parrot

parrot

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Also known as: Psittacidae

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parrot

Category: Animals & NatureRelated Topics: macaw Strigopinae Loriinae Nestorinae pygmy parrot

parrot, term applied to a large group of gaudy, raucous birds of the family Psittacidae. Parrot also is used in reference to any member of a larger bird group, order Psittaciformes, which includes cockatoos (family Cacatuidae) as well. Parrots have been kept as cage birds since ancient times, and they have always been popular because they are amusing, intelligent, and often affectionate. Several are astonishingly imitative of many sounds, including human speech.

Budgerigar (Melopsittacus undulatus), a popular parakeet. — Budgerigar (*Melopsittacus undulatus*), a popular parakeet.

The family Psittacidae numbers 333 species. The subfamily Psittacinae, the “true” parrots, is by far the largest subfamily, with members found in warm regions worldwide. These birds have a blunt tongue and eat seeds, buds, and some fruits and insects. Many members of the subfamily are known simply as parrots, but various subgroups have more specific names such as macaw, parakeet, conure, and lovebird.Britannica QuizWild Words from the Animal Kingdom Vocabulary Quiz

The African gray parrot (Psittacus erithacus) is unsurpassed as a talker; the male can precisely echo human speech. Captive birds are alert and, compared with other parrots, relatively good-tempered. Some are said to have lived 80 years. The bird is about 33 cm (13 inches) long and is light gray except for its squared, red tail and bare, whitish face; the sexes look alike. Gray parrots are common in the rainforest, where they eat fruits and seeds; they damage crops but are important propagators of the oil palm.

Among other proficient mimics are the Amazon parrots (Amazona). The 31 species of Amazons are chunky birds, mostly 25 to 40 cm (10 to 16 inches) long, with slightly erectile crown feathers and a rather short, squared tail. Their predominantly green plumage is marked with other bright colours, chiefly on the upper head; the sexes look alike. Amazon parrots live in tropical forests of the West Indies and Mexico to northern South America. They are difficult to breed and may be aggressive as well as squawky. Common in aviaries is the blue-fronted Amazon (A. aestiva) of Brazil; it has a blue forehead, a yellow or blue crown, a yellow face, and red shoulders. The yellow-crowned parrot (A. ochrocephala) of Mexico, Central America, and from Ecuador to Brazil has some yellow on the head and neck, a red wing patch, and a yellow tail tip.

Black-capped parakeet (Pyrrhura rupicola) — Black-capped parakeet (*Pyrrhura rupicola*)

The monk, or green, parakeet (Myiopsitta monachus) is one of the hardiest parrot species. It is native to South America, but some have escaped from captivity in the United States and now nest in several states. Its large stick nest is unique among psittaciforms. Other remarkable parrots of this subfamily include the hanging parrots (Loriculus), which sleep upside-down like bats. Caiques (Pionites) are small, short-tailed South American birds similar to conures in build and habits.

For decades the night parrot, or night parakeet (Geopsittacus occidentalis), of Australia was thought to be extinct, until a dead one was found in 1990. It feeds at night on spinifex grass seeds and dozes under a tussock by day. Its nest is a twig platform in a bush and is entered by way of a tunnel. Equally unusual is the ground parrot, or ground parakeet (Pezoporus wallicus). Rare local populations exist in the wastelands of coastal southern Australia and western Tasmania. It runs in the grass, flushes like a quail, and makes a sudden deceptive pitch, and it was formerly hunted with dogs. It eats seeds and insects; its nest is a leaf-lined depression under a bush.

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Rainbow lorikeet (Trichoglossus haematodus). — Rainbow lorikeet (*Trichoglossus haematodus*).

The lories (with short tails) and lorikeets (with longer, pointed tails) make up the Psittacidae subfamily Loriinae. The 53 species in 12 genera are found in Australia, New Guinea, and some Pacific islands. All have a slender, wavy-edged beak and a brush-tipped tongue for extracting nectar from flowers and juices from fruits.

The pygmy parrots of the subfamily Micropsittinae all belong to the genus Micropsitta. The six species are endemic to New Guinea and nearby islands. These are the smallest members of the family. They live in forests, where they eat insects and fungi.

The subfamily Nestorinae is found only in New Zealand. The kea (Nestor notabilis) occasionally tears into sheep carcasses (rarely, weakened sheep) to get at the fat around the kidneys. The kaka, N. meridionalis, a gentler forest bird, is often kept as a pet. The owl parrot, or kakapo (Strigops habroptilus), also lives only in New Zealand. It is the sole member of the subfamily Strigopinae. Rare and once thought extinct, it survives as a scant population on Stewart Island.

Sulfur-crested cockatoo (Cacatua galerita). — Sulfur-crested cockatoo (*Cacatua galerita*).

Flock of galahs, or roseate cockatoos (Eolophus roseicapillus). — Flock of galahs, or roseate cockatoos (*Eolophus roseicapillus*).

The cockatoo family (Cacatuidae) numbers 21 species from Australia, New Guinea, and nearby islands. The group includes the cockatiel (Nymphicus hollandicus), a smaller bird. All are crested and have heavy beaks for cracking nuts and seeds. The so-called sea parrot is unrelated to the psittaciforms (see puffin).

The Editors of Encyclopaedia BritannicaThis article was most recently revised and updated by Adam Augustyn.

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Also known as: Aves, fowl

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lesser flamingo

Category: Animals & NatureKey People: Jean Theodore Delacour Alfred Newton John James Audubon W.H. Hudson Ernst MayrRelated Topics: pigeon flightless bird songbird ornithology bird-watching

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basic body feather tracts on a generalized songbird

bird, (class Aves), any of the more than 10,400 living species unique in having feathers, the major characteristic that distinguishes them from all other animals. A more-elaborate definition would note that they are warm-blooded vertebrates more related to reptiles than to mammals and that they have a four-chambered heart (as do mammals), forelimbs modified into wings (a trait shared with bats), a hard-shelled egg, and keen vision, the major sense they rely on for information about the environment. Their sense of smell is not highly developed, and auditory range is limited. Most birds are diurnal in habit. More than 1,000 extinct species have been identified from fossil remains.

Since earliest times birds have been not only a material but also a cultural resource. Bird figures were created by prehistoric humans in the Lascaux Grotto of France and have featured prominently in the mythology and literature of societies throughout the world. Long before ornithology was practiced as a science, interest in birds and the knowledge of them found expression in conversation and stories, which then crystallized into the records of general culture. Ancient Egyptian hieroglyphs and paintings, for example, include bird figures. The Bible refers to Noah’s use of the raven and dove to bring him information about the proverbial Flood.

Various bird attributes, real or imagined, have led to their symbolic use in language as in art. Aesop’s fables abound in bird characters. The Physiologus and its descendants, the bestiaries of the Middle Ages, contain moralistic writings that use birds as symbols for conveying ideas but indicate little knowledge of the birds themselves. Supernatural beliefs about birds probably took hold as early as recognition of the fact that some birds were good to eat. Australian Aborigines, for example, drove the black-and-white flycatcher from camp, lest it overhear conversation and carry the tales to enemies. Peoples of the Pacific Islands saw frigate birds as symbols of the Sun and as carriers of omens and frequently portrayed them in their art. The raven—a common symbol of dark prophecy—was the most important creature to the Indians of the Pacific Northwest and was immortalized in Edgar Allan Poe’s poem “The Raven.” Eagles have long been symbols of power and prestige in many parts of the world, including Europe, where their representations are often seen in heraldry. Native Americans sprinkled eagle down before guests as a sign of peace and friendship, and eagle feathers were commonly used in rituals and headdresses. The resplendent quetzal—the national bird of Guatemala, which shares its name with the currency and is a popular motif in art, fabric, and jewelry—was worshipped and deified by the ancient Mayans and Aztecs. Highly symbolic birds include the phoenix, representing resurrection, and the owl, a common symbol of wisdom but also a reminder of death in Native American mythology. The bird in general has long been a common Christian symbol of the transcendent soul, and in medieval iconography a bird entangled in foliage symbolized the soul embroiled in the materialism of the secular world.

In modern times the recreational pleasures of bird-watching have grown in tandem with the rise of environmentalism. Evolving from the American and European “shoot-and-stuff” mania of the 19th century, bird-watching became a sportlike activity based on rapid identification—the rarest being the most rewarding—with the aid of binoculars and spotting scopes. The change from shooting to sighting coincided with campaigns, beginning about 1900, to halt the slaughter of wild birds for food and millinery. Bird-watching was advanced by the publication of excellent field guides and improvements in photography and sound recording. By mid-century the watcher’s enjoyable but rather unsophisticated tallying of “year lists” and “life lists” of species personally observed was being augmented, if not replaced, by interest in careful studies of bird behaviour, migration, ecology, and the like. This trend was abetted by bird banding (called ringing in the United Kingdom) and by such organizations as the British Trust for Ornithology and the National Audubon Society, which coordinate professional and amateur observations and efforts with scientific studies.Britannica QuizDeadliest Animals Quiz

General features

Birds arose as warm-blooded, arboreal, flying creatures with forelimbs adapted for flight and hind limbs for perching. This basic plan has become so modified during the course of evolution that in some forms it is difficult to recognize.

Among flying birds, the wandering albatross has the greatest wingspan, up to 3.5 metres (11.5 feet), and the trumpeter swan perhaps the greatest weight, 17 kg (37 pounds). In the largest flying birds, part of the bone is replaced by air cavities (pneumatic skeletons) because the maximum size attainable by flying birds is limited by the fact that wing area varies as the square of linear proportions, and weight or volume as the cube. During the Pleistocene Epoch (2.6 million to 11,700 years ago) lived a bird called Teratornis incredibilis. Though similar to the condors of today, it had a larger estimated wingspan of about 5 metres (16.5 feet) and was by far the largest known flying bird.

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The smallest living bird is generally acknowledged to be the bee hummingbird of Cuba, which is 6.3 cm (2.5 inches) long and weighs less than 3 grams (about 0.1 ounce). The minimum size is probably governed by another aspect of the surface-volume ratio: the relative increase, with decreasing size, in surface through which heat can be lost. The small size of some hummingbirds may be facilitated by a decrease in heat loss resulting from their becoming torpid at night.

When birds lose the power of flight, the limit on their maximum size is increased, as can be seen in the ostrich and other ratites such as the emu, cassowary, and rhea. The ostrich is the largest living bird and may stand 2.75 metres (9 feet) tall and weigh 150 kg (330 pounds). Some recently extinct birds were even larger: the largest moas of New Zealand and the elephant birds of Madagascar may have reached over 3 metres (10 feet) in height.

The ability to fly has permitted an almost unlimited diversification of birds, so that they are now found virtually everywhere on Earth, from occasional stragglers over the polar ice caps to complex communities in tropical forests. In general the number of species found breeding in a given area is directly proportional to the size of the area and the diversity of habitats available. The total number of species is also related to such factors as the position of the area with respect to migration routes and to wintering grounds of species that nest outside the area. In the United States, Texas and California have the most—approximately 620 for each (the figure varies based on criteria used for inclusion on state lists, such as unconfirmed, accidental, hypothetical, extirpated, and extinct species). More than 920 species have been recorded from North America north of Mexico. The figure for Europe west of the Ural Mountains and including most of Turkey is 514. More than 700 species live in Russia. At least 4,400 species live in North and South America. Although several South American countries boast well over 1,000 species, Costa Rica, with an area of only about 51,000 square km (about 20,000 square miles) and a known avifauna of more than 800 species, probably has the most diversity for its size. Asia accounts for more than 25 percent of the world’s species, with 2,700 species, and Africa slightly less, with about 2,300.

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Importance to man

In addition to their importance in literature and legend, birds have been significant to human society in myriad ways. Birds and their eggs have been at least incidental sources of food for humans since their origin and still are in most societies. The eggs of some colonial seabirds, such as gulls, terns, and murres, or guillemots, and the young of some muttonbirds are even now harvested in large quantities. With the development of agrarian human cultures, several species of chickens, ducks, geese, and pigeons were taken in early and have been selectively bred into many varieties. These domestic birds are descended, respectively, from the red jungle fowl (Gallus gallus), mallard duck (Anas platyrhynchos), greylag goose (Anser anser), and rock dove (Columba livia). After the discovery of the New World, the turkey (Meleagris gallopavo), which had already been domesticated by the Indians, and the Muscovy duck (Cairina moschata) were brought to Europe and produced several varieties. Guinea fowl (Numida meleagris) from Africa were also widely exported and kept not only for food but also because they are noisy when alarmed, thus warning of the approach of intruders.

Besides being a food source, pigeons have long been bred and trained for carrying messages, their wartime use dating to the Roman era, according to Pliny the Elder. Messenger pigeons were widely used by German, British, and American forces in World Wars I and II and by the United States in the Korean War. In the South Seas, the ability of frigate birds to “home” to their nesting colonies enabled island inhabitants to send messages by these birds.

With the development of modern culture, hunting evolved from a foraging activity to a sport, in which the food value of the game became secondary. Large sums are now spent annually on hunting waterfowl, quail, grouse, pheasants, doves, and other game birds. Sets of rules and conventions have been set up for hunting, and in one elaborate form of hunting, falconry, there is not only a large body of specialized information on keeping and training falcons but also a complex terminology, much of it centuries old.

Feathers have been used for decoration for many thousands of years. Their use in the headpieces of indigenous peoples throughout the world is well known. Feather robes were made by Polynesians and Eskimos; and down quilts, mattresses, and pillows are part of traditional European folk culture. Large feathers have often been used in fans, thereby providing an example of an object put to opposite uses—for cooling as well as for conserving heat. Whereas most feathers used in decorating are now saved as by-products of poultry raising or hunting, until early in the 20th century, egrets, grebes, and other birds were widely shot for their plumes alone. Ostrich farms have been established to produce plumes as well as meat, and some ostriches have been raised specifically for racing. Large quills were once widely used for writing, and feathers have long been used on arrows and fishing lures.

Many birds are kept as pets. Small finches and parrots are especially popular and easy to keep. Of these, the canary (Serinus canaria) and the budgerigar of Australia (Melopsittacus undulatus, often called a parakeet) are widely kept and have been bred for a variety of colour types. On large parks and estates, ornamental species such as peacocks (Pavo cristatus), swans, and various exotic waterfowl and pheasants are often kept. Zoological parks in many cities import birds from many lands and are a source of recreation and enjoyment for millions of people each year.

