather on waterfowl populations can be most drastic at the time of breeding. Late frosts and spring floods are catastrophic, especially when there is no time for renesting. Consequently, high-Arctic breeders may have near-complete reproductive failures. Brant geese (brant) (Branta bernicla) that winter in western Europe only produce successful broods of young in about half the seasons.

      Parasites of many kinds inhabit waterfowl, generally in a state of tolerance, gaining the upper hand only when the bird is stressed in some way, as by food shortage. This is also true for fungal infections, particularly by Aspergillus, a common cause of death in captivity and, exceptionally, in the wild. Bacterial diseases take their toll and sometimes cause wholesale die-offs. Thousands of ducks have succumbed to botulism when shallow, brackish waters dry out in warm countries and the bacteria responsible multiply, producing toxins that cause a fatal paralysis if ingested. Fowl cholera (Pasteurella) sometimes causes epidemics. Mass mortality is also triggered by pollution, especially oil and pesticide spills. A peculiarly insidious danger is lead poisoning caused by the ingestion of lead pellets that accumulate in the mud of waters frequently shot over.

      Humans are undoubtedly a major predator of waterfowl. Adult ducks, other than nesting females, suffer relatively little natural predation unless they are sick or weakened by hunger. The main predatory impact comes on the eggs and young. Crows and gulls, mink, raccoons, coyotes, foxes, ground squirrels, snakes, snapping turtles, bullfrogs, pike, and carp all take some toll. In normal circumstances such predation does not destabilize population dynamics, which are, after all, geared to a high rate of loss in the early stages. Predator-control measures have seldom produced any comparable increase in waterfowl production.

      Beneficial relations exist with other birds and animals. Thus stifftails (stifftail) and eiders (eider) actually prefer to nest in association with the smaller gulls (Larus species). The latter tend to drive off predators and, by increasing the number of available eggs, relieve the pressure of egg predators. The muskrat clears water routes in reed beds and builds “houses” that are utilized by ducks for standing and nesting. Beaver dams create very attractive duck habitats. And, of course, man himself is not wholly destructive but creates some new areas for waterfowl and provides them with food, both unintentionally and on purpose.

Form and function

Anatomy

      The characteristic features of the family Anatidae are the skin-covered, lamellate bill (beak) and thick, fleshy tongue. Screamers (screamer) have a short nonlamellate bill, slightly hooked, like that of a domestic hen, and a horny tongue. Their bill is adapted for tearing aquatic plants. Among the waterfowl the basic bill has undergone a wide adaptive radiation. The geese have evolved strong, deep bills with hard, sharp lamellae. In some, such as the red-breasted goose (Branta ruficollis), the bill is short and slight, used only for grazing; in others, such as the snow goose (Anser caerulescens), it is long and heavy enough to dig for roots and tubers. The massive digging bill reaches maximum development in the magpie goose. The little pygmy geese (Nettapus species) are so called for their gooselike bills, but they actually feed on lotus seeds and water vegetation and neither graze nor root for food. The European widgeon (Anas penelope), on the other hand, grazes extensively, but its bill differs little from the typical duck bill—flattened with platelike lamellae. This is used for sifting particles out of mud or picking up food items from the lake bottom as the bird upends itself. The sieving bill is yet further developed in the shovelers and the pink-eared duck (Malacorhynchus membranaceus), the lamellae becoming extremely fine, enabling particles as small as diatoms (diatom) to be taken from the surface film. The blue duck (Hymenolaimus malacorhynchos) has a rounded, expanded tip to the bill, which probably protects it when poking around sharp pebbles. The pochards have fewer lamellae and a narrower bill than the dabbling ducks (dabbling duck). In the mergansers (merganser) the lamellae have become toothlike projections in the long narrow bill, ideal for holding fish; additionally, the tongue has two dorsal rows of barbs. None has developed pickax bills or ones for crushing hard food items, such as mollusks. When such foods are eaten, they are broken up in the gizzard.

