— In belated recognition of the spring migration season in the Northern Hemisphere, Advocacy for Animals is pleased to publish the following primer on bird migration, adapted from Encylopædia Britannica’s article “migration.”

Migration is most evident among birds. Most species, because of their high metabolic rate, require a rich, abundant supply of food at frequent intervals. Such a situation does not always prevail throughout the year in any given region. Birds have thus evolved a highly efficient means for travelling swiftly over long distances with great economy of energy.

The characteristics of migratory birds do not differ greatly from those of nonmigratory forms; many intermediate types exist between the two groups. All transitional forms, in fact, may be manifested in a single species or in a single local population, which is then said to undergo partial migration.

In addition to regular migration, nomadic flights may also occur. This phenomenon takes place, for example, among birds of the arid zones of Australia, where ducks, parakeets, and seedeaters appear in a locality following infrequent and unpredictable rains, breed, and then move to other areas. Nomadism is a response to irregular ecological conditions.

In Europe

The populations of many northern and eastern European species of birds have pronounced migratory tendencies; the populations of western Europe, on the other hand, are more sedentary. Some birds are nomadic in winter, others spend the colder months in the southwestern part of the continent or in the Mediterranean region. Many migrant populations migrate to Africa south of the Sahara. Geographical conditions determine several main routes. The Alps are an important barrier to migratory birds. About 150 species travel westward and southwestward; others travel southeastward.

Tits, goldfinches, and blackbirds are usually sedentary in western Europe; they are usually migratory, however, in northern Europe, where their flights resemble a short migration. Starlings are sedentary in western Europe, where large numbers gather from eastern Europe. Large flocks also pass the winter in North Africa.

Insectivorous (insect-eating) species, such as warblers, flycatchers, and wagtails, are highly migratory and spend the winter in the tropics, chiefly in Africa. They migrate to Sierra Leone on the west coast, Tanzania on the east coast, and all the way southward to the tip of the continent. Most of these migrants use different routes to cross the Mediterranean, chiefly in the western portion, although some migrate only southeastward. Golden orioles and red-backed shrikes go to East Africa by way of Greece and Egypt. Swallows—particularly barn swallows and house martins—and swifts pass the winter in Africa south of 20 degrees N latitude, particularly in South Africa, in the Congo River region, and in some coastal areas of West Africa.

Among nonpasserines—i.e., nonperching birds—one of the best-known migrants is the stork, which migrates to tropical Africa along two well-defined flyways. The stork population nesting west of a line that follows the Weser River in Germany flies southwestward through France and Spain, past the Strait of Gibraltar, and reaches Africa by way of West Africa; the eastern population, by far more numerous, takes a route over the straits of the Bosporus, through Turkey and Israel, to east Africa. These well-separated routes are probably a result of the stork’s aversion to long flights over water.

Ducks, geese, and swans also are migrants. These birds winter partly in western Europe and partly in tropical Africa. In Africa they are likely to spend the winter in lake and river regions from Senegal in western Africa to the Sudan in eastern Africa, where thousands of garganeys and pintails congregate annually. Some ducks leave their breeding grounds to molt (a process by which old feathers are replaced) in areas where they are most secure from predators during the time they are unable to fly; this is known as a molt migration. After molting, the ducks fly to their final winter quarters.

Wading birds (shorebirds) are typical migrants, most of them nesting in tundra of the Arctic region and wintering along the seacoasts from western Europe to South Africa. Scientists have observed that shorebirds, such as the white-rumped sandpiper, risk increased mortality from exhaustion and severe weather during the course of their long migrations. They suspect that this cost is balanced by the benefit of reduced nest predation; the Arctic tundra—a preferred shorebird breeding region—supports lower predator population densities than areas farther south, and thus greater numbers of newly hatched young survive to adulthood.

In North and South America

North American birds must endure the same hazards of winter as European species. The geographical arrangement of the continent determines the main routes of migration, which run from north to south and include the Atlantic oceanic route, the Atlantic Coast route, the Mississippi flyway, the central flyway, the Pacific flyway, and the Pacific oceanic route. A great many birds pass the winter in the Gulf States, but the principal wintering area extends through Mexico and Central America to Panama, which has the greatest density of winter bird residents in the world.

The ruby-throated hummingbird nests in southern Canada and winters in Central America as far south as Panama. Some of these birds fly nonstop across the Gulf of Mexico. Because of their food requirements, many American flycatchers, which are mainly insectivorous, have the same migratory behaviour as the hummingbirds. Others, like the phoebe, spend the winter in the Gulf States. Birds such as the American robin and several species of grackles assemble in the Gulf States in enormous flocks. The seasonal flights of the American wood warblers are among the most spectacular on the North American continent. Some spend the winter in the Gulf States and in the West Indies; others, such as the blackpoll warbler, travel to Guiana, Brazil, and Peru by way of the West Indies. The spring migration routes of the Canada goose span the Continent of North America in an east–west direction from Hudson Bay as far south as Chesapeake Bay.