View a flock of red-billed queleas at the Etosha National ParkSee all videos for this article

With the rise of agriculture, man’s relationship with birds became more complex. Vast quantities of guano (bird excrement) were mined from island breeding colonies for use as fertilizer for crops. However, in regions where grain and fruit are grown, depredations by birds may be a serious problem. In North America various species of blackbirds (family Icteridae) are serious pests in grainfields; in Africa a grain-eating finch, the red-billed quelea (Quelea quelea), occurs like locusts, in plague proportions so numerous that alighting flocks may break the branches of trees. The use of city buildings for roosts by large flocks of starlings and blackbirds is also a problem, as is the nesting of albatrosses on airplane runways on Pacific islands. As a result of these problems, conferences on the control of avian pests are commonly held.

Although birds are subject to a great range of diseases and parasites, only a few of these are known to be capable of infecting man. Notable exceptions are ornithosis psittacosis, or parrot fever, a serious and sometimes fatal disease resembling viral pneumonia. The microorganism responsible for the disease is transmitted directly to man from pigeons, parrots, and a variety of other birds via their excrement. Encephalitis, an inflammation of the brain, is also serious, but this infection is transmitted to man and to his domestic animals via biting arthropods, including mosquitoes. West Nile virus can likewise be transmitted. Wild birds may also act as reservoirs for diseases that adversely affect domesticated birds.

Track how Galapagos finches underwent adaptive radiation from a single ancestral lineage and their contribution to Darwin's theory of evolution — Track how Galapagos finches underwent adaptive radiation from a single ancestral lineage and their contribution to Darwin’s theory of evolutionSee all videos for this article

The study of birds has contributed much to both the theoretical and practical aspects of biology. Charles Darwin’s studies of the Galapagos finches and other birds during the voyage of HMS Beagle were important in his formulation of the idea of the origin of species through natural selection. Collections of birds in research museums still provide the bases for important studies of geographic variation, speciation, and zoogeography, because birds are one of the best known of animal groups. Early work on the domestic fowl added to the development of both genetics and embryology. The study of animal behaviour (ethology) has been based to a large extent on studies of birds by Konrad Lorenz, Nikolaas Tinbergen, and their successors. Birds also have been the primary group in the study of migration and orientation and the effect of hormones on behaviour and physiology.

Man’s impact on bird populations is very strong. Since 1680 approximately 80 species of birds have become extinct, and a larger number are seriously endangered. While pollution and pesticides are important factors in the decline of certain large species, such as the peregrine falcon, osprey, and California condor, the destruction of natural areas and introduction of exotic animals and diseases have probably been the most devastating. Concerted efforts of research and conservation are required to ensure the survival of rare species.Robert W. Storer

Natural history

Locomotion

Because of their body structure and their feathered covering, birds are the best fliers among animals, better than insects and bats. There are, however, considerable differences in ability among various birds. Penguins cannot fly, instead spending much of their time in the water swimming with their paddlelike wings. Birds such as ostriches and emus have rudimentary wings but are permanently afoot. At the other extreme, long-winged swifts and frigate birds move from their perches only to fly, never to walk. Most birds alternate some walking or swimming with their flying.

Birds usually fly when they have any considerable distance to travel; there are exceptions, however. The mountain quail of California make their annual migrations up and down the mountains on foot. The guillemots of the Greenland coast migrate southward by swimming; they begin their journey before the young have grown their flight feathers and before some of the adults at least have regrown their recently molted ones. The Adélie penguins may ride northward on drifting ice floes; at the approach of nesting time, they swim back to the Antarctic continent and then walk over the ice to their breeding grounds many miles inland.

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Flight

Birds fly by flapping their wings, steering mainly with their tails. Compared to the parts of an airplane, a bird’s wing acts as both wing and propeller. The basal part of the wing supplies most of the supporting surface, the wing tip most of the propelling force. A bird’s wing has many adjustable features: it can be shortened or lengthened by flexion; the feathers of the tip can be spread or closed; the angle of the whole wing or its parts—on one side or both—can be altered. All of these adjustments make the aerodynamics of a bird’s wing much more complicated than those of the airplane; consequently, the flight of a bird is much more varied and adaptable.

Learn how a hummingbird can fly in any direction and about its iridescent plumageSee all videos for this article

The types of flight found in birds vary considerably, and different types of wings correlate with different types of flight. At least two major types of modifications for gliding or soaring are found. Albatrosses and some other seabirds have long, narrow wings and take advantage of winds over the oceans, whereas some vultures and hawks have broad wings with slotted tips that permit more use of updrafts and winds deflected by hills. Short, broad wings are characteristic of chickenlike birds, which fly up with a rush of rapid wing beats. Birds such as ducks, pigeons, and falcons, which fly rapidly with continuous wing beats, tend to have moderately long, pointed wings. Many songbirds use their short, rounded wings to move with quick wing beats from perch to perch or from ground to perch. Ducks have pointed wings that, beaten at high speed, provide rapid flight for long distances. Swallows, terns, and frigate birds have long, pointed wings that enable these birds to fly and maneuver gracefully for hours with leisurely wing beats. Large herons with long, broad wings travel far with slow, measured strokes, while buzzards soar high in the sky on their long, broad wings. Gulls and albatrosses, with their long, narrow wings flapping infrequently, sail along the beaches or over the waves. Swifts and hummingbirds, with their narrow, curved wings, fly rapidly and maneuver easily. A hummingbird can whir its tiny wings so rapidly that it can hover as it thrusts its long bill into a blossom; it can even fly backward as it leaves the bloom.

The shape of a bird’s tail also appears to be related to flight. The forked tails of frigate birds and terns enable quick changes of direction, and the barn swallow uses its deeply forked tail in making the intricate patterns of its graceful flight. A goshawk pursuing its prey through the forest uses its long tail in making quick turns. There is, however, such great diversity in birds’ tails that the precise size and shape probably is not of critical importance. For example, ducks, with their short tails, have a swift but direct flight, but long, graduated tails are often found in rapid, direct fliers such as some parrots and doves. Woodpeckers and some other climbing birds have strong tail feathers with stout shafts, which they use as props while on the trunks of trees.

The speed with which birds fly also varies greatly, and of course individual birds can vary their speed. Data on the speed of bird flight are difficult to evaluate. One of the complicating factors is that a bird’s speed in relation to the ground may depend on the force of the wind. Despite the variables involved in determining a bird’s speed of flight, the following generalized speeds, based on level flight in calm air, appear to be sound:Britannica QuizAnimal Group Names

15–30 km/hr (kilometres per hour) (10–20 mph [miles per hour])—many small songbirds such as sparrows and wrens
30–50 km/hr (20–30 mph)—many medium-sized birds such as thrushes and grackles, and larger, long-winged birds such as herons, pelicans, and gulls
30–60 km/hr (20–40 mph)—many small- and medium-sized birds such as starlings, chimney swifts, and mourning doves
60–100 km/hr (40–60 mph)—the faster-flying birds such as falcons, ducks, geese, and domestic pigeons. A homing pigeon has been timed at 152 km/hr (94 mph).
The fastest bird, however, is the peregrine falcon, whose speed in a dive has been measured in excess of 320 km/hr (200 mph).

The record long-range flight of a bird species in a single season is undoubtedly held by the Arctic terns that migrate from a summering ground in the Arctic to a wintering ground in the Antarctic, travelling more than 11,600 km (7,200 miles) each way. Some long-range flights are made very quickly: a blue-winged teal banded in Canada was recovered 6,100 km (3,800 miles) away in Venezuela only 30 days later; a Manx shearwater, trapped at its nest in Wales and transported 5,200 km (3,200 miles) to Massachusetts and released, returned home in 121/2 days. Some very small birds regularly make long water crossings in a single flight. Ruby-throated hummingbirds fly across the more than 800-km- (500-mile-) wide Gulf of Mexico, and many warblers fly from the American coast to Bermuda, a journey of about the same distance. For further information, see migration.

Flightlessness

Observe flightless cormorants in their natural habitat on coasts of Fernandina and Isabela islandsSee all videos for this article

Some birds have completely lost the power of flight during the course of evolution. The close similarity in the basic structure of flightless and flying birds, however, indicates that they all had a common flying ancestor. The rudimentary wings and the flightless condition of penguins and the ratites (ostriches and the like) is therefore a secondary, specialized condition. That flightlessness is a secondary condition is made still more apparent in other flightless birds that belong to families most of whose members are capable of flight. The extinct great auk of the North Atlantic is one of the best-known examples of such a flightless bird; the rail family also is noted for having many flightless species living on islands in the Pacific and the South Atlantic. Loss of flight seems to occur most often on isolated islands where there are no mammalian predators. In New Zealand, where there are no native land mammals of any kind, there were many species of extinct flightless moas, and there are still flightless kiwis, penguins, and rails as well as a duck, an owl, and several songbirds that are nearly flightless. The ratites of South America (rhea), Africa (ostrich), and Australia (cassowary) present an apparent contradiction to this correlation of mammal-free island habitats with bird flightlessness. Another adaptation, however—their great size—has enabled these birds to escape predation by mammals.

Walking and hopping

The bipedal gait of birds, dictated by modification of the forelimbs for flight, necessitates manipulation of food by the bill and feet. This poses problems in balance. The relative lengths of the segments of the legs must be such that, as the bird shifts from a standing to a sitting position, its centre of gravity remains over the feet. As some birds moved out of the trees and became terrestrial or aquatic, their legs were accordingly modified. In very large, slow-moving birds such as moas, the leg bones became very heavy. The toes became shorter, and the opposable first toe has been lost in rapidly running forms such as rheas and ostriches. The ostrich is the fastest runner, crossing stretches of savanna at a speed of 72.5 km/hr (45 mph). The toes became very long in birds that walk on aquatic vegetation or soft ground. Jacanas with their greatly elongated toes and claws walk over floating water weeds, and herons with long legs wade in shallow water. Wading birds developed long, thin legs, and climbing birds developed short legs with strongly curved, sharp claws. In swimming and diving birds, webs developed between the toes or lobes on the sides of the toes.

Terrestrial birds such as pheasants tend to walk; arboreal songbirds tend to hop as they travel from branch to branch. Tree dwellers such as woodpeckers, toucans, and the other members of order Piciformes, as well as parrots, can travel easily up and down trees because both of their outer toes face backward; in almost all other birds, only one toe faces to the rear. Parrots often walk along branches, and house sparrows hop when they come to the ground, while palm warblers walk on the ground and some songbirds, such as American robins and European blackbirds, may both walk and hop. Some birds with small feet, such as swifts, hummingbirds, bee-eaters, and many hornbills, use their feet only for perching and rarely walk at all. Other birds with robust feet, such as guinea fowl and rails, do most of their moving about on foot.

The usual position of a bird’s body in walking is more or less parallel to the ground. But the penguins, with their feet far to the rear of their bodies, stand upright as they waddle along. When the Adélie penguin makes its trek of many miles over the snow-covered ice to its breeding grounds, it may vary its awkward waddle with periods of tobogganing—i.e., sliding along on its breast and propelling itself with thrusts of its feet.

Swimming and diving

Some birds (auks, diving petrels, and certain ducks) use the wings for propulsion underwater as well as in the air. The wings of penguins have become highly modified into paddles that allow them to “fly” underwater; they use their webbed feet only for steering. Auks, on the other hand, use both their wings and webbed feet in swimming underwater. Several other water birds have become so adapted to swimming that they are practically helpless on land. In this class are loons, which shuffle awkwardly the few feet from the water to their nests. Swimming in birds is usually correlated with webbed feet, but coots and grebes, with only lobes on their toes, also swim and dive, and gallinules, with neither webs nor lobes, commonly swim. On the other hand, frigate birds, with partly webbed feet, never swim.

Some birds, such as the mallard, usually swim at the surface, feeding only as far underwater as they can reach by dipping their heads. Other ducks, such as scoters and pochards, commonly dive to the bottom for their food, and cormorants, auks, and loons pursue fish underwater. Sometimes loons are taken at remarkable depths in fishermen’s nets and on set lines, indicating that they may dive as deep as 61 metres (200 feet). Emperor penguins, however, are the best divers, having been recorded at depths of 483 metres (1,584 feet).

Pond ducks, such as mallards and teals, spring straight up from the water’s surface into the air in flight, but many swimming birds—for example, coots, grebes, cormorants, and diving ducks—take off with a long spattering run along the surface.Austin L. Rand

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Behaviour

Birds depend to a great extent on innate behaviour, responding automatically to specific visual or auditory stimuli. Even much of their feeding and reproductive behaviour is stereotyped. Feather care is vital to keep the wings and tail in flying condition and the rest of the feathers in place, where they can act as insulation. Consequently preening, oiling, shaking, and stretching movements are well developed and regularly used. Some movements, such as the simultaneous stretching of one wing, one leg, and half the tail (all on the same side) are widespread if not universal among birds. Stretching both wings upward, either folded or spread, is another common movement, as is a shaking of the whole body beginning at the posterior end. Other movements have evolved in connection with bathing, either in water or in dust. Such comfort movements have frequently become ritualized as components of displays.

Many birds maintain a minimum distance between themselves and their neighbours, as can be seen in the spacing of a flock of swallows perched on a wire. In the breeding season most species maintain territories, defending areas ranging from the immediate vicinity of the nest to extensive areas in which a pair not only nests but also forages. The frequency of actual fighting is greatly reduced by ritualized threat and appeasement displays. Birds range from solitary (e.g., many birds of prey) to highly gregarious, such as the guanay cormorants of the Peru Current off the west coast of South America, which nest in enormous colonies of hundreds of thousands and feed in large flocks with boobies and pelicans.

