ROUTES OF MIGRATION

While it is beyond question that certain general directions of flight are constantly followed by migratory birds, it is well to remember that the term "migration route" is to some extent a theoretical concept, concerned entirely with the lines of general advance or retreat of a species, rather than the exact course followed by individual birds. Even the records of banded birds usually show no more than the places of banding and recovery, and one must have recourse to intermediate records and to reasoning based on probabilities to fill in details of the flyway actually traversed between the two points.

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, in time of starting, in speed of flight, in geographical position, in latitudes of breeding and of wintering grounds, and in other factors, all contribute to this great variation of migration routes. Nevertheless, there are certain factors that serve to guide the avian travelers along more or less definite lines, and it is possible to define general lines of migration for the majority of species.

It has frequently been observed that migrating birds have a tendency to follow major topographic lines on the earth's surface when their trend is in the general direction of the birds' journey. Bird migration is generally thought of as a north-and-south movement, with the lanes of heavier concentration following the coasts, mountain ranges, and principal river valleys. To a considerable extent this is the case, particularly in North America, where the coast lines, mountain chains, and the larger rivers in general run north and south. Students of American birds thus have exceptionally good opportunities to study migratory movements. In cases where the migration is a long one, however, the notion must be abandoned that the birds' flight is restricted to particular narrow routes that follow river valleys and the like, as many species seem to disregard utterly such apparently good natural flyways as river valleys. For example, the Arkansas River has a general east and west course for a great part of its length, and while it does constitute a highway for many perching birds en route from the Mississippi Valley to the Rocky Mountain region, some of the hawks and many ducks and shore birds pay the valley scant attention. They may arrest their fall journey to feed among cottonwoods or along sand bars, but when ready to resume their flight they leave the river and fly directly south over the more or less arid region that lies between the Arkansas and the Rio Grande.

WIDE AND NARROW MIGRATION LANES

When birds start their southward migration, the movement necessarily involves the full width of the breeding range. Later, there is a convergence of the lines of flight taken by individual birds, owing to the conformation of the land mass, and as the species proceeds southward the width of the occupied region becomes less and less. An example of this is provided by the common kingbird (Tyrannus tyrannus), which breeds from Newfoundland to British Columbia, a summer range 2,800 miles wide. On migration, however, its paths converge, 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, and still farther south the migration path is further restricted. In the latitude of Yucatan it is not more than 400 miles wide, and it is probable that the great bulk of the species moves in a belt that is less than half that width.

A migration route, therefore, may be anything from a narrow path that adheres closely to some definite geographical feature, such as a river valley or a coast line, to a broad boulevard that leads in the desired direction and follows only the general trend of the land mass. Also it is to be remembered that whatever main routes are described, there remain a multitude of tributary and separate minor routes. In fact, with the entire continent of North America crossed by migratory birds, the different groups or species frequently follow lines that may repeatedly intersect those taken by others of their own kind or by other species. The arterial routes, therefore, must be considered merely as indicating paths of migration on which the tendency to concentrate is particularly noticeable.

In considering the width of migration lanes it will be obvious that certain species, as the knot (Calidris canutus) and the purple sandpiper (Arquatella maritima), which are normally found only along the coasts, must have extremely narrow routes of travel. They are limited on one side by the broad waters of the ocean and on the other by land and fresh water, both of which are unsuited to furnish the food that is desired and necessary to the well being of these species.

Among land birds that have a definite migration, the Ipswich sparrow (Passerculus princeps) has what is probably the most restricted migration range of any species. It is known to breed only on Sable Island, Nova Scotia, and it winters along the Atlantic coast south to Georgia. Living constantly within sound of the surf, it is rarely more than a quarter of a mile from the outer beach, and is entirely at home among the sand dunes and their sparse covering of coarse grass.

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Figure 16.—Breeding and wintering ranges and migration of Harris's sparrow, an example of a narrow migration route through the interior of the country. The heavy broken lines enclose the region traversed by the majority of these finches; the light broken line encloses the country where they occur with more or less regularity; while the spots indicate records of accidental or sporadic occurrence.