South America is winter quarters for several tanagers, such as the scarlet tanager and the bobolink; these birds migrate through the eastern United States and past Cuba to the swampy regions of Bolivia, southern Brazil, and northern Argentina. This area of South America is also winter quarters for the American golden plover, which travels in an enormous loop over much of the New World. After nesting in the tundras of Alaska and Canada, the plover assemble in Labrador in easternmost Canada and then fly to Brazil over an oceanic route (the shortest possible route) about 3,900 kilometres (2,400 miles) long. Their return flight traverses South America, Central America, and the Gulf of Mexico, then follows the Mississippi Valley.

In intertropical regions

Birds of tropical regions migrate according to the rhythmic succession of wet and dry seasons—a profoundly influential factor on the annual cycle of animals and plants alike.

The migratory behaviour of birds has a unique regularity in Africa, where life zones are arranged symmetrically by latitudes away from the Equator. Some migrants never cross the Equator. The standard-wing nightjar, which nests in a belt extending from Senegal in the west to Kenya in the east along the equatorial forest, migrates northward to avoid the wet season. The common nightjar, on the other hand, nests in a dry belt from Mali in the west to the Red Sea and Kenya in the east during the rains and then migrates southward to Cameroon and the northern Congo region during the dry season.

Other birds migrate across the Equator to their alternate seasonal grounds. Abdim’s stork nests in a belt extending from Senegal to the Red Sea; after the wet season, it winters from Tanzania through most of southern Africa. The pennant-wing nightjar, in contrast, nests in the Southern Hemisphere south of the Congo forests during the austral, or Southern Hemisphere, summer, then starts north with the onset of the rainy season. It spends its winters in savannas from Nigeria to Uganda.

In coastal and pelagic regions

Among the migrating seabirds, a distinction must be made between the coastal and the pelagic, or open-sea, species. Birds such as guillemots, auks, cormorants, gannets, and gulls—all common to the seashore—stay in the zone of the continental shelf. Except during the breeding season, they are dispersed over a vast area, often preferring specific directions of travel. Gannets nesting around the British Isles spread in winter along the Atlantic coast of Europe and Africa to Senegal, the young travelling farther than the adults. Pelagic birds, most of which belong to the order Procellariiformes (petrels and albatrosses), cover much greater distances and, from a few small nesting areas, roam over a large part of the oceans.

Wilson’s petrels, which nest in the western sector of the Antarctic (South Georgia Island, Shetland Islands, and South Orkney Islands), spread rapidly northward in April along the coasts of North and South America and stay in the North Atlantic during the summer. In September they leave the western Atlantic, travelling east, then southeast, along the coasts of Europe and Africa toward South America and their Antarctic breeding grounds, arriving there in November. These petrels thus travel in a great loop through the whole Atlantic Ocean, in a flight pattern correlated with the direction of prevailing winds. The same pattern is used by other seabirds normally carried by the winds. Albatrosses, such as the wandering albatross that nests on small Antarctic islands, circle the globe during their migrations. One such bird, banded as a chick at Kerguelen Island in the southern Indian Ocean and recovered at Patache, Chile, travelled in less than 10 months at least 13,000 kilometres (8,100 miles)—perhaps as much as 18,000 kilometres (11,200 miles)—by drifting with the prevailing winds.

In the Pacific, short-tailed shearwaters nest in enormous colonies along the coasts of southern Australia and in Tasmania, then migrate across the western Pacific to Japan, remaining in the North Pacific and the Arctic Ocean from June to August. On the return migration they go east and southeast along the Pacific coast of North America, then fly diagonally across the Pacific to Australia.

Arctic terns, whose breeding range includes the northernmost coast of Europe, Asia, and North America, spend the winter in the extreme southern Pacific and Atlantic, chiefly along Antarctic pack ice 17,600 kilometres (11,000 miles) from their breeding range. American populations of the Arctic tern first cross the Atlantic from west to east, then follow the coast of western Europe. Arctic terns thus travel further than any other bird species.

Modes of migration

The migration flights of birds follow specific routes, sometimes quite well defined over long distances. The majority of bird migrants, however, travel along broad airways. A single population of migrants may be scattered over a vast territory so as to form a broad front hundreds of miles in width. Such routes are determined not only by geographical factors—e.g., river systems, valleys, coasts—and ecological conditions but are also dependent upon meteorological conditions; i.e., birds change their direction of flight in accordance with the direction and force of the wind. Some routes cross oceans. Small passerine (perching) birds migrate across 1,000 kilometres (620 miles) or more of sea in areas such as the Gulf of Mexico, the Mediterranean Sea, and the North Sea. American golden plover, wintering in the Pacific, fly directly from the Aleutian Islands (southwest of Alaska) to Hawaii, the 3,300-kilometre (2,050-mile) flight requiring 35 hours and more than 250,000 wing beats.

The speed of migratory flights depends largely on the species and the type of terrain covered. Birds in migration go faster than otherwise. Rooks have been observed migrating at speeds of 51 to 72 kilometres (32 to 45 miles) per hour; starlings at 69 to 78 kilometres (43 to 49 miles) per hour; skylarks at 35 to 45 kilometres (22 to 28 miles) per hour; and pintails at 50 to 82 kilometres (31 to 51 miles) per hour. Although the speeds would permit steadily flying migrants to reach their wintering grounds in a relatively short time, the journeys are interrupted by long stops, during which the birds rest and hunt for food. The redbacked shrike covers an average of 1,000 kilometres (620 miles) in five days as follows: two nights for migration, three nights for rest, five days for feeding.