      The legs and feet (foot) vary according to whether the main mode of progression is walking (as in geese) or swimming (as in divers). The latter have flattened foot bones to reduce water resistance on the forward stroke, the hind toe being lobed. Webs are fully developed between the front three digits, except in the magpie goose (primitively) and the Hawaiian goose, or nene, where it is believed that the greater flexibility of the toes makes for easier walking on broken lava beds. The claws on the digits are particularly sharp in forms such as the perching ducks, which normally nest in treeholes.

      The wings (wing) are not of unusual structure but vary widely in shape, from the broad expanse of the slow and maneuverable fliers to the narrow sweeps of the fastest. Some of the whistling ducks have modified vanes on the outer primaries that produce a whirring noise in flight. The sighing creak of the wings of the mute swan (Cygnus olor) is well known. In a few cases the wing is so reduced in size that the bird is flightless, as is the Auckland Island (Auckland Islands) race of the brown teal (Anas aucklandica). Many waterfowl have horny knobs on the wrist joint located near the end of the wing; in some there is a distinctly projecting spur, as in the screamers, the spur-winged goose, and the torrent duck. In the first two, the spurs are actually used to strike adversaries; in the last, they may be of more use in progressing over slippery rocks in a raging stream.

      A structure that has received much attention, largely because of its taxonomic value, is the trachea. This may be straight (geese and ducks) or looped in various ways (magpie geese and swans), the most elaborate being that of the trumpeter swan (Cygnus buccinator). Here it first enters the sternum, flexing twice into horizontal and vertical bony pockets, then emerging again to coil around and back into the lungs. The analogy to an orchestral wind instrument is obvious, and a comparable sound is produced. Less obvious is the relation of shape to sound in the asymmetrical chamber (bulla) found at the base of the male trachea in most ducks. The male's whistle certainly differs from the female's quack, but it is difficult to account for the variation in bulla shapes—round and solidly ossified (dabbling ducks), angular with membranous “windows” (stifftails), or complexly membranous (mergansers).

      The anseriform birds share with the curassows, tinamous, and ratites (ostriches, emus, rheas, and the like) the possession of an intromittent male organ. This is not homologous with the mammalian penis but is instead a vascularized sac everted from the wall of the cloaca. It is protruded by muscular action and retracted by an elastic ligament. This gives it a spiral form, and sperm pass along a groove to the tip. This organ doubtless facilitates copulation on the water. Its presence permits easy sexing of birds that lack differences in plumage between the sexes. The size of the organ also allows an estimate of the bird's age. However, that is more safely done by determining the depth of the bursa of Fabricius, a glandular organ opening into the cloaca that diminishes with maturity.

Physiology

      There are interesting adaptations in the respiratory physiology of diving ducks (diving duck). The accumulation of carbon dioxide does not stimulate breathing as it does in land mammals. When the bird is under water, oxygen is drawn from stored hemoglobin and myoglobin in blood and muscles, instead of from the respiratory system. The air sacs are compressed, reducing buoyancy. glycogen is broken down by anaerobic and aerobic means, and the blood flow is modified so that the central nervous system receives the main supply.

      Rather little is known about the physiology of digestion in waterfowl. It appears that, despite being almost entirely grazers, geese lack the ability to digest cellulose, either by the secretion of enzymes or by the symbiotic activities of microorganisms in the gut. Such inefficient digestion is probably correlated with the extraordinarily rapid passage of ingested material, which begins to appear in the droppings within half an hour and can be completely eliminated in two hours.