Harris's sparrow (Zonotrichia querula) supplies an interesting example of a narrow migration route in the interior of the country (fig. 16). This fine, large finch is known to breed only in the region from Fort Churchill, on the west shore of Hudson Bay, northwest to the shores of Great Bear Lake. Very few actual breeding records of the species are available, but these are sufficient to indicate that the breeding range is in the strip of country characterized by more or less stunted timber just south of the limit of trees. When it begins its fall migration, this bird necessarily covers the full width of its breeding area. Then it proceeds almost directly south, or slightly southeasterly, the area covered by the majority of the species becoming gradually constricted, so that by the time it reaches the United States it is most numerous in a belt about 500 miles wide, extending across North Dakota to central Minnesota. Harris's sparrows are noted on migration with fair regularity east to the western shore of Lake Michigan, and west to the foothills of the Rocky Mountains, but the great bulk of the species moves north and south through a relatively narrow path in the central part of the continent. Present knowledge suggests that the reason for this narrow migration range is the close association that Harris's sparrow maintains with a certain type of habitat, including brushy places, thickets, edges of groves, and weed patches. While these environmental conditions are found in other parts of the country, the region crossed by this sparrow presents almost a continuous succession of habitat of this type. Its winter range extends from southeastern Nebraska and northwestern Missouri, across eastern Kansas and Oklahoma and through a narrow section of central Texas, at places hardly more than 150 miles wide.

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Figure 17.—Distribution and migration of the scarlet tanager. During the breeding season individual scarlet tanagers may be 1,900 miles apart in an east-and-west line across the breeding range. In migration, however, the lines converge until in southern Central America they are not more than 100 miles apart. For migration paths of other widths see figures 16, 18, and 19.

The scarlet tanager presents another extreme case of narrowness of migration route (fig. 17), its breeding range extending in greatest width from New Brunswick to Saskatchewan, a distance of about 1,900 miles. As the birds move southward in 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. Continuing to converge through Honduras and Costa Rica, the boundaries there are not more than 100 miles apart. The species winters in northwestern South America, where it spreads out over most of Colombia, Ecuador, and Peru.

The rose-breasted grosbeak also leaves the United States through the 700-mile stretch from eastern Texas to Appalachicola Bay, but thereafter the lines do dot further converge, as this grosbeak enters the northern part of its winter quarters in Central America and South America through a door of about the same width (fig. 18).

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Figure 18.—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 700 miles apart when the birds leave the United States. For migration paths of other widths see figures 16, 17, and 18.

While the cases cited represent extremes of convergence, a narrowing of the migratory path is the rule to a greater or less degree for the majority of North American birds. The shape of the continent tends to effect this, and so the width of the migration route in the latitude of the Gulf of Mexico is usually much less than in the breeding territory.

The redstart (Setophaga ruticilla) represents a notable case of a wide migration route, although even in the southern United States this is much narrower than the breeding range (fig. 19). These birds, however, cross all parts of the Gulf of Mexico and pass from Florida to Cuba and Haiti by way of the Bahamas, so that here their route has a width of about 2,500 miles.

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Figure 19.—Distribution and migration of the redstart. An example of a wide migration route, since birds of this species cross all parts of the Gulf of Mexico, or may travel from Florida to Cuba and through the Bahamas. Their fly way thus has an east and west width of more than 2,000 miles. For migration paths of greater or less extent see figures 16, 17, and 18.

In the following, the discussion of the principal routes of North American birds relates chiefly to the fall migration, for, except as otherwise noted, the spring flight generally retraces the same course. The routes indicated on the maps (figs. 20 and 21) must not be considered as representing paths with clearly defined borders, but rather as convenient subdivisions of the one great flyway that covers practically the entire width of the North American Continent and extends from the Arctic coast to South America.

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Figure 20.—Principal migration routes used by birds in passing from North America to winter quarters in the West Indies, Central America, and South America. Route no. 4 is the one used most extensively; only a few species make the 2,400-mile flight from Nova Scotia to South America.