Sound

Auditory signals, like visual ones, are almost universal among birds. The most familiar vocalization of birds is that usually referred to as “song” (see birdsong). It is a conspicuous sound (not necessarily musical) that is used, especially early in the breeding season, to attract a mate, to warn off another bird of the same sex, or both. As such it is usually associated with establishing and maintaining territories. Individual variation in songs of many species is well known, and it is believed that some birds can recognize their mates and neighbours by this variation. Many other types of vocalizations are also known. Pairs or flocks may be kept together by series of soft location notes. Alarm notes alert other individuals to the presence of danger; in fact, the American robin (and probably many other species) uses one note when it sees a hawk overhead and another when it sees a predator on the ground. Begging calls are important in stimulating parents to feed their young. Other calls are associated with aggressive situations, courtship, and mating. Nonvocal sounds are not uncommon. Some snipe and hummingbirds have narrow tail feathers that produce loud sounds when the birds are in flight, as do the narrowed outer primaries of the American woodcock. The elaborate courtship displays of grouse include vocalizations as well as stamping of the feet and noises made with the wings. Bill clapping is a common part of courtship in storks, and bill snapping is a common threat of owls.

Nesting

Most birds build nests in which the eggs are laid. Nests vary widely: they may be a scrape in the sand, a deep burrow, a hole in a tree or rock, an open cup, a globular or retort-shaped mass with a side entrance tube, or an elaborately woven hanging structure. The materials with which nests are made also vary widely. Some nests are lined with small stones, and others are built of dirt or mud with or without plant material. Sticks, leaves, algae, rootlets, and other plant fibres are used alone or in combination. Some birds seek out animal materials such as feathers, horsehair, or snakeskin. The nest materials may be held together by weaving, sewing, or felting the materials themselves or with mud or spider webs. Swifts use saliva to glue nest materials together and to attach the nest to the supporting structure. In at least one species of swift, the entire nest is made of saliva and is the prized ingredient of birds’ nest soup in the Orient.More From BritannicaWhy Don’t Birds Have Teeth?

European cukoo egg in a reed warbler nest

fledgling European cuckoo being fed by an adult reed warbler

All birds incubate their eggs, except megapodes (mound builders), which depend on the heat generated by decaying vegetation or other external sources, and brood parasites such as cuckoos and cowbirds, which lay their eggs in the nests of other species. Murres and the king and emperor penguins build no nest but incubate with the egg resting on top of the feet.

In most birds a brood patch on the abdomen is developed. This bare area is fluid-filled (edematous) and highly vascularized; it directly contacts the eggs during incubation. Its development during the breeding season is under hormonal control. When the parent is off the nest, adjacent feathers are directed over the brood patch, and it is usually not apparent. A few birds (e.g., boobies) keep their webbed feet over the eggs during incubation.

Incubation takes from 11 to 80 days, depending at least in part on the size of the bird and the degree of development at hatching. Most songbirds and members of some other groups are hatched nearly absent of feathers and helpless (altricial), and they are brooded until well able to regulate their body temperature. They are fed by the parents even after they are capable of flight. The young of numerous other birds, such as chickens, ducks, and shorebirds, are hatched with a heavy coat of down and are capable of foraging for themselves almost immediately (precocial). Still others, such as the petrels and the auks, are downy when hatched but remain in the nest and are fed by their parents.

The length of time that parents care for young birds varies widely. Young megapodes can fly shortly after hatching and are entirely independent of their parents; young royal albatrosses may spend more than eight months at the nest and in the area immediately around it before they can fly. The length of time needed to attain independence is related to size and condition at hatching. Ground-nesting birds tend to take less and hole-nesting birds more time than the average.

The number of eggs in a set (clutch) varies from 1 to about 20. Some species invariably lay the same number per clutch (determinate laying), whereas in the majority the number is variable (indeterminate laying). In species of the latter category, clutch size tends to be smaller in tropical regions than in cold ones. There is also a tendency for birds in warm regions to make more nesting attempts in a given season. In the Arctic, where the season is very short, the cycle of breeding and the molt that follows it are compressed into a minimum of time.

Feeding habits

See the black skimmer water bird skim the calm surface with its longer lower mandible in search of preySee all videos for this article

The earliest birds were probably insectivorous, as are many modern ones, and the latter have evolved many specializations for catching insects. Swifts, swallows, and nightjars have wide gapes for catching insects on the wing; some woodpeckers can reach wood-boring grubs, whereas others can catch ants by probing anthills with their long, sticky tongues; thrashers dig in the ground with their bills; tree creepers and woodhewers probe bark crevices; and warblers glean insects from many kinds of vegetation. Raptorial birds (raptors and owls) have evolved talons and hooked bills for feeding on larger animals, and vultures have bare heads and tearing bills for feeding on carrion. Herons have spearlike bills and trigger mechanisms in the neck for catching fish, while kingfishers, terns, and boobies plunge into the water after similar prey. Long-billed waders probe for worms and other invertebrates. Of the many kinds of birds that feed on plant material, most use seeds, fruit, or nectar, which are high in food value; leaves and buds are eaten by fewer species. While some kinds of birds feed entirely on a single kind of food, others may take a wide range of foods, and many have seasonal changes in diet.

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Form and function

The structures associated with flight, even if they are vestigial or specialized for terrestrial or aquatic locomotion, easily distinguish birds from other animals. Whereas various skeletal and internal features are diagnostic of birds, feathers are unique to and present on all birds. Also unique to birds is their sound-producing organ, the syrinx. This avian analog to the voice box (larynx) is most highly developed in the songbirds. The syrinx is located where the trachea (windpipe) divides into the bronchial tubes. Sound is produced by airflow that vibrates membranes formed from part of the trachea, bronchi, or both.

Feathers

Like the scales of reptiles, and those on the feet of birds, feathers are made of keratin, a fibrous protein also found in hair. Feathers vary considerably in structure and function. Contour feathers form most of the surface of the bird, streamlining it for flight and often waterproofing it. The basal portion may be downy and thus act as insulation. The major contour feathers of the wing (remiges) and tail (rectrices) and their coverts function in flight. Contour feathers grow in tracts (pterylae) separated by bare areas (apteria) and develop from follicles in the skin.

The typical contour feather consists of a tapered central shaft, the rachis, with paired branches (barbs) on each side. An unbranched basal section of the rachis is called the calamus, part of which lies beneath the skin. The barbs, in turn, have branches, the barbules. The barbules on the distal side of each barb have hooks (hamuli) that engage the barbules of the next barb. The barbs at the base of the vane are often plumaceous—i.e., lacking in hamuli and remaining free of each other. In many birds each contour feather on the body (but rarely on the wings) is provided with a complex branch, the aftershaft, or afterfeather, that arises at the base of the vane. The aftershaft has the appearance of a second, smaller feather, growing from the base of the first. Down feathers have loose-webbed barbs, all rising from the tip of a very short shaft. Their function is insulation, and they may be found in both pterylae and apteria in adult birds. They also constitute the first feather coat of most young birds. Filoplumes are hairlike feathers with a few soft barbs near the tip. They are associated with contour feathers and may be sensory or decorative in function. Bristlelike, vaneless feathers occur around the mouth, eyes, and nostrils of birds. They are especially conspicuous around the gape (corners of the mouth) of birds that catch insects in the air. Some bristles function as eyelashes on ground-dwelling birds, and the bristles over the nostrils may serve as filters.

Molting

The contour feathers are shed and replaced (molted) at least once a year, usually just after the breeding season. In addition, many birds have at least a partial molt before the breeding season. A typical series of molts and plumages would be juvenal plumage, postjuvenal (also called first prebasic) molt, first winter (or first basic) plumage, first prenuptial (or pre-alternate) molt, first nuptial (or alternate) plumage, first postnuptial (first annual, or second prebasic) molt, second winter (or basic) plumage, etc. Molt of the remiges and rectrices usually occurs as part of the annual molt and can be serial, from the innermost feather out (centrifugal), from the outermost in (centripetal), or simultaneous. Normally the process is symmetrical between the right and left sides.

Flight, so characteristic of birds, is maintained during the molt in most species by a gradual replacement of the flight feathers. However, ducks and geese, some rails and loons, and auks shed all of their flight feathers at one time, immediately after the nesting season. Not until these feathers are replaced are the birds able to fly again. Most of these are birds that find their food by walking or swimming, as would be expected. Some ducks living in the marshes become very shy and retiring at this season, skulking in the reeds, but geese nesting in the Arctic barrens continue to walk about over the tundra, feeding. In the hornbills of Africa and Asia, only females lose the flight and tail feathers at the same time. During this time they stay in the nest until the feathers grow out again, being fed by the males.

Colour

Rivoli's hummingbird — Rivoli’s hummingbird

Colour in birds is caused by pigments and structure. Buffs, red browns, dark browns, and blacks are caused by melanins, pigments synthesized by the bird and laid down in granules. Yellows, oranges, and reds come from carotenoid or lipochrome pigments; these originate at least in part from the food and are diffused in the skin and feathers. Porphyrin feather pigments occur in birds but less frequently than melanins and carotenoids. Blue colours in feathers are structural, based on a thin, porous layer of keratin overlying melanin pigment. Most greens result from the addition of yellow pigment to the structural blue colour. Iridescent colours result from the thinly laminated structure of the barbules and are enhanced by underlying melanin deposits.

Other external features

Birds’ feet are covered with scales like those of reptiles. The scales are occasionally shed, but the timing of this molt is not known. The toes are tipped with claws, and vestigial claws are not infrequently found on the tips of the first two digits of the wing.

The bill is covered with a sheet of keratin, the rhamphotheca, which in petrels and a few other birds is divided into plates. In birds that probe for food (kiwis, woodcocks, etc.), many sensory pores are found near the tip of the bill. Both melanins and carotenoids are found in the rhamphotheca and in the scales of the feet.

The skin of a bird is almost without glands. The important exception is the preen gland, which lies on the rump at the base of the tail. The secretion of this gland contains approximately one-half lipids (fats and oils) and is probably important in dressing and waterproofing the plumage. In a few birds, the secretion has a strong, offensive odour. Some birds, in which the gland is small or absent, have a specialized type of feather (powder down) that grows continuously and breaks down into a fine powder, believed to be used in dressing the plumage.

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Skeleton

The avian skeleton is notable for its strength and lightness, achieved by fusion of elements and by pneumatization (i.e., presence of air cavities). The skull represents an advance over that of reptiles in the relatively larger cranium with fusion of elements, made possible by the fact that birds have a fixed adult size. Birds differ from mammals in being able to move the upper mandible rather than the lower, relative to the cranium. When the mouth is opened, both the lower and upper jaws move: the former by a simple, hingelike articulation with the quadrate bone at the base of the jaw, the latter through flexibility provided by a hinge between the frontal and nasal bones. As the lower jaw moves downward, the quadrate rocks forward on its articulation with the cranium, transferring this motion through the bones of the palate and the bony bar below the eye to the maxilla, the main bone of the upper jaw.

The number of vertebrae varies from 39 to 63, with remarkable variation (11 to 25) within the neck (cervical) series. The principal type of vertebral articulation is heterocoelous (saddle shaped). Each of the 3 to 10 (usually 5 to 8) chest (thoracic) vertebrae normally bears a pair of complete ribs consisting of a dorsal vertebral rib articulating with the vertebra and with the ventral sternal rib, which in turn articulates with the breastbone (sternum). Each vertebral rib bears a flat, backward-pointing spur, the uncinate process, characteristic of birds. The sternum, ribs, and their articulations form the structural basis for a bellows action, by which air is moved through the lungs. Posterior to the thoracic vertebrae is a series of 10 to 23 fused vertebrae, the synsacrum, to which the pelvic girdle is fused. Posterior to the synsacrum is a series of free tail (caudal) vertebrae and finally the pygostyle, which consists of several fused caudal vertebrae and supports the tail feathers.

The sternum consists of a plate lying ventral to the thoracic cavity and a median keel extending ventrally from it. The plate and keel form the major area of attachment for the flight muscles. The bones of the pectoral girdle consist of the wishbone (furcula) and the paired coracoids and shoulder blades (scapulae). The sword-shaped scapula articulates with the coracoid and upper “armbone” (humerus) and lies just dorsal to the rib basket. The coracoid articulates with the forward edge of the sternum and with the scapula, humerus, and furcula. The furcula connects the shoulder joints with the anterior edge of the keel of the sternum. It consists of paired collarbones (clavicles) and, probably, the median, unpaired interclavicle.

The bones of the forelimb are modified for flight with feathers. Major modifications include restriction of the motion of the elbow and wrist joints to one plane, reduction of the number of digits, loss of functional claws, fusion of certain bones of the “hand” (the metacarpals and most of the carpals) into a carpometacarpus, and modification of the elements, especially those toward the tip of the limb (distal), for the attachment of feathers. The wing bones are hollow, and the cavity in the humerus is connected with the air-sac system. As a general rule, large flying birds have proportionally greater pneumaticity in the skeleton than small ones. The highly pneumatic bones of large flying birds are reinforced with bony struts at points of stress. The humerus, radius, and ulna are well developed. The secondary flight feathers are attached to the ulna, which thus directly transmits force from the flight muscles to these feathers and is therefore relatively heavier than the radius. Two small wrist bones are present: the radiale, or scapholunar, and the ulnare, or cuneiform. The former lies between the distal end of the radius and the proximal part (the part toward the body) of the carpometacarpus. When the elbow joint is flexed (bent), the radius slides forward on the ulna and pushes the radiale against the carpometacarpus, which in turn flexes the wrist. Thus the two joints operate simultaneously. The U-shaped ulnare articulates with the ulna and the carpometacarpus. Anatomists differ on which bones of the reptilian “hand” are represented in the bird’s wing. Embryological evidence suggests that the digits are II, III, and IV, but it is possible that they are actually I, II, and III. The carpometacarpus consists of fused carpals (bones of the wrist) and metacarpals (bones of the palm), metacarpals II and III (or III and IV) contributing the greater part of the bone. The bones of the “fingers” (phalanges) are reduced to one each on the outer and inner digits and two on the middle one. The primary flight feathers are attached to the carpometacarpus and digits, the number attached to each being characteristic of the various major groups of birds.

The pelvic girdle consists of three paired elements, the ilia, ischia, and pubes, which are fused into a single piece with the synsacrum. The ilium is the most dorsal element and the only one extending forward of the socket of the leg (acetabulum). The ilium is fused with the synsacrum and the ischium, the latter of which is fused with the pubis. All three serve as attachments for leg muscles and contribute to the acetabulum, which forms the articulation for the femur. The leg skeleton consists of the thighbone (femur), main bone of the lower leg (tibiotarsus), fibula, fused bones of the ankle and middle foot (tarsometatarsus), and toes (phalanges). The fibula is largest at its upper end, where it forms part of the knee joint and tapers to a point distally, never forming part of the ankle joint. The latter joint is simplified, there being but two bones involved: the tibiotarsus, consisting of the tibia (the so-called shinbone in man) fused with the three upper ankle bones (proximal tarsals), and the tarsometatarsus, resulting from the fusion of metatarsals I through IV and the distal row of tarsals. Metatarsals II through IV contribute most to the tarsometatarsus. The basic number of phalanges (sections) on the toes is two, three, four, and five, respectively, one more than the number of the toe. Most birds have four toes, the fifth being always absent, but there are many variations in the number of digits, or phalanges, representing reductions of the basic arrangement.

