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Migration of Birds

Routes of Migration

General Considerations
While it is beyond question that certain general directions of flight are consistently followed by migratory birds, it is well to remember the term "migration route" is to some extent a theoretical concept referring to the lines of general advance or retreat of a species, rather than the exact course followed by individual birds or a path followed by a species with specific geographic or ecological boundaries. Even the records of banded birds usually show no more than the place of banding and recovery. One ought to have recourse to intermediate records and reasoning based on probabilities to fill in details of the route actually traversed between the two points. In determining migration routes, one must constantly guard against the false assumption that localities with many grounded migrants are on the main path of migration and localities where no grounded migrants are observed are off the main path.

 There is also infinite variety in the routes covered during migration by different species. In fact, the choice of migration highways is so wide that it seems as if the routes of no two species coincide. Differences in distance traveled, time of starting, speed of flight, geographical position, latitudes of breeding and wintering grounds, and other factors contribute to this great variation of migration routes. Nevertheless, there are certain factors that serve to guide individuals or groups of individuals along more or less definite lines, and it is possible to define such lines of migration for many species.

 Except in a few species, individuals probably do not follow precisely the same route twice. This is especially true in the group of soaring birds that utilize thermals. Mueller and Berger (1967b) recaptured only three migrants in subsequent years at Cedar Grove, Wisconsin, after banding over 50,000 birds there. In general, those populations of species with very discernible breeding or wintering grounds have readily discernible migration routes. However, even the whole migration process of certain species may show great yearly fluctuations (Rudebeck 1950).

 Aldrich et al. (1949) showed from banding data great variation in migration patterns between species of waterfowl. In some species there was considerable diversity in direction of movement, not only of different breeding populations within a species but also for different individuals of the same breeding population. The impression is inescapable; waterfowl migration is even more complicated than originally supposed, and it is difficult to make generalizations with regard to migration pathways for even a single species let alone waterfowl in general.

Flyways and Corridors
Through plotting accumulated banding data in the 1930's, investigators became impressed by what appeared to be four broad, relatively exclusive flyway belts in North America. This concept, based upon analyses of the several thousand records of migratory waterfowl recoveries then available, was described by Lincoln (1935a). In this paper (p. 10), Lincoln concluded that:

. . . because of the great attachment of migratory birds for their ancestral flyways, it would be possible practically to exterminate the ducks of the West without seriously interfering with the supply of birds of the same species in the Altantic and Mississippi flyways, and that the birds of these species using the eastern flyways would be slow to overflow and repopulate the devasted areas of the West, even though environmental conditions might be so altered as to be entirely favorable.
Since 1948, this concept served as the basis for administrative action by the Fish and Wildlife Service in setting annual migratory waterfowl hunting regulations.

 The concept of bird populations being confined to four fairly definite and distinct migration "flyways" is probably most applicable to those birds that migrate in family groups, namely geese, swans, and cranes, but does not appear to be very helpful in understanding the movements of the more widely dispersing ducks. The "pioneering spirit" in Canada geese, for example, is limited by their social structure the young travel to and from specific breeding and wintering areas with their parents. These young later in life usually breed in the same areas as did their parents. If a goose population is decimated in one flyway, either by hunting or natural calamities, other goose populations in other flyways are not seriously endangered, but also these populations are very slow to repopulate an area where the previous goose population had been decimated. This is not the case with ducks because these birds are not always bound by their intrinsic behavior to return to specific breeding areas. Consequently, vacant breeding areas are more rapidly repopulated by ducks than by geese.

 Although Lincoln's analysis was confined to ducks and geese, some thought that it applied to other groups of birds as well. Everyone now realizes that the concept of four flyways, designated as the Atlantic, Mississippi, Central, and Pacific Flyways, was an oversimplification of an extremely complex situation involving crisscrossing of migration routes, varying from species to species. It can be considered meaningful only in a very general way, even for waterfowl, and not applicable generally to other groups of birds. Nevertheless the four "Flyway" areas have been useful in regionalizing the harvest of waterfowl for areas of different vulnerability of hunting pressure.

GIF - Migration corridors
Figure 13. Migration corridors used by dabbling ducks east of the Rocky Mountains during their fall migration (After Bellrose 1968). 

Bellrose (1968) identified corridors of southward migrating waterfowl east of the Rocky Mountains and determined, through statistical analyses, the relative abundance of birds in each. He showed major corridors of dabbling duck movements down the Great Plains and Missouri-Mississippi river valleys with minor offshoots at various points from these corridors eastward to the Atlantic coast where they joined equally minor eastern movements from the North (Fig. 13). Bellrose's map of migration corridors for the diving ducks showed heavy traffic similar to that of dabbling species down the Great Plains and relatively heavily used corridors from these central arteries eastward across the Great Lakes area to the Atlantic coast, terminating particularly in the vicinity of Chesapeake Bay. A fairly well-used corridor extends along the Atlantic coast.

GIF - Harris' sparrow distribution and migration
Figure 14. Distribution and migration of Harris' sparrow. This is an example of a narrow migration route through the interior of the country. 