ATLANTIC OCEANIC ROUTE

By reference to figure 20 it will be noted that route no. 1 is almost entirely oceanic, passing directly over the Atlantic Ocean from Labrador and Nova Scotia to the Lesser Antilles, and then through this group of small islands to the mainland of South America. It is not used by any of the smaller land birds, but is followed chiefly by thousands of water birds and by shore birds of several species, the adult golden plover being a notable example. Since it lies entirely over the sea, this route is definitely known only at its terminals and from occasional observations made on Bermuda and other islands in its course. Some of the shore birds that breed on the Arctic tundras of Mackenzie and in Alaska fly southeastward across Canada to the Atlantic coast and finally follow the oceanic route to the mainland of South America. The golden plover may accomplish the whole 2,400 miles without pause or rest, in fair weather the flocks passing Bermuda and sometimes even the islands of the Antilles without stopping. Although most birds make their migratory flights either by day or by night, the golden plover in this remarkable journey flies both day and night. As it swims lightly and easily it may make a few short stops along the way, and it has been actually seen resting on the ocean. Other shore birds have been observed busily feeding in that great area of ocean known as the Sargasso Sea, where thousands of square miles of floating seaweed teem with marine life.

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Figure 21.—Migration routes of North American birds. Though this map was prepared chiefly to show the flyways used by waterfowl, most of these routes also are utilized by innumerable land birds. For example, the important Mackenzie Valley-Great Lakes-Mississippi Valley route is shown (with its tributaries) from the Arctic coast to the delta of the Mississippi River.

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Figure 22.—Distribution and migration of the golden plover, Pluvialis dominica. Adults of the eastern form (P. d. dominica) migrate across northeastern Canada and then by a nonstop flight reach South America. In spring they return by way of the Mississippi Valley. Their entire route is therefore in the form of a great ellipse with a major axis of 8,000 miles and a minor axis of about 2,000 miles. The Pacific golden plovers (P. d. fulva), which breed in Alaska, apparently make a nonstop flight across the ocean to Hawaii, the Marquesas Islands, and the Low Archipelago, returning in spring over the same route.

The annual flight of the adult golden plover is so wonderful that it may be given in some detail, particularly since it is one of the exceptions to the general rule that spring and fall movements are over the same routes (fig. 22). After reaching the South American coast the birds make a short stop and then continue overland to the pampas of Argentina, where they remain from September to March. Leaving their winter quarters, they cross northwestern South America and the Gulf of Mexico, reaching the North American mainland on the coasts of Texas and Louisiana. Thence they proceed slowly up the Mississippi Valley, and by the early part of June are again on their breeding grounds, having performed a round-trip journey in the form of an enormous ellipse with the minor axis about 2,000 miles and the major axis 8,000 miles, reaching from the Arctic tundras to the pampas of Argentina. The older birds are probably accompanied by some of the young, perhaps those from early nestings, but most of the immatures leave their natal grounds late in summer and move southward through the interior of the country, re turning in spring over essentially the same course. The elliptical route is therefore used chiefly by fully adult birds.

ATLANTIC COAST ROUTE AND TRIBUTARIES

The Atlantic coast is a regular avenue of travel, and along it are many famous points for observing both land and water birds. About 50 different kinds of land birds that breed in New England follow the coast southward to Florida and travel thence by island and mainland to South America (fig. 20, route 2). As will be seen from the map, a seemingly natural and convenient highway extends through the Bahamas, Cuba, Hispaniola, Puerto Rico, and the Lesser Antilles to the South American coast. Resting places are afforded at convenient intervals, and at no time need the aerial travelers be out of sight of land. It is not, however, the favored highway, and only about 25 species of birds go beyond Cuba to Puerto Rico along this route to their winter quarters, while only 6 species are known to reach South America by way of the Lesser Antilles. The obvious drawback is lack of adequate food. The total area of all the West Indies east of Puerto Rico is less than that of Rhode Island, so that if only a small part of the birds of the eastern United States were to travel this way, it is doubtful whether even the luxuriant flora and fauna of tropical habitats would provide food sufficient for their needs.