The basic avian foot is adapted for perching. The first, or hind, toe (hallux) opposes the other three, and the tendons for the muscles that bend the toes pass behind the ankle joint in such a way that when the ankle is bent the toes are also. The weight of a crouched bird thus keeps the toes clasped around the perch.

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Muscles and organs

The cardiac (heart) muscles and smooth muscles of the viscera of birds resemble those of reptiles and mammals. The smooth muscles in the skin include a series of minute feather muscles, usually a pair running from a feather follicle to each of the four surrounding follicles. Some of these muscles act to raise the feathers, others to depress them. The striated (striped) muscles that move the limbs are concentrated on the girdles and the proximal parts of the limbs. Two pairs of large muscles move the wings in flight: the pectoralis, which lowers the wing, and the supracoracoideus, which raises it. The latter lies in the angle between the keel and the plate of the sternum and along the coracoid. It achieves a pulleylike action by means of a tendon that passes through the canal at the junction of the coracoid, furcula, and scapula and attaches to the dorsal side of the head of the humerus. The pectoralis lies over the supracoracoideus and attaches directly to the head of the humerus. In most birds the supracoracoideus is much smaller than the pectoralis, weighing as little as one-twentieth as much; in the few groups that use a powered upstroke of the wings (penguins, auks, swifts, hummingbirds, and a few others), the supracoracoideus is relatively large.

Avian striated muscles contain a respiratory pigment, myoglobin. There are relatively few myoglobin-containing cells in “white meat,” whereas “dark meat” derives its characteristic colour from their presence. The former type of muscle is used in short, rapid bursts of activity, whereas the latter is characteristic of muscles used continuously for long periods and especially in muscles used during diving.

The circulatory system of birds is advanced over that of reptiles in several ways: (1) there is a complete separation between pulmonary circulation (lungs) and systemic (body) circulation, as in the mammals, (2) the left systemic arch (aortic artery) is lost, blood passing from the heart to the dorsal aorta via the right arch, (3) the postcaval vein is directly connected with the renal portal that connects the kidneys with the liver, and (4) the portal circulation through the kidneys is greatly reduced. Birds’ hearts are large—0.2 to over 2.4 percent of body weight, as opposed to 0.24 to 0.79 percent in most mammals.

The avian lung differs from the type found in other land vertebrates in that several pairs of nonvascular air sacs are connected with the lungs. These extend into the pneumatic parts of the skeleton. Muscles between the sternum and trachea or along the trachea and bronchi vary tension on the membranes.

The avian digestive system shows adaptations for a high metabolic rate and flight. Enlargements of the esophagus, collectively called the crop, permit the temporary storage of food prior to digestion. The stomach is typically divided into a glandular proventriculus and a muscular gizzard, the latter lying near the centre of gravity of the bird and compensating for the lack of teeth and the generally weak jaw musculature. Otherwise, the digestive system does not vary markedly from the general vertebrate type.

Like reptiles, birds possess a cloaca, a chamber that receives digestive and metabolic wastes and reproductive products. A dorsal outpocketing of the cloaca, the bursa of Fabricius, controls antibody-mediated immunity in young birds. The bursa regresses with age, and thus its presence or absence may be used to determine age.

The testes of the male bird are internal, like those of reptiles. Intromittent organs are found in only a few groups (waterfowl, curassows, tinamous, ratites). The distal part of the vas deferens (the seminal sac) becomes enlarged and convoluted in the breeding season and takes on both secretory and storage functions. In songbirds this enlargement and the adjacent part of the cloaca form a cloacal protuberance, a swelling visible on the outside of the bird. Usually only the left ovary and oviduct are functional. As the egg moves down the oviduct, the albumen, membranes, and shell are laid down. The gonads and accessory sexual organs of both sexes enlarge and regress seasonally. In the breeding season, the testes of finches may increase more than 300 times their winter size.

Birds are homeothermic (warm-blooded) and maintain a body temperature of approximately 41 °C (106 °F). This temperature may be slightly less during periods of sleep and slightly higher during intense activity. Feathers, including down, provide effective insulation. In addition, layers of subcutaneous fat add further insulation in penguins and some other water birds. Heat loss through the feet in cold weather is minimized by reducing blood flow to the feet and by a heat-exchange network in the blood vessels of the upper leg, so that the temperature of blood flowing into the unfeathered part of the leg is very low.

Birds do not possess sweat glands. Excess heat is dissipated by rapid panting, which reaches 300 respirations per minute in domestic hens. Some heat can also be lost by regulation of blood flow to the feet. In hot climates, overheating is often prevented or reduced via behavioral means by concentrating activities in the cooler parts of the day and seeking shade during the hot periods. Temporary hypothermia (lowered body temperature) and torpor are known for several species of nightjars, swifts, and hummingbirds. Torpor at night is believed to be widespread among hummingbirds. The heart rate of birds varies widely—from 60 to 70 beats per minute in the ostrich to more than 1,000 in some hummingbirds.

The kidneys lie in depressions that are located on the underside of the pelvis. The malpighian bodies, which are the active tubules of the kidney, are very small in comparison with those of mammals, ranging from 90 to 400 per cubic mm. More than 60 percent of the waste nitrogen is excreted as uric acid or its salts. There is some reabsorption of water from the urine in the cloaca, with uric acid remaining. There is no urinary bladder, the urine being voided with the feces. In marine birds, salt is excreted in a solution from glands lying above the eyes through ducts leading to the nasal cavity.Robert W. Storer

Evolution and paleontology

The origin of birds, feathers, and avian flight have long been hotly debated. That birds evolved from reptilian ancestors is universally accepted, but did they evolve from thecodont reptiles or theropod dinosaurs? How and when did feathers evolve from the scaly reptilian integument? Did avian flight evolve from a terrestrial ancestor or an arboreal one? Such debates continue to be fueled by regular discoveries in China and Mongolia of remarkably preserved fossils from a critical period in avian evolution—the Early Cretaceous (145.5 million to 99.6 million years ago). The diversity of theropod dinosaurs, some with feathers, has greatly expanded our perspective of the evolution and early diversification of birds. Two major groups of early birds diversified dramatically in the Cretaceous Period: the Enantiornithines, an extinct subclass of birds; and Odontornithes, a primitive offshoot of the subclass Ornithurae that gave rise to modern birds at the end of the Cretaceous. Paleontologists now position Archaeopteryx, once thought to be a direct ancestor of birds, and several other fossil taxa at the base of the extinct enantiornithine lineage.

The origin of birds

The debate over the origin of birds centres on whether birds descended directly from thecodont reptiles about 230 million years ago (during the Triassic Period) or from a later lineage, the carnivorous theropod dinosaurs. This debate has been long-standing and divisive. At the beginning of the 21st century, the pendulum has swung decisively toward the theropod ancestor hypothesis—that today’s birds are feathered dinosaurs. This hypothesis is supported by analyses of shared characteristics (synapomorphies) combined with improved samples of early bipedal theropods.

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The origin of feathers

Feathers are complex and novel evolutionary structures. They did not evolve directly from reptilian scales, as once was thought. Current hypotheses propose that they evolved through an invagination of the epidermis around the base of a dermal papilla, followed by increasing complexity of form and function. They evolved before birds and even before avian flight. Thus, early feathers functioned in thermal insulation, communication, or water repellency, but not in aerodynamics and flight. Among extinct life-forms, feathers are no longer considered a unique and diagnostic characteristic of birds. Feathers with modern features were present in a variety of forms on a variety of theropod dinosaurs. At least nine Cretaceous dinosaurs had featherlike structures. The details of some are questionable, but some, such as those of Sinornithosaurus and other basal dromaeosaurs, bear a resemblance to modern pennaceous feathers. Feathered dinosaurs did not survive the end of the Cretaceous Period, but birds did, and then they flourished.

The origin of flight

Experts continue to debate whether flight evolved through gliding by an arboreal ancestral bird or through aerial launching by a running terrestrial ancestor. Historically these two hypotheses have been strongly linked to, respectively, the thecodont origin hypothesis and theropod origin hypothesis. The shift of opinion toward the theropod hypothesis, however, does not resolve this debate, since feathers on the forelimbs of early birds could have facilitated the early stages of flight through either mode. Precursors of an effective flight stroke of the forelimbs were present in terrestrial bipedal theropods. In either case, the evolution of avian flight required a decoupling of coordinated movements of the forelimbs and hind limbs. It also depended on new neural links between forelimb and tail movements as well as on other elaborations essential to controlled flight without major (initial) compromises of terrestrial locomotion. Once controlled flight had evolved, the avian body plan was transformed into a powerful flight engine. The transformation was then followed by the loss of other capabilities—or, in some cases, of flight itself.

Fossil birds

Archaeopteryx lithographica, which was first discovered in southern Germany in 1861, was once known as the world’s earliest fossil bird. A series of fossils, each dated to approximately 150 million years ago during the Jurassic Period, were about the size of a magpie. It resembled some reptiles and differed from modern birds in many ways, notably: (1) the jaw contained teeth set in sockets, (2) the sternum was short and not keeled, (3) the bones were not pneumatic, (4) the first two metacarpals were free, resulting in three movable digits of the “hand,” all with functional claws. Avian characteristics of Archaeopteryx included the possession of feathers and other skeletal features indicating that the creature represented an intermediate stage between reptiles and modern birds. The absence of a keel on the short sternum indicates that Archaeopteryx did not fly but glided. An opposable hallux, indicative of a perching type of foot, and clawed digits on the hand point to an arboreal existence.

However, early in the 21st century, the discovery of Xiaotingia zhengi, a fossilized feathered dinosaur in China that shares several birdlike characteristics with Archaeopteryx, cast doubt on the notion that all birds descended from Archaeopteryx. X. zhengi appears in the fossil record some five million years before Archaeopteryx, and this fact suggests that the single ancestor from which all bird lineages originate has yet to be found. The earliest fossil bird known to be ancestral to the lineage of modern birds is Archaeornithura meemannae. The species was described in 2015 after having been found in rocks of the Huajiying Formation of northeastern China that date to 130.7 million years ago.

The other major group of toothed Cretaceous birds, the Odontornithes, included one of the best-known groups of fossil birds, Hesperornis and its relatives. These birds were highly specialized foot-propelled divers of the Late Cretaceous. Hesperornis was up to 1.8 metres (6 feet) long and had completely lost the power of flight. The sternum lacked a keel, the humerus was small and weak, and other elements of the wing were missing entirely. The pelvis and hind limb had a strong but superficial resemblance to those of modern loons and grebes. However, two major features and several less obvious ones indicate that the resemblance was the result of convergent evolution rather than common ancestry. Hesperornis was remarkable for three features. It had (1) teeth set in grooves, not sockets, (2) a stout fourth toe with a unique rotary ball-and-flange type of articulation, and (3) tail (caudal) vertebrae with limited vertical motion, making the tail somewhat beaverlike in its action. Baptornis, a contemporary relative of Hesperornis, was smaller and less strongly modified. Though flightless, its wings were less reduced than Hesperornis, and it lacked the peculiar modifications of the fourth toe and caudal vertebrae.

In the Late Cretaceous also appeared the first modern birds, assigned to the infraclass Neornithes, or Carinata. Living alongside Hesperornis and other Odontornithes was a group of flying birds that included Ichthyornis and Apatornis. Although not related to gulls, these birds resembled them superficially and may well have been their ecological counterparts. It was long believed that Ichthyornis had teeth, like Hesperornis, but it is now thought that the toothed jaws formerly thought to belong to Ichthyornis were really those of a small mosasaur, a marine reptile.

A genetic study of bird lineages estimated that birds began to lose the outer covering of enamel from their teeth, and possibly their teeth as well, some 116 million years ago during the Early Cretaceous. The study also showed that this change was accompanied by an increase in beak development that assisted in the diversification of living birds.

Modern birds

In the evolution of modern birds from an Archaeopteryx-like form, the development of active flight must have occurred early. This meant an increase in size of the muscles moving the wing and the development of a keel on the sternum as an added area of attachment for these muscles. As the tail took on more of a steering function and less of a supportive one, it became shorter and more readily moved as a unit. Feathers became specialized for different functions, and at the same time the eyes, brain, and respiratory and circulatory systems continued to develop in a manner associated with the evolution of homeothermic, arboreal, gliding animals. By the time birds became strong fliers, they were ready to exploit many new environments, and by the Cretaceous Period they had begun to do so, producing the wide array of adaptive types known today.

The major diversification of modern birds probably took place in the Cretaceous, and it must have started early in that period because fragmentary fossil evidence of foot-propelled divers (Enaliornis) and of an early relative of the flamingos (Gallornis) are known from Lower Cretaceous deposits in Europe. Upper Cretaceous deposits have yielded, besides Hesperornis and Ichthyornis and their relatives, diving birds similar to Enaliornis, other early flamingo-like birds, and species in the same suborders as gannets, ibises, rails, and shorebirds.

Deposits from the Paleocene Epoch (65.5 million to 55.8 million years ago) have yielded the earliest known loons, cormorants, New World vultures, and gulls. In addition, large, flightless predatory birds culminating in Diatryma made their appearance during this period. From the far richer Eocene Epoch (55.8 million to 33.9 million years ago) have come the earliest known fossil representatives of most of today’s bird orders.

Almost certainly all living orders and most living families of birds were in existence by the end of the Eocene. One of the most interesting finds from this period was fossils of Neocathartes, a long-legged bird allied to the New World vultures. There are several anatomical similarities between this group of vultures and the storks, and the existence of this fossil lends support to the idea that the storks and New World vultures are more closely related to each other than each family is to the birds with which it is usually grouped.