Most migrations occur at relatively low altitudes. Small passerine birds often fly at less than 60 metres (200 feet). Some birds, however, fly much higher. Migrating passerines, for example, have been observed at altitudes as great as 4,000 metres (14,000 feet). The highest altitude recorded thus far for migrating birds is 9,000 metres (29,500 feet) for geese near Dehra Dun in northwest India.

Pelicans, storks, birds of prey, swifts, swallows, and finches are diurnal (daytime) migrants. Waterbirds, cuckoos, flycatchers, thrushes, warblers, orioles, and buntings are mostly nocturnal (nighttime) migrants. Studies of nocturnal migrants using radar on telescopes focussed on the Moon show that most migratory flights occur between 10 PM and 1 AM, diminishing rapidly to a minimum at 4 AM.

Most birds are gregarious during migration, even those that display a fierce individualism at all other times, such as many birds of prey and insectivorous passerines. Birds with similar habits sometimes travel together, a phenomenon observed among various species of shorebirds. Flocks sometimes show a remarkable cohesion; the most characteristic migratory formation of geese, ducks, pelicans, and cranes is a V with the point turned in the direction of flight.

Navigation

A compass sense has been demonstrated in birds; that is, they are able to fly in a particular constant direction, regardless of the position of the release point with respect to the bird’s home area. It has also been shown that birds are capable of relating the release point to their home area and of determining which direction to take, then maintaining that direction in flight. The navigational ability of birds has long been understood in terms of a presumed sensitivity to both the intensity and the direction of the Earth’s magnetic field. It has also been suggested that birds are sensitive to forces produced by the rotation of the Earth (Coriolis force); however, no sense organ or physiological process sensitive to such forces has yet been demonstrated to support this hypothesis.

Experiments have shown that the orientation of birds is based on celestial bearings. The Sun is the point of orientation during the day, and birds are able to compensate for the movement of the Sun throughout the day. A so-called internal clock mechanism in birds involves the ability to gauge the angle of the Sun above the horizon. Similar mechanisms are known in many animals and are closely related to the rhythm of daylight, or photoperiodism. When the internal rhythm of birds is disturbed by subjecting them first to several days of irregular light–dark sequences, then to an artificial rhythm that is delayed or advanced in relation to the normal rhythm, corresponding anomalies occur in the homing behaviour.

Two theories have been formulated to explain how birds use the Sun for orientation. Neither, however, has so far been substantiated with proof. One theory holds that birds find the right direction by determining the horizontal angle measured on the horizon from the Sun’s projection. They correct for the Sun’s movement by compensating for the changing angle and thus are able to maintain the same direction. According to this theory, the Sun is a compass that enables the birds to find and maintain their direction. This theory does not explain, however, the manner in which a bird, transported and released in an experimental situation, determines the relationship between the point at which it is released and its goal.

The second theory, proposed by British ornithologist G.V.T. Matthews, is based on other aspects of the Sun’s position, the most important of which is the arc of the Sun—i.e., the angle made by the plane through which the Sun is moving in relation to the horizontal. Each day in the Northern Hemisphere, the highest point reached by the Sun lies in the south, thus indicating direction; the highest point is reached at noon, thus indicating time. In its native area a bird is familiar with the characteristics of the Sun’s movement. Placed in different surroundings, the bird can project the curve of the Sun’s movement after watching only a small segment of its course. By measuring maximum altitude (the Sun’s angle in relation to the horizontal) and comparing it with circumstances in the usual habitat, the bird obtains a sense of latitude. Details of longitude are provided by the Sun’s position in relation to both the highest point and position it will reach—as revealed by a precise internal clock.

Migrant birds that travel at night are also capable of directional orientation. Studies have shown that these birds use the stars to determine their bearings. In clear weather, captive migrants head immediately in the right direction using only the stars. They are even able to orient themselves correctly to the arrangement of night skies projected on the dome of a planetarium; true celestial navigation is involved because the birds determine their latitude and longitude by the position of the stars. In a planetarium in Germany, blackcaps and garden warblers, under an artificial autumn sky, headed “southwest,” their normal direction; lesser whitethroats headed “southeast,” their normal direction of migration in that season.

It is known, then, that birds are able to navigate by two types of orientation. One, simple and directional, is compass orientation; the second, complex and directed to a point, is true navigation, or goal orientation. Both types apparently are based on celestial bearings, which provide a navigational “grid.”

Images: Barn swallow—© Vadim Andrushchenko/Fotolia; Blackpoll warbler—© Stubblefield Photography/Shutterstock.com; Male common nightjar—Frank V. Blackburn; Wandering albatross—Mark Jobling; Golden plover—Kenneth W. Fink/Root Resources; Bird migration at Eddystone Lighthouse, illustration by Charles Samuel Keene for “Punch”—Photos.com/Jupiterimages.