In the northern part of the Atlantic coast route is a tributary route used by the brant (Branta bernicla) that is of special interest. The southward movement of these birds is chiefly along the western shores of Hudson Bay and thence southeastward to the Atlantic coast. Returning in spring, they follow the coast line north to the Gulf of St. Lawrence and then fly almost due north to their breeding grounds on the west coast of Greenland and the islands of the Arctic archipelago. The round trip is therefore in the form of a great ellipse, probably 3,000 miles long by 1,000 miles wide.

The Atlantic flyway receives accretions of waterfowl from three or four interior migration paths, one of which is of first importance, as it includes great flocks of canvasbacks (Nyroca valisineria), redheads (N. americana), scaup ducks (N. marila and N. affinis), Canada geese, and many of the black ducks that winter in the waters and marshes of the coastal region south of Delaware Bay. The canvasbacks, redheads, and scaups come from their breeding grounds on the great northern plains of central Canada, follow the general southeasterly trend of the Great Lakes (fig. 21), cross Pennsylvania over the mountains, and reach the Atlantic coast in the vicinity of Delaware and Chesapeake Bays. Black ducks, mallards, and blue-winged teals (Querquedula discors) that have gathered in southern Ontario during the fall leave these feeding grounds and proceed southwest over a course that is apparently headed for the Mississippi Valley. Many do continue this route down the Ohio Valley, but others, upon reaching the vicinity of Lake St. Clair, between Michigan and Ontario, swing abruptly southeast and, crossing the mountains in a single flight, reach the Atlantic coast south of New Jersey. This route, with its Mississippi Valley branch, has been fully demonstrated by the recovery records of ducks banded by a cooperator of the Biological Survey at Lake Scugog, Ontario.

The white-winged scoter (Melanitta deglandi), which also breeds in the interior country from northern North Dakota north to the Arctic coast, is another bird having an elliptical migration route, so far as those wintering on the Atlantic coast are concerned. This duck breeds only near fresh water and winters entirely on the ocean along both the Atlantic and Pacific coasts of the United States. Those wintering on the Atlantic side leave their breeding grounds west of Hudson Bay and fly 1,500 miles almost due east to the most eastern part of Labrador, whence they proceed southward across the Gulf of St. Lawrence to their winter home, which extends from southwestern Maine to Chesapeake Bay. The spring flight is made by an interior route that traverses the valleys of the Connecticut, Hudson, and Ottawa Rivers, and thence passes west and north to the breeding grounds.

A study of the Canada geese that winter abundantly in the waters of Back Bay, Va., and Currituck Sound, N. C., reveals another important tributary to the Atlantic coast route. Banding has shown that the principal breeding grounds of these birds are among the islands and on the eastern shores of Hudson Bay (fig. 21). From this region they move south in fall to the point of lower Ontario between Lakes Erie and Huron. Some of the banded geese are recovered in the Mississippi Valley, but the great majority are retaken either on their breeding grounds or on the Atlantic coast south of Delaware Bay, showing another instance of a long cross-country flight by waterfowl. Although Canada geese are abundant in migration on the coast of New England, the birds taken there do not include any that were banded in southern Ontario. Again, banding has shown that the New England visitants come from other breeding areas, chiefly Newfoundland and the desolate coast of Labrador, and that their migration is entirely coastwise.

Still another cross-country route between the Mississippi Valley and the Atlantic coast may be briefly described. While not yet well understood, a hitherto unsuspected migration route across the Alleghenies to the Mississippi Valley has been revealed by the banding of blue-winged teal, on the coastal sawgrass marshes of South Carolina. Birds marked in these marshes have been retaken in Tennessee and Kentucky as well as in States farther north in the Mississippi Valley. Several species of shoal-water ducks, including this dainty little teal and the shoveler (Spatula clypeata), are more or less common winter residents in the South Carolina marshes, but are less common or even decidedly rare in most of the coastal marshes farther north, so this cross-country route connecting two main arteries of migration seems to be of considerable importance.

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Figure 23.—Distribution and migration of the bobolink. In crossing to South America most of the bobolinks use route no. 3 (fig. 20), directly from Jamaica across an unbroken stretch of ocean. Colonies of these birds have established themselves in several areas in the western United States, but in migration they adhere to the ancestral flyways and show no tendency to take the short cut across Arizona, New Mexico, and Texas.