By Pliocene times (5.3 million to 2.6 million years ago), most modern genera were probably in existence. After the extinction of the dinosaurs and before large carnivorous mammals evolved, two groups of large flightless birds evolved to fill a similar niche. From the late Paleocene to the middle Eocene, Diatryma and its relatives were major predators in the Northern Hemisphere. The largest species stood over 2.25 metres (7 feet) tall and had stout hooked beaks. They are of uncertain relationships but may have been distantly related to the cranes and rails (order Gruiformes). The second group, that of Phororhacos and related genera, had a long history (from the lower Oligocene to the middle Pliocene) in South America, which was without large carnivores until relatively recent times. The Phororhacos line evidently evolved from seriema-like stock and radiated into numerous genera and species, the largest of them (Onactornis) standing 2.5 metres (8 feet) tall and having a skull 80 cm (31 inches) long and 40 cm (16 inches) high.

Large grazing or browsing birds appear to have evolved several times. On continents where there are large predators, these birds have always been rapid runners (ostriches, rheas, emus), but on islands lacking such predators, they were slow-moving, heavy-bodied birds. Two such groups were the elephant birds of Madagascar and the moas of New Zealand, the largest in each group approaching 3 metres (10 feet) in height. Fragmentary fossil material from Eocene and Oligocene deposits in Egypt indicates that similarly adapted birds occurred there before the advent of large carnivores.Frank Gill

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Classification

Distinguishing taxonomic features

In classifying birds, most systematists have historically relied upon structural characteristics to infer evolutionary relationships. Plumage characteristics include the number of various feather types; the presence or absence of down on the feather tracts and on the preen gland; and the presence or absence of an aftershaft. Characteristics of the bill and feet are also useful, as is the arrangement of bones in the palate and around the nostrils. The presence or absence of certain thigh muscles is considered, as are the arrangement of the carotid arteries, the syrinx, and the deep flexor tendons of the toes as well as the condition of the young when hatched. Advances in the study of DNA sequences and computerized construction of phylogenetic trees have provided new means of testing hypotheses of taxonomic relationships.

Critical appraisal

It has frequently been stated that birds are one of the best known of animal groups. This is true in the sense that most of the living species and subspecies in the world have probably been described; but because of inadequacies in the fossil record and repeated cases of convergent evolution within the group, our knowledge of the phylogenetic relationships between orders, suborders, and families of birds is inferior to that of mammals and reptiles. Most, if not all, of the major lineages of modern birds arose rapidly in the Late Cretaceous and the Paleogene Period (about 100 million to 23 million years ago). DNA data continue to resolve the relationships among major groups of birds. The penguins (Sphenisciformes), tube-nosed seabirds (Procellariiformes), and pelicans (Pelecaniformes) form a triad of related lineages. Waterfowl (Anseriformes) and chickenlike birds (Galliformes) are linked and together may be the oldest assemblage of modern birds. Some caprimulgiforms (owlet frogmouths) seem clearly related to swifts (Apodiformes) through a link between owlet frogmouths and treeswifts.

The taxonomic positions of several bird groups remain open to question. The hoatzin, included below in the Cuculiformes, is often given its own order, Opisthocomiformes. The sandgrouse are listed separately in order Pteroclidiformes. The turacos, sometimes included in the Cuculiformes, are considered by many authors to warrant separation and are listed here as Musophagiformes. Diatryma and several related genera of extinct flightless predators are often placed in a distinct order, Diatrymiformes, near Gruiformes. The flamingos, which constitute the order Phoenicopteriformes in some classifications, are placed in the Ciconiiformes in this classification, but their relationships are still unknown.

One area particularly in need of study is the relationships among the various groups of ratites (ostriches, rheas, emus, moas, and others). Formerly, some authorities argued that these birds and the penguins arose independently from cursorial reptiles, but it is now generally agreed that all of them passed through a flying stage in the course of their evolution. The ratite groups differ greatly in morphology and yet show remarkable similarities in palate and bill characters. The principal unanswered questions are how many different flightless lines evolved from flying ancestors and from how many different groups the flying ancestors evolved. On zoogeographic grounds, it is likely that the isolated kiwi-moa, elephant bird, and emu-cassowary lines arose independently from each other and from ratites on the other continents. But the ostriches and rheas could be descended from a common flightless ancestor because of the known former land connections from Asia to North and South America. Kiwis, ostriches, rheas, emus, and cassowaries are contained within order Struthioniformes in this classification.

The evolutionary sequence of the bird orders starts with ratites and marine seabirds and ends with songbirds. Beginning in the 1980s, Charles Sibley proposed radically different listings of the nonpasserine orders on the basis of his pioneering DNA analyses.

Annotated classification

This classification is a synthesis of current information compiled by American ornithologist Frank Gill (2002).

CLASS AVES (birds)10,100 living species of vertebrate (backboned) animals primarily adapted for flight with feathers. Warm-blooded with a 4-chambered heart; left systemic arch lost. Lower jaw articulates with cranium via the quadrate; teeth absent in living forms. Reproduction by hard-shelled eggs, nearly always incubated by one or both parents.
- Order Passeriformes (songbirds, or perching birds)5,700 species in 74 families (depending on the authority), worldwide; complex assemblage containing more than half of all known bird species; bill, plumage, and habits highly varied; length 7.5–125 cm (3–49 inches).
- Order Apodiformes (swifts, hummingbirds)Approximately 425 species in 3 families including crested swifts, worldwide except in the extreme north; hummingbirds limited to New World; rapid-flying birds that feed in flight upon insects or nectar; “hand” and primary flight feathers constitute a relatively great proportion of the wing; feet weak; length 6.3–23 cm (2.5–9.1 inches).
- Order Piciformes (woodpeckers and allies)Approximately 400 species in 6 families including jacamars, puffbirds, barbets, honey guides, toucans; worldwide in forests; hole-nesting birds that feed upon insects and fruit; outer toes able to face rearward; woodpeckers specialized for climbing; honey guides are brood parasites; length 7.5–61 cm (3–24 inches).
- Order Charadriiformes (gulls, sandpipers, auks, and allies)370 species in 17 families including plovers, jacanas, stilts, avocets, thickknees, terns, and murres; worldwide. Three basic body plans: suborder Charadrii—waders (shorebirds) that usually feed on small animals in mud or water; bill variable but often long and used for probing; Lari—web-footed, dense-plumaged water birds that feed by plunging into water for fish, robbing other birds, or scavenging; Alcae—dense-plumaged, web-footed, marine, wing-propelled divers that feed on fish or invertebrates; length 12–78 cm (4.7–30.7 inches).
- Order Pteroclidiformes (sandgrouse)16 species in 1 family. Stocky, pigeonlike ground birds with short legs but fast flight; feed on seeds and insects; deserts of Africa and Asia; length 22–40 cm (about 9–16 inches).
- Order Psittaciformes(parrots, lorikeets, cockatoos, kea, and kakapo)About 368 species in 2 families, 10 species extinct since 1600; tropical, with some temperate-zone species; often brightly coloured; strong-flying, seed-, fruit-, or nectar-eating birds with very stout, hooked bills and zygodactyl feet (i.e., outer toe facing rearward); length 8–100 cm (3.2–39 inches).
- Order Columbiformes (pigeons and doves)300-plus species in 1 family, worldwide except in the extreme north; fast-flying birds with pointed wings and weak bills; feed on seeds and fruit; length 15–120 cm (5.9–47.2 inches).
- Order Falconiformes (diurnalbirds of prey)309 species in 5 families including hawks, falcons, eagles, the secretary bird, Old World vultures, and condors; length 14–150 cm (5.5–59 inches), condor wingspan more than 3 metres (10 feet); some fossil forms larger.
- Order Galliformes (chickenlike birds)About 290 species in 5 families including pheasants, megapodes, guinea fowl, curassows, and guans; nearly worldwide, except southern South America; terrestrial or arboreal, with strong, scratching feet; short, rounded wings; feathers with long aftershafts; length 15 to more than 200 cm (5.9 to more than 79 inches).
- Order Gruiformes (cranes and allies)About 210 species in 11 families including rails, coots, moorhens; worldwide and diverse group, ranging from small quail-like hemipodes to large long-legged cranes, marsh-inhabiting rails, swimming coots and finfoots, and cursorial bustards; length 12–176 cm (4.7–70 inches). The carnivorous phororhacoids of the early Cenozoic Era belong here, as may the very large Diatryma and its relatives; fossils to 200 cm (6.6 feet) tall.
- Order Procellariiformes(tubenosed seabirds)117 species in 4 families including albatrosses, shearwaters, and petrels; oceans worldwide but most numerous in Southern Hemisphere; web-footed marine birds with tubular nostrils; possess a musky smell; most have narrow wings and stiff, gliding flight; length 13–200 cm (5.1–79 inches), albatross wingspan more than 3 metres (10 feet).
- Order Coraciiformes (kingfishers and allies)211 species in 10 families including hornbills, bee-eaters, rollers, hoopoes, todies, motmots; worldwide except in the extreme north; heterogeneous group of hole-nesting birds; many with long, pointed bills and blue or green in plumage; all have 2nd and 3rd or 3rd and 4th toes joined at base; food largely animal, except hornbills, which eat much fruit; length 10–120 cm (4–47 inches).
- Order Strigiformes (owls)180 species in 2 families worldwide, nocturnal raptorial birds with hooked beaks, strong talons, and soft plumage; length 12–69 cm (4.7–30 inches).
- Order Musophagiformes (turacos)18 species in 1 family, colourful plumage, fruit-eating; length 35–70 cm (14–28 inches); Africa.
- Order Cuculiformes (cuckoos and allies)141 species in 2 families including anis, roadrunners, and the hoatzin; one species extinct since 1600; worldwide except in the extreme north; long-tailed birds with rearward or sideward facing toes; feed on both fruits and small animals; most arboreal, a few terrestrial; some are brood parasites; length 16–76 cm (6.3–30 inches).
- Order Anseriformes (screamers, waterfowl)150 species 2 families worldwide, including ducks, geese, and swans; web-footed birds with broad bills containing fine plates or lamellae except for screamers, large-footed marsh birds with chickenlike bills; length 34–180 cm (13–71 inches).
- Order Ciconiiformes (herons, storks, and allies)120 species in 6 families including shoebills, New World vultures, ibises, bitterns; worldwide except in the extreme north; long-legged wading birds with long bills; feet not webbed; length 25–152 cm (9.7–60 inches).
- Order Caprimulgiformes (nightjars)121 species in 5 families including frogmouths, potoos, and the oilbird; worldwide except in the extreme north; nocturnal and concealingly coloured, with weak feet, soft plumage, and very large mouths; most feed on insects caught in flight; length 15–60 cm (6–24 inches).
- Order Pelecaniformes (pelicans and allies)66 species in 6 families worldwide, including cormorants, boobies, gannets, tropic birds, and frigate birds. Water birds with all 4 toes webbed; bill hooked or straight and sharply pointed; length 48–188 cm (19–74 inches).
- Order Tinamiformes (tinamous)47 species in 1 family; Central and South America; ground-dwelling birds resembling quails or pheasants with flat, elongated, and rather weak bills and very small tails; length 20–53 cm (8–21 inches).
- Order Trogoniformes (trogons)37 species in 1 family; tropical, except Australasia; extremely soft-plumaged arboreal birds that feed on insects and small fruit; feet weak; 1st and 2nd toes directed backward; length 23–40 cm (9.1–16 inches).
- Order Podicipediformes (grebes)22 species in 1 family worldwide, 2 species recently extinct; foot-propelled diving birds with lobed toes, minute tails, and silky plumage; length 20–78 cm (8–31 inches).
- Order Sphenisciformes (penguins)17 species in 1 family in oceans of the Southern Hemisphere; wings flipperlike for propulsion underwater; webbed feet short and stout; stance upright; feathers short and dense, molted in patches; length 35–115 cm (14–45 inches); fossil forms to 180 cm (71 inches).
- Order Gaviiformes (loons)5 species in 1 family of the Northern Hemisphere; foot-propelled diving birds with webbed feet and pointed bills; length 53–91 cm (21–36 inches).
- Order Coliiformes (colies, or mousebirds)6 species in 1 family of Africa south of the Sahara; soft plumage with long, pointed tails and all 4 toes directed forward; largely vegetarian, some insects; length 29–36 cm (11–14 inches).
- Order Struthioniformes (ostriches, rheas, emus, cassowaries, and kiwis)10 species in 6 families in Africa, South America, New Zealand, Australia, and Oceania, with fossils from southern Europe and Asia, including India and Mongolia; cursorial (running); height 35 cm to 2.7 metres (14 inches to almost 9 feet). Many species have small tails with little or no aftershaft. Some forms are nearly wingless. Order includes the largest living birds.

Frank Gill

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Also known as: Chordata

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Last Updated: Jun 18, 2023 • Article History

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Chordate

Category: Science & TechKey People: William BatesonRelated Topics: reptile vertebrate tunicate cephalochordate protochordate

chordate, any member of the phylum Chordata, which includes the vertebrates (subphylum Vertebrata), the most highly evolved animals, as well as two other subphyla—the tunicates (subphylum Tunicata) and cephalochordates (subphylum Cephalochordata). Some classifications also include the phylum Hemichordata with the chordates.

As the name implies, at some time in the life cycle a chordate possesses a stiff, dorsal supporting rod (the notochord). Also characteristic of the chordates are a tail that extends behind and above the anus, a hollow nerve cord above (or dorsal to) the gut, gill slits opening from the pharynx to the exterior, and an endostyle (a mucus-secreting structure) or its derivative between the gill slits. (A characteristic feature may be present only in the developing embryo and may disappear as the embryo matures into the adult form.) A somewhat similar body plan can be found in the closely related phylum Hemichordata.

General features

Tunicates are small animals, typically one to five centimetres (0.4 to 2.0 inches) long, with a minimum length of about one millimetre (0.04 inch) and a maximum length slightly more than 20 centimetres; colonies may grow to 18 metres (59 feet) in length. Cephalochordates range from one to three centimetres. Vertebrates range in size from tiny fish to the whales, which include the largest animals ever to have existed.