With our present knowledge of bird migration it is difficult at best to recognize distinct broad belts of migration down the North American continent encompassing groups of distinct populations or species. It seems that so much intermingling of populations occurs that distinctions between broad "flyway" belts are not discernible. About all we can say for sure now is that birds travel between certain breeding areas in the North and certain wintering areas in the South and that a few heavily traveled corridors used by certain species, and more generalized routes followed by one or more species, have become obvious.

Narrow Routes
Some species exhibit extremely narrow routes of travel. The red knot and purple sandpiper, for example, are normally found only along the coasts because they are limited on one side by the broad waters of the ocean, and on the other by land and fresh water; neither of these habitats furnish conditions attractive to these species.

 The Ipswich race of the Savannah sparrow likewise has a very restricted migration range. It is known to breed only on tiny Sable Island, Nova Scotia, and it winters from that island south along the Atlantic coast to Georgia. It is rarely more than a quarter of a mile from the outer beach and is entirely at home among the sand dunes with their sparse covering of coarse grass.

 The Harris' sparrow supplies an interesting example of a moderately narrow migration route in the interior of the country (Fig. 14). This fine, large sparrow is known to breed only in the narrow belt of stunted timber and brush at or near the limit of trees from the vicinity of Churchill, Manitoba, on the west shore of Hudson Bay, to the Mackenzie Delta 1,600 miles to the northwest. When this sparrow reaches the United States on its southward migration, it is most numerous in a belt about 500 miles wide, between Montana and central Minnesota and continues south through a relatively narrow path in the central part of the continent. Knowledge of habitat preference by Harris' sparrows suggests the narrow migration range is restricted to the transition between woodland and prairie, a type of habitat approaching the woodland-tundra transition of its breeding area. Development of this migration route, of course, preceded destruction of the heavy eastern forests by colonists from Europe. Its winter range lies primarily in similar country extending from southeastern Nebraska and northwestern Missouri, across eastern Kansas and Oklahoma and through a narrow section of eastern Texas, at places hardly more than 150 miles wide.

Converging Routes
When birds start their southward migration the movement necessarily involves the full width of the breeding range. Later, in the case of landbirds with extensive breeding ranges, there is a convergence of the lines of flight taken by individual birds owing, in part, to the conformation of the land mass and in part to the east-west restriction of habitats suitable to certain species. An example of this is provided by the eastern kingbird, which breeds in a summer range 2,800 miles wide from Newfoundland to British Columbia. On migration, however, the area traversed by the species becomes constricted until in the southern part of the United States the occupied area extends from Florida to the mouth of the Rio Grande, a distance of only 900 miles. Still farther south the migration path continues to converge, and, at the latitude of Yucatan, it is not more than 400 miles wide. The great bulk of the species probably moves in a belt less than half this width.

 The scarlet tanager presents another extreme case of a narrowly converging migration route starting from its 1,900-mile-wide breeding range in the eastern deciduous forest between New Brunswick and Saskatchewan (Fig. 15). As the birds move southward in the fall, their path of migration becomes more and more constricted, until, at the time they leave the United States, all are included in the 600-mile belt from eastern Texas to the Florida peninsula. The boundaries continue to converge through Honduras and Costa Rica where they are not more than 100 miles apart. The species winters in the heavily forested areas of northwestern South America including parts of Colombia, Ecuador, and Peru.

GIF - Scarlet tanager distribution and migration
Figure 15. Distribution and migration of the scarlet tanager. During the breeding season individual scarlet tanagers may be 1,500 miles apart in an east-and-west line across the breeding range. In migration, however, the lines gradually converge until in South America they are about 500 miles apart. 

The rose-breasted grosbeak also leaves the United States through the 600-mile stretch from eastern Texas to Apalachicola Bay, but thereafter as this grosbeak crosses the Gulf of Mexico and enters the northern part of its winter quarters in southern Mexico the lines do not further converge. However, the pathway of those individuals that continue on to South America is considerably constricted by the narrowing of the land through Central America to Panama (Fig. 16).

GIF - Distribution and migration of the rose-breasted grosbeak
Figure 16. Distribution and migration of the rose-breasted grosbeak. Though the width of the breeding range is about 2,500 miles, the migratory lines converge until the boundaries are only about 1,000 miles apart when the birds leave the United States. 

Although the cases cited represent extremes of convergence, a narrowing of the migratory path is the rule to a greater or lesser degree for the majority of North American birds. Both the shape of the continent and major habitat belts tend to constrict southward movement so that the width of the migration route in the latitude of the Gulf of Mexico is usually much less than in the breeding territory.

 The American redstart represents a case of a wide migration route, but even in the southern United States, this is still much narrower than the breeding range (Fig. 17). These birds, however, cross all of the Gulf of Mexico and pass from Florida to Cuba and Haiti by way of the Bahamas so here their route is about 2,500 miles wide.

GIF - Distribution and migration of the redstart
Figure 17. Distribution and migration of the redstart. An example of a wide to migration route, birds of this species cross all parts of the Gulf of Mexico, or may travel from Florida to Cuba and through the Bahamas. Their route has an east-and-west width of more than 2,000 miles. 

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