Referring again to figure 20, it is noted that route no. 3 presents a much more direct line of flight for the Atlantic coast migrants to South America than the others, although it involves much longer flights. It is used almost entirely by land birds. After taking off from the coast of Florida the migrants find only two land masses on the way where they can pause for rest and food. Nevertheless, tens of thousands of birds of some 60 species cross the 150 miles from Florida to Cuba, where about half of them elect to remain for the winter. The others fly the 90 miles between Cuba and Jamaica. From that point to the South American coast, however, there is a stretch of unbroken ocean fully 500 miles across, and scarcely a third of the North American migrants leave the forested mountains of Jamaica to risk the perils of this ocean trip. Chief among those that do is the bobolink (Dolichonyx oryzivorus), which so far outnumbers all other birds using this fly way that route no. 3 may well be called "the bobolink route" (fig. 23). As traveling companions along this route the bobolinks may meet vireos, kingbirds, and nighthawks from Florida; the chuck-will's-widow (Antrostomus carolinensis) of the Southeastern States; black-billed and yellow-billed cuckoos (Coccyzus erythropthalmus and C. americanus) from New England; gray-cheeked thrushes from Quebec, bank swallows (Riparia riparia) from Labrador; and blackpoll warblers from Alaska. Sometimes this scattered assemblage will be joined by a tanager or a wood thrush but "the bobolink route" is not popular with the greater number of migrants, and although many individuals traverse it, they are only a small fraction of the multitudes of North American birds that spend the winter in South America.

MACKENZIE VALLEY-GREAT LAKES-MISSISSIPPI VALLEY ROUTE AND TRIBUTARIES

Easily the longest flyway of any in the Western Hemisphere is that extending from the Mackenzie Valley past the Great Lakes and down the Mississippi River, including its tributaries. Its northern terminus is on the Arctic coast in the regions of Kotzebue Sound, Alaska, and the mouth of the Mackenzie River, while its southern end lies in Patagonia (fig. 21). During the spring migration some of the shore birds traverse the full extent of this great path, and it seems likely that the nighthawk, the barn swallow, the blackpoll warbler, and individuals of several other species that breed north to Yukon and Alaska must twice each year cover the larger part of it.

For more than 3,000 miles—from the mouth of the Mackenzie to the Delta of the Mississippi—this flyway is uninterrupted by mountains. In fact, there is not even a ridge of hills on the route high enough to interfere with the movements of the feathered travelers, and the greatest elevation above sea level is less than 2,000 feet. Well timbered and watered, the entire region affords ideal conditions for the support of its great hosts of migrating birds. This route is followed by such vast numbers of ducks, geese, shore birds, blackbirds, sparrows, warblers, and thrushes, that observers stationed at favorable points in the Mississippi Valley during the height of migration can see a greater number of species and individuals than can be noted anywhere else in the world.

Starting in the region of Kotzebue Sound, Alaska, the route extends eastward across northern Alaska and joins another that has its origin at the mouth of the Mackenzie River (fig. 21). The line of flight then trends a little east of south through the great lake system of central Canada, where it is joined by 2 or 3 other routes from the northeast that have their origin on the central Arctic coast. Continuing southward the migrating flocks are constantly augmented by additions to their numbers as they pass over the great breeding grounds of central and southern Canada. Upon reaching the headwaters of the Missouri and Mississippi Rivers the route follows these streams to the Gulf coast. Arriving in this latitude many species, including ducks and geese, the robin, the myrtle warbler, and some others spread out east and west for their winter sojourn. Others, despite the perils of a trip involving a flight of several hundred miles across the Gulf of Mexico, strike out boldly for Central America and South America. This part of the route is a broad "boulevard" extending from northwestern Florida to eastern Texas and reaching southward across the Gulf of Mexico to Yucatan and the Isthmus of Tehuantepec (fig. 20, route 4).