      Salt excretion is achieved by paired nasal glands (nasal gland) situated on indentations of the skull over the eyes. Populations living in marine or brackish environments have enlarged glands, considerably modifying the profile of the head. In a botulism infection, it is probable that the toxins cause these glands to malfunction, resulting in the characteristic salt imbalance and paralysis.

evolution and paleontology

      Anseriforms represent an ancient lineage. Fossil and genetic evidence suggest that their ancestors were one of the earliest lineages in the evolution of modern birds, going as far back as the Cretaceous Period more than 65 million years ago. The rich fossil record of modern waterfowl starts in the Late Eocene Epoch (37 to 33.7 million years ago). Diversification of this order probably started in the Southern Hemisphere, where old and apparently primitive lineages such as the magpie goose and screamers (screamer) as well as many endemic and monotypic genera still reside.

      Ancestral forms were probably long-legged unspecialized birds with features similar to the galliform curassows (curassow) and to screamers and magpie geese. While the screamers have probably not advanced much, the Anatidae have diversified rapidly and extensively. Two main lines may be distinguished. One, typified by the swans and geese, became large-bodied, long-lived, slow-breeding birds, with prolonged pair and family bonds and associated parental care of the young. The other line, characterized by most of the ducks, has favoured small-bodied, short-lived birds having brief pair bonds and minimal care of the young, necessarily correlated with prolific production of young. Superimposed on these general trends were adaptive radiations from omnivorous to specialized food habits and from a largely terrestrial to an almost fully aquatic existence. An adaptation for the latter, the double molt, enabled males to develop striking nuptial plumage, which prevented interspecific hybridization where many species coexisted. Migratory habits strengthened this requirement, and gene flow within such species was rapid. Racial differences developed only in nonmigratory or otherwise isolated species, or in those migratory forms that maintained family groupings when away from the breeding grounds, and so developed learned, traditional attachments to the area where they were born.

Classification

Distinguishing taxonomic features

      Broad classification divisions are based on skeletal and other internal structures. Finer divisions are decided on plumage type and patterns, including those of the downy young, and on behavioral features such as displays associated with pair formation and maintenance.

Annotated classification

      There is general agreement on the family and subfamily divisions of the order and at the generic level, where most workers agree on the lumping of species into large genera. The study, in recent years, of new taxonomic characters such as behaviour patterns and biochemistry has substantially improved our understanding of relationships within the Anseriformes.

Geoffrey Vernon Townsend Matthews Frank Gill Order Anseriformes

 165 species in 2 families of 49 genera.

      Family Anatidae (waterfowl)

 147 species in 41 genera belonging to 7 subfamilies, found virtually worldwide. Skin-covered bill, lamellate edges, tipped with horny “nail”; occipital fontanelles generally present; tongue thick, bordered with spiny processes; primary wing feathers generally molted simultaneously; male copulatory organ present; body length about 30–150 cm (12–59 inches), weight 300 grams–18 kg (10.6 ounces–40 pounds).

      Subfamily Anatinae ( dabbling ducks, perching ducks, and atypical geese)

 69 species in 16 genera including pintails, pochards, scaups, shovelers, teals, wigeons, wood ducks, and the canvasback, gadwall, mallard, and redhead. Sexes differ in voice and also usually in plumage and behaviour; brief pair bond and family life. Double annual molt; males often alternate a cryptic “eclipse” plumage with a conspicuous nuptial one; downy young strongly patterned. Female “inciting” display present. One recently extinct species.

      Subfamily Anserinae ( swans and typical geese (goose))

 24 species in 6 genera with breeding distribution primarily in the Northern Hemisphere. Legs short to medium; neck medium to long. Sexes similar in plumage, voice, and behaviour, includes “triumph” ceremony and precopulatory head dipping. Long pair bond and extended family life; young mature in second to fourth year. Single annual molt. Vegetarian. Wings usually moderately long, pointed; tail short. All species swim capably, many primarily aquatic. Simple trachea and syrinx.

      Subfamily Tadorninae (sheldgeese (sheldgoose), shelducks, and steamer ducks)

 18 species in 6 genera, worldwide except North and Central America. Iridescent green wing speculum and, generally, white wing coverts; sexes alike in majority; mature in second year. Pugnacious. Downy young black and white. One recently extinct species.