Tunicates are marine animals, either benthic (bottom dwellers) or pelagic (inhabitants of open water), that often form colonies by asexual reproduction. They feed by taking water in through the mouth, using the gill slits as a kind of filter. The feeding apparatus in cephalochordates is similar. They have a well-developed musculature and can swim rapidly by undulating the body. Cephalochordates usually live partially buried in marine sand and gravel.Britannica QuizAnimal Group Names

Vertebrates retain traces of a feeding apparatus like that of tunicates and cephalochordates. The gill slits, however, ceased to function as feeding structures, and then later as respiratory devices, as the vertebrate structure underwent evolutionary changes. Except in some early branches of the vertebrate lineage (i.e., agnathans) a pair of gill arches has become modified so as to form jaws. The fishlike habitus that evidently began with cephalochordates became modified by the development of fins that were later transformed into limbs. With the invasion of the vertebrates into fresh water and then onto land, there was a shift in means of breathing—from gills to lungs. Other modifications, such as an egg that could develop on land, also emancipated the vertebrates from water. Elaboration of the locomotory apparatus and other developments allowed a diversification of structure and function that produced the amphibians, reptiles, birds, and mammals.

Natural history

Reproduction and life cycle

The chordate life cycle begins with fertilization (the union of sperm and egg). In its primitive form, fertilization occurs externally, in the water. Asexual reproduction takes place in tunicates and in some vertebrates (females of some fish and lizards can reproduce without fertilization). Hermaphroditism (possessing both male and female reproductive organs) is found in tunicates and some fishes, but otherwise the sexes are separate. Larvae (very young forms that differ considerably from the juveniles and adults), when they do occur, differ in structure from the larvae of nonchordates. Internal fertilization, viviparity (giving birth to young that have undergone embryological development), and parental care are common in tunicates and vertebrates.

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Ecology and habitats

Chordates are common in all major habitats. Tunicate larvae either seek out a place where they can attach and metamorphose into an adult or develop into adults that float in the open water. Cephalochordates develop in the open water, but as adults they lie partially or entirely buried in sand and gravel. In either case, they are filter feeders with simple behaviour. Vertebrates are much more complex and, in keeping with their more active manner of obtaining food, highly varied in their ecology and habits.

Locomotion

Chordates are capable of locomotion by means of muscular movements at some stage in life. In tunicate larvae, this is accomplished using a tail; in cephalochordates, by undulations of the body; and in vertebrates, by general body movements (as in eels and snakes) and by the action of fins and limbs, which in birds and some mammals are modified into wings.

Associations

Chordates enter into a wide variety of symbiotic relationships and are especially noteworthy as hosts for parasites. Family groups and societal relationships, in both a broad and narrow sense, are particularly well developed in vertebrates, due primarily to their elaborate nervous systems. This phenomenon is seen in schools of fish, flocks of birds, and herds of mammals, as well as in the primate associations that suggest the beginnings of human society.

Form and function

General features

Chordates have many distinctive features, suggesting that there has been extensive modification from simple beginnings. The early stages of chordate development show features shared with some invertebrate phyla, especially the mouth that forms separately from the anus, as it does in the phyla Hemichordata, Echinodermata, and Chaetognatha. Likewise, as in these phyla, the coelom, or secondary body cavity around the viscera, develops as outpouchings of the gut. A coelom also is present in some more distantly related phyla, including Annelida, Arthropoda, and Mollusca, but the main organs of the body are arranged differently in these phyla. In chordates the main nerve cord is single and lies above the alimentary tract, while in other phyla it is paired and lies below the gut. Cephalochordates and vertebrates are segmented, as are the annelids and their relatives; however, segmentation in the two groups probably evolved independently. The gill slits and some other features that are common among the hemichordates and the chordates originated before the chordates became a separate group. Hemichordates have no tail above the gut and no mucus-secreting endostyle between the gill slits.

External features

An ancestral chordate, as suggested by the adult lancelet and the tadpole larva of tunicates, had a distinct front and hind end, an anterior mouth, a posterior tail above an anus, unpaired fins, and gill slits that opened directly to the exterior. A free-swimming tunicate larva metamorphoses into an attached, sessile adult with an atrium that surrounds the gills. The atrium of lancelets probably evolved independently.

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Internal features

Skeleton and support

The chordate notochord is a stiff rod with a turgid core and fibrous sheath. It keeps the animal from shortening when locomotory waves are produced through muscular contraction. The chordate body is supported by fluid in the body cavities. In tunicates, added support is provided by the tunic. Cartilaginous material supports the gills and other body parts of tunicates and cephalochordates. Immature vertebrate skeletons generally consist largely of cartilage, which becomes increasingly bony with age. The cartilaginous skeletons of sharks and some other vertebrates are thought to have evolved from more highly mineralized ones.

Tissues and muscles

In both cephalochordates and vertebrates, muscles used in locomotion are well developed and organized segmentally. The tail musculature of tunicates is simpler and without clear indications of segmentation. There is at least a small amount of musculature throughout the body of all chordates. As jaws, limbs, and other body parts have evolved in vertebrates, so have the muscles that operate them.

Nervous system and sense organs

The anterior end of the main nerve cord in chordates is enlarged to form at least the suggestion of a brain, but a brain is well developed only in vertebrates. Tunicate larvae have visual organs sensitive to light and sense organs responsive to the direction of gravity. Pigment spots and light receptors in the nerve cord of lancelets detect sudden changes in light intensity. The eyes and other sense organs of vertebrates are more elaborate and complex.

The presence in cephalochordates and vertebrates of a nervous system with segmentally repeated nerves arising from the dorsal hollow nerve cord is suggestive of a common ancestry. The tunicate nervous system does not have the segmentally repeated nerves. The brains of all vertebrates are greatly enlarged and subdivided into functionally specialized regions.

Digestion and nutrition

Both tunicates and cephalochordates are filter feeders of small particles of food suspended in the water. Beating cilia (hairlike cellular extensions) on the gill slits draw a current of water into the mouth and through the pharynx, where a sheet of mucus, secreted by the endostyle (a glandular organ lying below the two rows of gill slits), filters suspended food particles from the water. Cilia lining the pharynx move the food-rich sheet of mucus upward over the gill slits, and it is then rolled up and transported to the posterior part of the gut. The water current passes into the atrium and exits through the atrial opening.

Something similar to this arrangement occurs in the vertebrates in the “ammocoetes” larva stage of the primitive jawless fish called the lamprey. The difference is that the food consists of somewhat larger particles that have been deposited on the bottom (detritus), and, instead of the feeding current being driven by cilia, the pharyngeal musculature pumps water and food particles across the gill slits. The earliest fishes probably fed on detritus, and a sucking action is retained by their extant representatives (lampreys and hagfishes). With the development of jaws, it became possible for the vertebrates to capture and seize larger food items.

The lower digestive tract of the primitive chordate is a simple tube with a saclike stomach. There are only indications of the specialized areas and of glandlike structures, such as the liver and pancreas, that occur in vertebrates.

Excretion

The excretion of wastes and the control of the chemical composition of the internal environment are largely effected by kidneys, although other parts of the body, including the gills, may play an important role. Tunicates and cephalochordates have a salt content essentially the same as seawater, but vertebrates, even marine species, have body fluids of low salt content, with the exception of hagfishes. A possible explanation is that the vertebrates evolved in fresh water, but it seems reasonable that hagfishes branched off while still marine and that the freshwater form evolved later.

Respiration

A primitive chordate gill is present in tunicates and cephalochordates, where it serves in both respiration and feeding. The vertebrate gill may retain some role in feeding, although the current is now produced by the action of muscles, not cilia. The gills became reduced in number in various lineages, and they were strengthened by supporting elements, some of which evolved into jaws. Lungs, already present in fishes, became the main respiratory organs of terrestrial vertebrates.

Circulatory system

The circulatory system in chordates has a characteristic pattern. In tunicates and vertebrates the blood is propelled by a distinct heart; in cephalochordates, by contraction of the blood vessels. Unoxygenated blood is driven forward via a vessel called the ventral aorta. It then passes through a series of branchial arteries in the gills, where gas exchange takes place, and the oxygenated blood flows to the body, much of it returning to its origin via a dorsal aorta. The blood of vertebrates passes through the tissues via tiny vessels called capillaries. In tunicates and cephalochordates, capillaries are absent and the blood passes through spaces in the tissues instead.

Hormones

In vertebrates, endocrine glands (those of internal secretion) produce hormones that regulate many physiological activities. In tunicates and cephalochordates, organs have been identified that correspond in anatomical position to the pituitary gland of vertebrates, but which hormones, if any, they secrete is uncertain. In vertebrates, the thyroid gland produces thyroxine, an iodine-containing hormone that helps regulate metabolism. The thyroid is a modified endostyle, as can be illustrated by larval lampreys in which the thyroid still secretes mucus for use in feeding. The endostyles of lancelets take up iodine and form thyroxine, but the thyroxine formed may not function as a hormone in the lancelets themselves.

Features of defense and aggression

Tunicates largely rely upon the passive defense afforded by their heavy tunic. Lancelets move rapidly through the substrate, and their well-developed locomotory apparatus evolved largely to provide a means of escaping predators. Vertebrates have ceased to feed on detritus brought to them by water currents. They have shifted to consuming larger foodstuffs and to actively locating, pursuing, and subduing what they eat.

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Evolution and paleontology

Many scientists maintain that chordates originated sometime earlier than 590 million years ago; that is, they predate the fossil record. Such early representatives were soft-bodied and therefore left a poor fossil record. The oldest known fossil chordate is Pikaia gracilens, a primitive cephalochordate dated to approximately 505 million years ago. There is disagreement over whether older animals—such as Yunnanozoon lividum and Haikouella (both of which date to 530 million years ago and possess several chordate features)—should be considered chordates. An extensive vertebrate fossil record begins about 400 million years ago.

Embryological evidence places the phylum Chordata within the deuterostomes (bilaterally symmetrical animals with undeterminate cleavage and whose mouth does not arise from the blastopore), which also includes the phyla Hemichordata, Echinodermata, and Chaetognatha. The closest relatives of the chordates are probably the hemichordates, since these animals possess gill slits and other features not found in other animal phyla. A slightly more remote relationship to the echinoderms is inferred on the basis of resemblances between the larvae in some groups of hemichordates and echinoderms. The derivation of chordates from certain fossil echinoderms has been argued on the basis of features such as what appear to be gill slits. Theories that derive them from other phyla (e.g., Annelida, Nemertea, Arthropoda) have been proposed, but such theories have few contemporary advocates.

Whether the first ancestral chordate was more like a tunicate or a cephalochordate has been extensively debated. The classical theory is that the ancestor was like a cephalochordate and that one lineage became attached to hard surfaces and evolved into tunicates, whereas another remained unattached and evolved into vertebrates. An alternative theory is that the ancestor was like a tunicate and that the other two subphyla arose by modification of the tadpole larva. There is some preference for the classical theory because it provides the most satisfactory way of accounting for the similarities between chordates and hemichordates of the subphylum Enteropneusta. Within the chordates, the tunicates probably branched off before the common ancestor of cephalochordates and vertebrates arose, for the latter resemble each other in some details of neuroanatomy and biochemistry.

Classification

Annotated classification

PHYLUM CHORDATADeuterostomatous eucoelomates; gill clefts; endostyle or its derivative in pharynx; notochord; hollow dorsal nerve cord; tailposterior and dorsal to anus.
- Subphylum Tunicata (or Urochordata; tunicates)Notochord, when present, restricted to tail; body covered with tunic, but sometimes only cuticle; atrium, absent in Appendicularia, dorsal and often paired in embryonic development; heart present; generally sessile (attached) as adults; see below Tunicates.
  - Class Ascidiacea (sea squirts)Sessile; benthic; solitary or colonial within a common tunic.
  - Class Appendicularia (larvacea)Free-swimming; pelagic; resembles tadpole larvae of ascidians; 1 pair of gill slits; no distinct atrium.
  - Class ThaliaceaPelagic; forms aggregations or colonies.
- Subphylum Cephalochordata (or Acrania; lancelets)Notochord extends entire body length, with tip anterior to nerve cord; atrium a single cavity with single, ventral opening; segments well developed; head poorly developed; no paired fins; no heart; see below Cephalochordates.
- Subphylum Vertebrata (or Craniata; vertebrates)Notochord extends to the back of a well-developed head; no atrium; segments well developed; paired fins or limbs usually present; heart present; see below Vertebrates.

Critical appraisal

This outline gives the major groups of chordates. Modern systematic biology attempts to arrange groups of organisms in a way that suggests the genealogical relationships (branching sequences) and therefore presents an epitome of evolutionary history. It also may attempt to show where there are important differences among the various groups. These goals often conflict. In a purely genealogical system, each group must correspond to a single lineage (clade) composed of the common ancestor and all of its descendants. A group that does not meet both of these requirements is called a grade and may be used as an informal group. Groups that do not contain the common ancestor, and therefore had two separate origins, are said to be polyphyletic. Such polyphyletic grades, which would put whales together with fish or birds together with bats, have generally been abandoned as soon as they were recognized. Another kind of grade, which does not include all the descendants of the common ancestor, is said to be paraphyletic and is retained in more conservative systems. Within the vertebrates the class Aves is a clade, but the class Reptilia is a grade, for the birds are modified dinosaurs. Some systems do not recognize Reptilia as a formal group. Likewise, birds, mammals, reptiles, and amphibians are all modified fish, and the old class of fishes (Pisces) is now rarely used. Vertebrata is a single clade, but “invertebrate” is a grade consisting of all animals except vertebrates. Therefore there is no formal group called Invertebrata.Britannica QuizAnimal Factoids

Many differences among systems are quite subjective. This is often the case when a group may be ranked either as a class or as a subphylum. The organizational limits of some groups are also largely a matter of opinion. Some authors have placed the phylum Hemichordata within the Chordata, expressing the close genealogical relationship. Others prefer to keep them as a separate phylum because hemichordates lack what are considered important chordate features.Michael T. GhiselinThe Editors of Encyclopaedia Britannica

vertebrate

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vertebrate

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Also known as: Craniata, Vertebrata

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major vertebrate groups

Category: Animals & NatureKey People: Alfred Sherwood Romer Edward Drinker Cope Marie-Jean-Pierre Flourens Othniel Charles Marsh Sir John Graham KerrRelated Topics: reptile fish bird mammal chondrichthyan

vertebrate, also called Craniata, any animal of the subphylum Vertebrata, the predominant subphylum of the phylum Chordata. They have backbones, from which they derive their name. The vertebrates are also characterized by a muscular system consisting primarily of bilaterally paired masses and a central nervous system partly enclosed within the backbone.