Many of the birds that breed east of the Allegheny Mountains parallel the sea coast as they move southwestward in fall and, apparently maintaining the same direction from northwestern Florida, cross the Gulf to the coastal regions of eastern Mexico. On the other hand, the birds that have come south directly through the Mississippi Valley and the region west to the Rocky Mountains reach the coastal plains of Mississippi, Louisiana, and Texas and continue directly across the Gulf. The great majority of North American birds seeking winter homes in the Tropics thus elect the short cut across the Gulf of Mexico in preference to the longer although safer land or island journey by way of Texas or Florida. During the height of the migration some of the islands off the coasts of Louisiana and Texas are wonderful observation places for the student of birds, as the feathered travelers literally swarm over them.

One of the short cuts (fig. 20, route 5) that may be considered a part of this great water artery of migration extends a few hundred miles from the coast of Texas to the northern part of the State of Vera Cruz. As the neighboring coast is arid and thus entirely unsuited to the needs of birds that are frequenters of moist woodlands, it is not surprising to find that this Gulf route is used by such woodland species as the golden-winged warbler (Vermivora chrysoptera), the worm-eating warbler (Helmitheros vermivorus), and the Kentucky warbler (Oporornis formosus).

Formerly it was thought that most of the North American birds that migrate to Central America made a leisurely trip along the Florida coast, crossed to Cuba, and thence made the short flight from the western tip of Cuba to Yucatan. A glance at the map would suggest this as a most natural route, but as a matter of fact it is practically deserted except for a few swallows and shore birds, or an occasional land bird storm-driven from its accustomed course.

Present knowledge of the chief tributaries to the Mackenzie Valley-Great Lakes-Mississippi Valley highway relates chiefly to waterfowl. Reference already has been made to the flight of the black ducks (p. 42) that reach the Mississippi Valley from southern Ontario. Some individuals of this species banded at Lake Scugog, Ontario, have been recaptured in succeeding seasons in Wisconsin and Manitoba, but the majority have been retaken at points south of the junction of the Ohio River with the Mississippi, definitely indicating their route of travel from southern Ontario.

A second route that joins the main artery on its eastern side is the one used by the blue goose (fig. 10), the migration route of which is probably more nearly due north and south than that of any other North American bird. The breeding grounds, which only recently have been discovered, are in the Fox Basin region of Baffin Island and on Southampton Island. In fall these geese work southward, chiefly along the eastern shore of Hudson Bay, and upon reaching the southern extremity of James Bay they take off for what is practically a nonstop flight to the great coastal marshes of Louisiana west of the delta of the Mississippi River (fig. 21). In some seasons the flocks make intermediate stops among the islands and sand bars of the Mississippi, as they are occasionally common in the general vicinity of Memphis, Tenn. Most of the birds push on, however, and during the period from the first of November to the last of March fully 90 percent of the species are concentrated in the area between the Sabine and the Mississippi Rivers. On the return trip northward there is sometimes a tendency for some of the blue geese to veer off toward the Northwest, as they are occasionally abundant in eastern South Dakota and southeastern Manitoba. It is of particular interest to note that while some other geese and many ducks start their northward journey at the first sign of awakening spring, the blue goose remains in its winter quarters until the season there is far advanced, seemingly aware that its own breeding grounds in the Arctic are still in the grip of winter.

As shown by the route map (fig. 21) a great western highway also has its origin in the Mackenzie River delta area and in Alaska. This is used chiefly by the pintail and the baldpate (Mareca americana), which fly southward through eastern Alberta to western Montana. Some localities in this area, as for example, the National Bison Range, at Moiese, Mont., normally furnish food in such abundance as to induce these birds to pause in their migratory movement. Upon resuming travel, some flocks move almost directly west across Idaho to the valley of the Columbia River, from which they turn abruptly south to the interior valleys of California. Others leave the Montana feeding and resting areas and turn southeastward across Wyoming and Nebraska to join the flocks that either are moving southward through the Great Plains, or are to continue across Arkansas to the main Mississippi Valley fly way.

Many redheads that breed in the Bear River marshes in Utah, take a westerly route across Nevada to California, but some leave these breeding grounds and fly northeastward across North Dakota and Minnesota to join the flocks of these ducks that come out of the prairie regions of Canada, and travel southeastward to the Atlantic coast. This route can be well traced by the records of ducks banded in summer in the Bear River marshes and retaken the following fall at points in eastern Montana, Wyoming, South Dakota, North Dakota, Minnesota, Wisconsin, Michigan, and Maryland.