      Subfamily Merginae (sea ducks)

 18 species in 6 genera including eiders, goldeneyes, mergansers, scoters, and the bufflehead, goosander, and smew. Typically of the Northern Hemisphere; 1 species in South America. Mostly saltwater divers, feeding on animals. Sexual dimorphism in plumage, voice and behaviour; male courtship patterns very varied; maturity requires at least two years. Two recently extinct species.

      Subfamily Dendrocygninae (whistling ducks (whistling duck))

 9 species in 2 genera widely distributed in the tropics and subtropics. Whistling voice. Relatively long-legged, with upright stance. Enclosed bony eye socket. Strongly patterned adult plumage; downy young with light stripe below eyes and around nape. Mature at one year. Mutual postcopulatory step-dance.

      Subfamily Oxyurinae (stiff-tailed duck (stifftail)s)

 8 species in 3 genera including ruddy duck and musk duck. Mainly of the Southern Hemisphere. Freshwater divers with legs so far to rear that walking is difficult; sound production by inflatable sacs in neck; long, stiff tail feathers; elaborate male displays; large eggs and precocial young.

      Subfamily Anseranatinae ( magpie goose)

 1 species of northern Australia and New Guinea. Features shared with both the screamers and the swans and geese.

      Family Anhimidae (screamer) ( screamers)

 3 species in 2 genera of South America. Short, curved, nonlamellate bill; feet unwebbed; plumage uniformly distributed; extensive air cells between skin and body.

Additional Reading

Kurt M. Bauer and Urs N. Glutz von Blotzheim, Handbuch der Vögel Mitteleuropas, vol. 2 and 3, Anseriformes (1968–69), provides much wider geographic coverage than the name implies and is a complete sourcebook on the biology of Holarctic waterfowl. Classic and still useful works include J.G. Millais, The Natural History of the British Surface-feeding Ducks (1902), and British Diving Ducks, 2 vol. (1913); and John C. Phillips, A Natural History of the Ducks, 4 vol. (1922–26, reprinted 1986), covering many aspects of waterfowl biology. H. Albert Hochbaum, Travels and Traditions of Waterfowl (1955, reissued 1967), reviews waterfowl migration and orientation. Paul A. Johnsgard, Handbook of Waterfowl Behavior (1965), inventories behaviour patterns, especially social and sexual, and his Waterfowl: Their Biology and Natural History (1968), contains photographs, many coloured, of nearly every species. Joseph P. Linduska (ed.), Waterfowl Tomorrow (1964), contains authoritative articles on all aspects of waterfowl conservation, management, and research. Worldwide coverage is provided in Jean Delacour, The Waterfowl of the World, 4 vol. (1954–64, reissued 1973), a treatment of all species of Anatidae; Peter Scott et al., The Swans (1972), with coverage of all 8 species; Johnsgard's Ducks, Geese, and Swans of the World (1978); Frank S. Todd, Waterfowl (1979); Peter Scott, A Coloured Key to the Wildfowl of the World, rev. ed. (1988), containing coloured plates of all species of the Anatidae, with details of distribution and aids to identification; and Steve Madge and Hilary Burn, Wildfowl (1988). Regional studies include G.L. Atkinson-Willes and D.G. Salmon, Wildfowl in Great Britain, 2nd ed. (1986), on Anatidae distribution and populations; H.J. Frith, Waterfowl in Australia (1967), devoted to the waterfowl of the Southern Hemisphere; Hochbaum's The Canvasback on a Prairie Marsh, 2nd ed. (1959, reissued 1981), a classic species monograph; Frank C. Bellrose et al., Ducks, Geese & Swans of North America, 3rd ed. (1980), a thorough treatment, with excellent illustrations; and John Gooders and Trevor Boyer, Ducks of North America and the Northern Hemisphere (1986).Geoffrey Vernon Townsend Matthews Ed.

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