The subphylum is one of the best known of all groups of animals. Its members include the classes Agnatha, Chondrichthyes, and Osteichthyes (all fishes); Amphibia (amphibians); Reptilia (reptiles); Aves (birds); and Mammalia (mammals).

General features

Although the vertebral column is perhaps the most obvious vertebrate feature, it was not present in the first vertebrates, which probably had only a notochord. The vertebrate has a distinct head, with a differentiated tubular brain and three pairs of sense organs (nasal, optic, and otic). The body is divided into trunk and tail regions. The presence of pharyngeal slits with gills indicates a relatively high metabolic rate. A well-developed notochord enclosed in perichordal connective tissue, with a tubular spinal cord in a connective tissue canal above it, is flanked by a number of segmented muscle masses. A sensory ganglion develops on the dorsal root of the spinal nerve, and segmental autonomic ganglia grow below the notochord. The trunk region is filled with a large, bilateral body cavity (coelom) with contained viscera, and this coelom extends anteriorly into the visceral arches. A digestive system consists of an esophagus extending from the pharynx to the stomach and a gut from the stomach to the anus. A distinct heart, anteroventral to the liver, is enclosed in a pericardial sac. A basic pattern of closed circulatory vessels is largely preserved in most living forms. Unique, bilateral kidneys lie retroperitoneally (dorsal to the main body cavity) and serve blood maintenance and excretory functions. Reproductive organs are formed from tissue adjacent to the kidneys; this original close association is attested by the tubular connections seen in males of living forms. The ducts of the excretory organs open through the body wall into a cloacal chamber, as does the anus of the digestive tract. Reproductive cells are shed through nearby abdominal pores or through special ducts. A muscular tail continues the axial musculature of the trunk.

Approximately 45,000 living species constitute the vertebrates. Species of several classes are found from the high Arctic or Antarctic to the tropics around the Earth; they are missing only from interior Antarctica and Greenland and from the North Polar ice pack. In size, vertebrates range from minute fishes to elephants and whales (of up to 100 tons), the largest animals ever to have existed. Vertebrates are adapted to life underground, on the surface, and in the air. They feed upon plants, invertebrate animals, and one another. Vertebrate faunas are important to humans for food and recreation.Britannica QuizWild Words from the Animal Kingdom Vocabulary Quiz

Natural history

In order to give a broad and comparative view of their life histories, the vertebrates are subdivided here into major groups based on morphology: the cyclostomes (jawless fishes), the chondrichthyes (cartilaginous fishes), the teleostomes (bony fishes), and the tetrapods.

The cyclostomes

Observe the hagfish's horny teeth and scaleless exterior covering a cartilaginous skeleton — Observe the hagfish’s horny teeth and scaleless exterior covering a cartilaginous skeletonSee all videos for this article

The cyclostomes include two classes of living, jawless fishes (agnathous)—Petromyzontiformes (lamprey eels) and Myxiniformes (hagfishes). The hagfishes are totally marine, often living in deep waters associated with muddy bottoms. The lampreys may be marine as adults but spawn in fresh waters, where the larvae spend some time before metamorphosing to the adult. Some lampreys live entirely in fresh water and may change only slightly in habit as a result of metamorphosis. Without lateral fins, lampreys swim by undulations of the body and can control direction only for short distances.

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The living agnaths are predatory, the lampreys being well known for attacking salmonoid fishes. The lamprey attaches to its prey using its round, suctorial mouth, and it rasps a hole through the outer tissues using a tongue armed with keratinized teeth. It suctions off bits of tissue, blood, and body fluids. The hagfishes feed somewhat similarly, but on a variety of prey—invertebrates (worms and soft-bodied forms) and dead fishes.

The lampreys produce small eggs, which develop directly into larvae that burrow into the muddy bottom of the stream. With its mouth at the surface of the mud, the larva filter feeds until large enough to metamorphose and swim off as a small adult. In contrast, the hagfishes produce relatively large encapsulated, yolky eggs up to two centimetres in length. When laid, these eggs attach to any available object by terminal hooks. The encased egg develops more or less directly into a miniature adult.

The chondrichthyes

southern stingray (Dasyatis americana or Hypanus americanus) — southern stingray (*Dasyatis americana* or *Hypanus americanus*)

The sharks, rays, and chimaerids are usually marine, but some sharks have entered fresh waters (the Amazon) or even live there permanently (Lake Nicaragua). In size, sharks range from the whale shark, nearly 10 metres in length, to rather small species, three centimetres in length. They usually weigh 25 to 200 kilograms (55 to 440 pounds). Sharks are predatory animals. Some large shark species (basking and whale sharks) filter feed on small crustaceans. Herbivorous sharks are unknown. Sharks swim by undulations of the tail, but rays “fly” through the water by undulations of the pectoral fins. Most species occur in near-shore waters, but some range widely throughout the oceans. A few are found in deep water.

A few sharks produce live young (viviparous) after internal fertilization. The posterior angle of the male’s pelvic fins are modified into a clasper, which acts as an intromittent organ in copulating with the female. Most sharks lay large yolky, encapsulated eggs with hooks for attachment. The young develop directly and begin life as miniature adults. The young that develop in the mother’s uterus obtain nutrients from the large yolk sac until they are born alive. In a few cases, the uterine wall secretes nutrients.

The teleostome, or osteichthyian, fishes (those having an internal bony skeleton) can be divided into two groups: the subclasses Actinopterygii (ray-finned fishes) and Sarcopterygii (lobe-finned fishes). The latter group includes the lungfishes, which live in marshes, ponds, or streams, and are frequent air breathers. They lay fairly large eggs, with a limited amount of yolk, that are enclosed in jelly coats like those of an amphibian. The eggs develop into small fishes that feed on live prey. The larvae of the African lungfish have external gills to supplement oxygen intake.

The teleostomes

American paddlefish (Polyodon spathula) — American paddlefish (*Polyodon spathula*)

Actinopterygian fishes are the common bony fishes of modern aquatic environments. They range in size from fishes that are only millimetres in size to those two or more metres (6.6 or more feet) in length, weighing 500 kilograms or more. Large species (sturgeons) are found in fresh waters (several other large species are found in the Amazon) as well as in marine environments. The diet may include plants, animals, and carrion. Most species are midwater swimmers, but many spend much time lying on the bottom. Tail, pectoral, and even dorsal fins are used in swimming. Reproduction in this group is by way of large numbers of small eggs, which produce small larvae or develop directly to the adult.

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The tetrapods

sandhill cranes (Grus canadensis) — sandhill cranes (*Grus canadensis*)

The tetrapods live primarily on land and are rather similar in habit. Members include the amphibians, reptiles, birds, and mammals. Amphibians are widespread in the warmer parts of the continents, being absent only in the far north and in the Antarctic. Three orders are recognized: Candata (the salamanders), the frogs and toads (Anura, or Salientia), and the Apoda or Gymnophiona (caecilians). Modification takes many forms, from the moist glandular skin (some scale remnants persist in apodans) to the loss of many of the bones of the skull. Like their ancestors, amphibians are cold-blooded and tend to be aquatic or limited to moist surroundings. Salamanders are seemingly the least modified in body form. They do not actively pursue prey and at best are only marginal swimmers. In swimming or crawling, the salamander’s body and tail undulate. Frogs and toads hop using hind-limb propulsion and the forelimbs as body props. This dominance of the hind limb in locomotion is best seen in swimming when the forelimbs are drawn back against the body. In contrast to the salamanders and frogs, the burrowing, wormlike apodans are without limbs.

Amphibians usually trap food using a tongue that can be shot out of the mouth, or they use the mouth itself to grasp and ingest food. There is great variation in foods; only the larvae of frogs and toads appear to be plant feeders, a specialization that is reflected in the highly modified jaws and guts of the tadpoles.

Amphibians have retained a simple egg cell with a gelatinous cover. The eggs are laid in ponds, streams, or even in damp places high in trees, usually in great numbers. Fertilized eggs develop into free-swimming larvae, which then metamorphose to adults, but in highly specialized forms.

The class Reptilia retains many of the structural characteristics of the ancestral amphibian. While most reptiles are carnivorous, feeding on other organisms, a few are herbivorous (e.g., tortoises). As cold-blooded animals, reptiles tend to be limited to temperate and tropical areas, but, where found, they are relatively common, although not as large or conspicuous as birds or mammals. Most reptiles are terrestrial, but a few are aquatic. As basic tetrapods, reptiles move about by creeping or swimming in a fashion similar to amphibians. Some reptiles, however, can lift the body from the ground and run rapidly either in a quadrupedal or bipedal fashion. Reptiles lay relatively large, shelled eggs. In a few instances, the eggs and young are cared for by the female; in others, the young are born alive (ovovivipary).

Birds are warm-blooded, and, although most are capable of flight, others are sedentary and some are flightless. Like their relatives the reptiles, birds lay shelled eggs that differ largely in the amount of calcification (hardening) of the shell. The young are usually cared for in a nest until they are capable of flight and self-feeding, but some birds hatch in a well-developed state that allows them to begin feeding immediately or even take flight. The megapods lay their eggs in mounds of rotting vegetation, which supplies the heat for incubation. (Nesting activities similar to those of some birds are seen in the crocodilians.)

The mammals range in size from tiny shrews or small bats weighing only a few grams to the largest known animals, the whales. Most mammals are terrestrial, feeding on both animal and vegetable matter, but a few are partially aquatic or entirely so, as in the case of the whales or porpoises. Mammals move about in a great variety of ways: burrowing, bipedal or tetrapedal running, flying, or swimming. Reproduction in mammals is usually viviparous, the young developing in the uterus, where nutritive materials are made available through an allantoic placenta or, in a few cases, a yolk sac. The fertilized egg develops directly into the adult. The monotremes (platypus and echidna) differ from other mammals in that they lay eggs which hatch, and the relatively undeveloped young are carried in a pouch or kept in a nest; the growing young lap up a milk nutrient fluid exuded from the belly of the mother.

Form and function

External features

The evolution of the notochord, dorsal nerve tube, and pharyngeal slits in chordate structure suggests improved swimming capability and probably greater ability to capture prey. Specialization in the vertebrate for the active capture of larger prey is evident both in the structure of the mouth and in the relatively simple structure of the pharynx, with its strong gill development. Specialization for feeding is again seen in the two basic groups of vertebrates, the agnathans and gnathostomes. Swimming adaptations are also numerous and involve variations both in body form and in medial fins and the two pairs of lateral fins.

Internal features

The skeletal system

Support and protection are provided by the exoskeletal and endoskeletal divisions of the skeletal system. The exoskeleton, when present, is basically protective but functions in tooth support in the mouth region. The endoskeleton protects the brain and spinal cord and assists primarily with locomotion in the trunk and tail regions. The endoskeleton begins as cartilage and may remain so or may develop into bone. The cartilaginous endoskeleton, found in the shark or chimaerid, is usually calcified so as to be stiffer and stronger. Bone is distinctive but highly variable; some types of bone contain cells, others do not, or the bone may be laminar, spongy, or arranged in sheathing layers around blood channels.

Tissues and muscles

Tissue development in the vertebrate is unique in its complexity; tissues in the strict sense (defined as a mass or sheet of similar cells with a similar function), however, do not exist. The simplest situation is seen in the epidermis, but even here there is a layered system in which different cell types provide different functions (such as protection and secretion). The stratified epithelium of the vertebrate is highly characteristic of that group (a similar one is seen in only one invertebrate group, the class Chaetognatha).

Other tissues of the vertebrate are more complex than the epithelium. For example, skeletal muscle consists not only of striated muscle fibres but also of connective tissue, which binds it together and attaches it by way of tendons. This contractible tissue includes nerves and blood vessels and their contained blood. Skeletal muscles thus appear as simple organs, just as do the smooth muscles in the wall of the gut or the iris muscles of the eye. Such unique histological complexity runs through the entire body of the vertebrate.

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Nervous system and organs of sensation

The dorsal position, tubular structure, and epidermal origin of the central nervous system are definitive of the chordates, although some may see similarities with the hemichordates. The sensory structures are distinctive of the chordates and include the paired nasal, optic, and otic organs (along with the strongly differentiated head).

The nasal vesicle is variously open to the environment, and its sensory cells, as chemical receptors, are not unlike those in the taste buds of the mouth. The eye is the most complex organ of the head and is a lateral outpocketing of the anterior end of the brain tube. Later it acquires a lens of epidermal origin. The act of focusing the eye (accommodation) shows extensive adaptive variation among the different groups of vertebrates.

The otic vesicle starts from a simple sac formed by the invagination of an ectodermal placode. These developmental changes also include the changes of innervation. Whereas the original structure was basically an equilibrium adaptation, other functions, such as an awareness of movement or the sensation of the proximity of prey, developed.

The lateral-line system of canals and sensory organs is a unique vertebrate feature. The elements of this system are found on the head as well as the body. This system is related to the ear and presumably at its origin served a similar function. This system is lost in terrestrial vertebrate forms.Britannica QuizAnimal Factoids

The digestive system

The digestive system of the vertebrate is distinctive in its structure but not in its function. The mouth and pharynx can be considered as parts of this system; the latter as an expanded cavity in the head is unmatched in any other group. The stomach and gut have been discussed above.

Presumably the original condition of the digestive glands was that of a ventral diverticulum which may have received the food mass into its cavity. This diverticulum, matched by the diverticulum seen in the amphioxus or the “intestine” of the tunicate, produced the secretions (bilelike and enzymes) of both liver and pancreas. Through time, the liver gradually differentiated from the pancreas. The size and separation of the liver from the gut suggest its separate blood and metabolic activities. The most obvious by-product of the liver, bile, necessitated the formation of a gall bladder and a duct connection with the gut. The pancreas, in contrast, continued to produce digestive enzymes, but its secretory cells were no longer in direct contact with the food mass. Because the pancreas was only a partial source of intestinal digestive enzymes, it was sometimes reduced in size and enclosed in the gut wall itself (agnaths) or dispersed as tiny bits of tissue in the mesentary supporting the gut (actinopterygians).