Another route from these great marshes crosses the mountains in an easterly direction, but almost immediately turns southward through Colorado and New Mexico, and continues to winter quarters in the Valley of Mexico (fig. 21 and fig. 20, route 6). This route also represents the travels of many of the land birds of the Rocky Mountain region. Such birds perform comparatively short migrations, most of them being content to stop when they reach the middle districts of Mexico, only a few passing east beyond the southern part of that country.

This account of the Mackenzie Valley-Great Lakes-Mississippi Valley route shows the great importance of this highway and also its complicated nature. It receives accretions from both the northeast and the northwest, while branch routes make wide detours toward the Pacific coast before turning back to the parent fly way. This flyway is notable for its great length, as it extends from the Arctic-coast to Patagonia; and for its width east and west, as in North America it reaches from the Allegheny Mountains westward to the Great Basin.

PACIFIC COAST ROUTE

Although it does present features of unusual interest, the Pacific-coast route is not of so great importance as some of the others described. Because of the equable conditions that prevail, many species of birds along the coast from the Northwestern States to Southern Alaska either do not migrate at all or else make relatively short journeys. This route has its origin chiefly in Alaska, the general region of the delta of the Yukon River marking its northern terminus, although a few species join it after a flight westward along the Alaskan Arctic coast (fig. 21). Some of the scoters (Melanitta and Oidemia) and other sea ducks of the north Pacific region, and the diminutive cackling goose (Branta canadensis minima), which breeds in the delta of the Yukon River, use the coastal sea route for all or most of their southward flight. The journey of the cackling geese, as shown by return records from birds banded at Hooper Bay, Alaska, has been traced southward across the Alaska Peninsula and apparently across the Gulf of Alaska to the Queen Charlotte Islands, the birds following the coast line south to near the mouth of the Columbia River. There the route swings toward the interior for a short distance before continuing south by way of the Willamette River Valley. The winter quarters of the cackling geese are chiefly in the vicinity of Tule Lake, on the Oregon-California line, and in the Sacramento Valley of California, though a few push on to the San Joaquin Valley.

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Figure 24.—Probable breeding range, the winter range, and the migration route of Ross's goose. This is the only species of which all members apparently breed in the Arctic regions, migrate south through the Mackenzie Valley, and upon reaching the United States turn to the southwest rather than the southeast. The southern part of this route, however, is followed by some mallards, pintails, baldpates, and possibly by other ducks.

A tributary of this flyway is followed by Ross's goose (Chen rossi), which is believed to breed on the Arctic islands north of Mackenzie (fig. 24). Its fall migration is across the barren grounds to Great Slave and Athabaska Lakes, where it joins thousands of other waterfowl bound for their winter homes along the eastern coast of the United States and the Gulf of Mexico. But when Ross's geese have traveled south approximately to the northern boundary of Montana, they separate from their companions, and turning to the southwest cross the Rocky Mountains and settle for the winter in California.

The route taken by the white-winged scoters that winter on the Atlantic coast already has been indicated (p. 43). Some birds of this species, however, winter on the Pacific coast from Puget Sound south to southern California. Their passage by thousands up and down the coast has been noted as far north as northwestern British Columbia. The species is known to nest in Alaska, which may be the home of some at least of the scoters that winter on the Pacific coast. If such be the case, however, it must be admitted that a part of the route taken by the birds when on migration is unknown, though very few observations are available from the interior of northern British Columbia, across which the route may lie.

The southward route of those migratory land birds of the Pacific coast that in winter leave the United States extends chiefly through the interior of California to the mouth of the Colorado River and on to winter quarters in western Mexico.

The movements of the western tanager (Piranga ludoviciana) show a migration route that is in some ways remarkable. The species breeds in the mountains from the northern part of Baja California and western Texas north to northeastern British Columbia and southwestern Mackenzie. Its winter range is in two discontinuous areas—southern Baja California and eastern Mexico south to Guatemala (fig. 25). On the spring migration the birds enter the United States about April 20, appearing first in western Texas and the southern parts of New Mexico and Arizona (fig. 26). By April 30 the van has advanced evenly to an approximate east-and-west line across central New Mexico, Arizona, and southern California. But by May 10 the easternmost birds have advanced only to southern Colorado, while those in the far West have reached northern Washington. Ten days later the northward advance of the species is shown as a great curve, extending northeastward from Vancouver Island to central Alberta and thence southeastward to northern Colorado. Since these tanagers do not reach northern Colorado until May 20, it is evident that those present in Alberta on that date, instead of traveling northward through the Rocky Mountains, which from the location of their summer and winter homes would seem to be the natural route, reached there by the Pacific coast route to southern British Columbia and thence across the mountains, despite the fact that these are still partly covered with snow at that time.

PACIFIC OCEANIC ROUTE

The route of the Pacific golden plover (Pluvialis dominica fulva) is fully as interesting and as remarkable as the elliptical course followed by its eastern cousin (P. d. dominica) (fig. 22). The breeding range of the eastern golden plover extends through Arctic America west to the northern coast of Alaska, where in the vicinity of Point Barrow it meets the nesting grounds of the Pacific form, which is really an Asiatic subspecies. It breeds chiefly in the Arctic coast region of Siberia and merely overflows onto the Alaskan coast, some of the birds probably migrating south along the coast of Asia to winter quarters in Japan, China, India, Australia, New Zealand, and Oceania, including the Hawaiian Islands, the Marquesas Islands, and the Low Archipelago. Golden plovers in migration have been observed at sea on a line that apparently extends from these islands to the Aleutians, and it therefore appears certain that at least some of the Alaskan birds make a nonstop flight across a landless sea from Alaska to Hawaii. While it would seem incredible that any birds could lay a course so straight as to attain these small oceanic islands, 2,000 miles south of the Aleutians, 2,000 miles west of Baja California, and nearly 4,000 miles east of Japan, the evidence admits only the conclusion that year after year this transoceanic round-trip journey between Alaska and Hawaii is made by considerable numbers of golden plovers.

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Figure 25.—Breeding and wintering ranges of the western tanager. See figure 26 for the spring route taken by the birds breeding in the northern part of the range.

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Figure 26.—Migration of the western tanager. The birds that arrive in eastern Alberta by May 20 do not travel northward along the eastern base of the Rocky Mountains, as in that region the van has then only reached northern Colorado. Instead the isochronal lines indicate that they migrate north through California, Oregon, and Washington, and then cross the mountains of British Columbia.

The Pacific oceanic route probably is used also by the arctic terns that breed in Alaska, and possibly by those from the more western tern colonies of Canada. This species is of regular occurrence on the western coasts of both the United States and South America, indicating that the western representatives travel southward to the Antarctic winter quarters without the spectacular migration features that appear to characterize the flight of those from the eastern part of the continent (fig. 15).

ARCTIC ROUTES

In the discussion of the migration of the Arctic tern (p. 31) it was noted that this species makes a very distinct west-to-east movement across northern Canada, continuing the flight eastward across the Atlantic Ocean toward the western coast of Europe. It seems likely that there are other species, including the parasitic jaeger (Stercorarius parasiticus) that regularly breed in the northern part of the Western Hemisphere but migrate back to the Old World for their winter sojourn. Some others, as the red-legged kittiwake (Rissa brevirostris) and Ross's gull (Rhodostethia rosea), remain near the Arctic region throughout the year, retreating southward in winter only a few hundred miles. The emperor goose (Philacte canagica) in winter is found only a relatively short distance south of its breeding grounds, and eider ducks (Somateria and Arctonetta), although wintering in latitudes well south of the breeding grounds, nevertheless remain farther north than do the majority of other species of ducks (fig. 21).

The routes followed by these birds are chiefly coastwise, and in the final analysis may be considered as being tributary either to the Atlantic or to the Pacific coast routes. The passage of gulls, ducks, and other water birds at Point Barrow, Alaska, and at other points on the Arctic coast, has been noted by several observers, and from present knowledge it may be said that the best defined Arctic route in North America is the one that follows around the coast of Alaska.