The excretory system

The excretory system is unique in its nephrons, which filter the blood in the glomeruli and remove a variety of wastes from the body through selective secretion and reabsorption. In the shark or the coelacanth Latimeria, urea is used to raise the osmotic pressure of the blood to that of the marine habitat, thus saving these organisms considerable metabolic energy. The large intestine (sometimes centred in a rectal gland) acts as an auxiliary excretory organ, as do also the gills of fishes or the sweat glands of mammals.

Respiration and gas exchange

Respiration, like excretion, involves specialized body structures, such as lungs or gills, but also can involve other areas, such as the skin itself. Respiration involves exchange of gases both between the body of the organism and the environment and between the blood system and the body tissues. It also involves cellular respiration where oxygen is used and carbon dioxide is produced. There is nothing characteristic of the vertebrate in this functional area; even the hemoglobin of the blood is suggested in the respiratory pigments of other animals.

The circulatory system

The circulatory system of vertebrates is closed in that fluids course through vessels, but there is free movement of cells in and out of blood. Some leukocyte (white blood cell) movement out of the capillaries and fluid leakage are observed in all tissues. Blood tissues are distinctive in the range of specialized cells, although these vary in detail among animals. The immune function of the blood is best developed in the vertebrate.

The endocrine system

The endocrine system is characterized by its separate organs. The occurrence of a pituitary or a thyroid gland is suggestive of the evolutionary change and specialization that took place within this group. The relatively unspecialized nature of some parts of this system is seen in certain scattered cells in the gut wall or even the clumps of islet cells of the pancreas.

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Evolution and paleontology

The knowledge of vertebrates as revealed by fossils has grown rapidly during the past few decades, but there is much still to be discovered. The ancestral vertebrate (protovertebrate) has been sought for more than 100 years, and the likelihood of finding it today is not much greater than in the past. It can be assumed that the protovertebrate was small and soft-bodied, two factors that suggest the improbability of finding a fossilized form in a recognizable condition. There are Cambrian fossils that have been suggested to be fossil cephalochordates and there are scales of agnath fishes, but the first type of fossil is too simple and the second already too complex to explain the transition.Malcolm T. Jollie

Classification

Annotated classification

SUBPHYLUM VERTEBRATA (OR CRANIATA)Bilaterally symmetrical; internal skeletal support with skull enclosing a highly developed brain and a vertebral column and nerve cord; paired, jointed appendages; skin; advanced organ systems; sense organs concentrated in head.
- Class Agnatha (hagfishes, lampreys)Primitive; jawless; paired fins are poorly developed or lacking; rasping tongue; notochord without bone; skin is soft, glandular, and slimy; true gill arches absent; marine habitat.
- Class Placodermi (placoderms)†Extinct; fishlike; jaws supported by both cranium and hyoid arch (amphistylic); partly ossified cranium; primitive; head and trunk have armour that is jointed at the neck; pelvic fins present or absent; pectoral fins or finlike structures often present; gill arches.
- Class Chondrichthyes (sharks, rays, and skates)Cartilaginous fishes; jaws; paired fins; no swim bladder; pelvic fins in males often modified to form claspers; gill arches internal to gills; reduced notochord; lateral-line system; paired nostrils; internal nares absent; separate sexes; internal fertilization and direct development; oviparous, ovoviviparous, or viviparous.
  - Subclass Elasmobranchii (sharks and rays)Numerous teeth derived of placoid scales; 5 to 7 gill clefts; operculum absent; cloaca; upper jaw not fused with braincase; dorsal fin nonerectile; with spiracles; worldwide distribution.
  - Subclass Holocephali (chimeras)Teeth fused to bony plates; no scales; 4 gill pairs under 1 gill opening on each side; no cloaca; no spiracles; operculum present; upper jaw fused to braincase; dorsal fin erectile; whiplike tail; claspers present in males; temperate marine freshwater.
- Class Osteichthyes (bony fishes)Jaws; partly or fully ossified skeleton; usually a swim bladder; paired fins; gills covered by a bony operculum; scales; paired nostrils with or without internal nares; lateral-line system; mostly oviparous with external fertilization; some ovoviviparous or viviparous.
  - Subclass Actinopterygii (ray-finned fishes)Generally lack choanae; no fleshy base to paired fins; no internal nares; air sacs usually function as swim bladder; skeleton usually well ossified.
  - Subclass Sarcopterygii (lobe-finned fishes)Usually possess a choana; paired fins with a fleshy base over a bony skeleton; persisting notochord; 2 dorsal fins; nares are internal.
- Class AmphibiaCold-blooded; respire by lungs, gills, skin, or mouth lining; larval stage in water or in egg; skin is usually moist with mucous glands and without scales; tetrapods; freshwater and terrestrial; paired appendages are legs; 10 pairs of cranial nerves; separate sexes; external fertilization with development into tadpole larvae; some have internal development, ovoviviparous or viviparous.
  - - Order Aponda (or Gymnophiona; caecilians)Wormlike; no limbs or girdles; compact skull; lidless, minute eyes; persistent notochord; tail; scales present in some species.
    - Order Anura (or Salientia; frogs and toads)Tailless; elongated hind limbs modified for jumping; larvae lack true teeth and external gills.
    - Order Caudata (or Urodela; salamanders)Tail; limbs normal; many skeletal elements cartilaginous; larvae with true teeth and external gills.
- Class ReptiliaCold-blooded; no larval stage; breathing by lungs; well-ossified skull; dry skin; scales; no glands; 5-toed limbs; claws; 3- or 4-chambered heart with incomplete ventricle separation; 12 pairs of cranial nerves; internal fertilization, direct development; oviparous and ovoviviparous.
  - Subclass Anapsida (turtles, tortoises, terrapins)No temporal skull openings; body encased in bony shell; no teeth in living members; oviparous.
  - Subclass LepidosauriaNo bipedal specializations; 2 complete temporal openings; complete palate; oviparous; male is without penis.
  - Subclass Archosauria (ruling reptiles)Some ancient forms had bipedal locomotion; longer hind legs; semiaquatic; webbed feet; teeth in sockets; single penis; oviparous; includes extinct dinosaurs.
  - Subclass Synaptosauria†Extinct; single temporal opening on area of cheek.
  - Subclass Ichthyopterygia†Extinct; temporal openings high up on skull; fishlike; spindle-shaped body; high tail fin; triangular dorsal fin; paddlelike legs; marine.
  - Subclass Synapsida†Extinct; mammal-like; lateral temporal opening.
- Class AvesWarm-blooded; skull has only 1 condyle; front limbs primarily modified for flight; hind limbs are legs with 4 or fewer toes; body covered with feathers; scales on feet; 4-chambered heart; no teeth; horny beak; lungs with extended air sacs; 12 pairs of cranial nerves; internal fertilization; oviparous.
  - Subclass Archaeornithes†Extinct; teeth in both jaws; long, feathered tail; less specialized for flight; body elongated and reptilelike; forelimb had 3 clawed digits; small brain and eyes; nonpneumatic bones.
  - Subclass Neornithes (true birds)Well-developed sternum; tail is not long; no teeth; forelimbs modified to wings; teeth replaced by horny rhamphoteca over bill.
- Class MammaliaWarm-blooded; mammary glands; lower jaw is composed of 1 bone; hair; advanced brain; skin with different glands and hair; ears with 3 middle-ear bones; 12 pairs of cranial nerves; 4-chambered heart; young nourished by milk from mammary gland; internal fertilization; mostly viviparous, some oviparous.
  - Subclass PrototheriaPrimitive; egg-laying; hair; mammary glands without nipples; pectoral girdle; separate oviducts that open into cloacal chamber that is shared with excretory ducts; oviparous.
  - Subclass TheriaMammary glands with nipples; functional teeth; oviducts partly fused; with or without a cloaca; uterus and vagina; viviparous.(Ed.)

Critical appraisal

The classification of animals is presently in a state of flux. The classification presented here is traditional and conservative. Because traditional theories of taxonomy tend to be nonquantitative, various interpretations of relationships or patterns can be presented and defended.

The alternative cladistic style of taxonomy is an attempt to force taxonomy into a testable, highly objective operation. One tentative classification based in cladistics separates the vertebrates into two superclasses (Agnatha and Gnathostomata). Agnathans are jawless, while the gnathostomates encompass the remainder of the jawed vertebrates. Living agnathans are placed in the class Cyclostomata. Gnathostomates can be further divided into the epiclasses Elasmobranchiomorphi (sharks and rays) and Teleostomi (bony fishes and tetrapods). The former group are identified primarily by a cartilaginous skeleton, while the latter group have developed a bony skeleton. Two subepiclasses of the teleostomes are Ichthyopterygii (or Osteichthyes; bony fishes) and Cheiropterygii (tetrapods), the latter being further divided into the classes Amphibia, Reptilia, Aves, and Mammalia.

Although this classification includes and uses traditional taxonomic categories, their position in the hierarchy may be changed. Separation of agnath and gnathostome is opposed by those cladists who chart the origin of gnathostomes from the agnath, believing that the differences in mouth and tooth structure are a result of modification. The Gnathostomata is subdivided into the Elasmobranchiomorphi and the Teleostomi largely on the basis of mouth and tooth structure. The creation of epiclasses and subepiclasses in the alternative classification is not important in itself; the creation of a dichotomy between Ichthyopterygii and Cheiropterygii, however, is important, although from the evolutionary view it is evident that the one evolved from the other.More From BritannicaWhy Do Animals Have Tails?Malcolm T. Jollie

psittaciform

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psittaciform

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Also known as: Psittaciformes, parrot

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Glen E. WoolfendenSee All

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Last Updated: Article History

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Black-capped parakeet (Pyrrhura rupicola

)

Category: Animals & NatureRelated Topics: parrot macaw cockatoo Loriinae pygmy parrot

Uncover interesting tidbits about the flightless kakapo, the African gray parrot, and blue and gold macawsSee all videos for this article

psittaciform, (order Psittaciformes), any member of the group of more than 360 species of generally brightly coloured noisy birds to which the general name parrot may be applied. All belong to just two families. In the family Psittacidae are parakeets (including the budgerigars, rosellas, and conures), lovebirds, amazons, macaws, and parrotlets (or parrolets), in addition to the lorikeets (including lories) as well as the kea and the kakapo of New Zealand. Members of the cockatoo family, Cacatuidae, live only in the region of Australia and New Guinea. This group also includes the cockatiel.

Parrots are primarily birds of the tropics. Their distribution encompasses the tropical and southern temperate regions of the world, including Madagascar, many Pacific Islands, and the West Indies. In Asia they occur throughout almost all of India but extend northward only to the Himalayas and southern China. They are absent from Europe. In North America one species, the thick-billed parrot (Rhynchopsitta pachyrhyncha), once ranged north into the extreme southwestern United States. Prior to the early 1900s, however, the Carolina parakeet (Conuropsis carolinensis) inhabited most of the eastern United States; it was rendered extinct by human persecution. The last captive died in the Cincinnati Zoological Garden in 1914, but the last generally accepted observation in the wild was a flock seen in Florida in 1920, although it has been claimed that they existed in South Carolina until 1938. In the Southern Hemisphere a number of parrots range to Tasmania and New Zealand, and in South America one species is found on Tierra del Fuego, and they are absent from parts of extreme southern Africa.

General features

Scarlet macaw (Ara macao). — Scarlet macaw (*Ara macao*).

Body plans of some smaller psittaciforms.

Body plans of some larger psittaciforms.

Parrots vary in total length from 8 to 100 cm (3 to nearly 40 inches), the latter in long-tailed forms such as macaws. The short neck and sturdy body, along with the stout feet and thick bill, give them a bulky appearance. The broad wings are often pointed; the tail is highly variable in both length and shape. In some species the tail is short and rounded or square; in others, such as the macaws, it is extremely long and pointed. In numerous species the central tail feathers are very long, surpassing the body in total length. In the five species of racquet-tailed parrots (Prioniturus), the central tail feathers are longer than the others and are spatulate, the middle part of the feather shaft being bare. No parrot has a forked tail. Pointed wings and a long tail usually are found in species that fly great distances; rounded wings and blunt tails typify the more adept climbers. Most parrots are swift on the wing, although they generally fatigue quickly.

Importance to humans

Lovebirds (Agapornis personata). — Lovebirds (*Agapornis personata*).

The first accurate written reference to a parrot is frequently credited to the Greek historian Ctesias, of the 5th century BC, who described clearly what is now called the blossom-headed parakeet (Psittacula cyanocephala) of India. However, there is no doubt that parrots were associated with humans much earlier, for natives on all continents have had parrots as pets. Parrots of many kinds have been long transported to zoos and private collections. Affluent citizens of early Rome often kept parrots in their homes and even esteemed them as delicacies of the dinner table.Britannica QuizWild Words from the Animal Kingdom Vocabulary Quiz

The qualities of parrots, especially the ability of many species to imitate human sounds, make them popular as pets. The African gray parrot (Psittacus erithacus) and some species of amazons (Amazona) from the New World tropics are particularly good mimics. There is no evidence to suggest, however, that talking parrots realize what they are saying. Another appealing attribute of parrots is their display of affection, not only to others of their own species but also to humans. Pairs of many species, especially the lovebirds (Agapornis), are together almost constantly, nibbling each other’s feathers with seeming affection; if one bird disappears, its mate sometimes dies, apparently of loneliness. Many parrots seem to delight in being petted and scratched, which is rare among birds. Parrots have extremely powerful jaws, however, and an indiscriminate attempt to pet them can result in a severe bite. The use of toes for climbing and food handling, in much the same manner as humans use hands, also makes parrots appealing. Their longevity, bright colours, intent gaze, ability to learn tricks, and willingness to remain on a perch instead of fluttering about contribute to the fondness people may feel for various kinds of parrots as pets. Finally, most species are vegetarians and thrive on a varied diet. This circumstance, as well as the fact that their droppings typically are dry and compact, means that parrots require little care.

Most species of parrots have been kept in captivity at one time or another, and most have been bred. A large zoological garden may have more than 100 species on display at one time. No parrot has been domesticated in the sense of chickenlike birds (order Galliformes) and waterfowl (order Anseriformes), although breeders have produced a variety of colour strains of the budgerigar (Melopsittacus undulatus), also called the shell parakeet. In the mid-1950s “budgies” became popular household pets in the United States; within a decade more than 5.5 million people had at least one in their homes. Captive parrots, especially the larger species, are long-lived. Claims of 80 or even 100 years are frequent and perhaps true, although thus far they have been impossible to document.

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