The Migration of North American Birds (1935)
Part 6
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.
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, md 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.
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, continning 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.
=EVOLUTION OF MIGRATION ROUTES=
From the foregoing descriptions of migration routes it will be observed that the general trend of migration in most species of North American birds is northwest and southeast. It is comparatively easy to trace the probable steps in the evolution of the migrations of some species, and some routes have developed so recently that they still plainly show their origin.
The tendency is for eastern species to extend their ranges by pushing westward, particularly in the north. For example in the Stikine River Valley of northern British Columbia and southwestern Alaska the eastern nighthawk (_Chordeiles minor minor_), eastern chipping sparrow (_Spizella passerina passerina_), rusty blackbird (_Euphagus carolinus_), eastern yellow warbler (_Dendroica aestiva aestiva_), redstart, and others have established breeding stations at points 20 to 100 miles from the Pacific Ocean. The robin, flicker (Colaptes auratus), slate-colored junco (_Junco hyemalis hyemalis_), blackpoll warbler, myrtle warbler (Dendroica coronata), and ovenbird (_Seiurus aurocapillus_), all common eastern species, also are established as breeding birds in western Alaska, the ovenbird having been detected on the lower Yukon River. These birds, however, do not migrate in fall by any of the Pacific routes, but instead retrace their journey across the mountains and move southward along the broad flyways of the interior.
The red-eyed vireo (_Vireo olivaceus_), a striking example of an obundant woodland bird, is essentially an inhabitant of States east af the Great Plains, but an arm of its breeding range extends northwest to the Pacific coast in British Columbia (fig. 27). It seems evident that this is a range extension that has taken place comparatively recently by a westward movement from the upper Missouri Valley, and that the invaders retrace in spring and fall the general route by which they originally entered the country.
In the case of the bobolink, a new extension of the breeding range, and a consequent change in the migration of the species, has taken place since the spread of settlement in this country (fig. 23). A bird of damp meadows, it was originally cut off from the Western States by the intervening arid regions. But with the advent of irrigation and the bringing of large areas under cultivation, small colonies of nesting bobolinks have appeared at various western points, and now the species is established as a regular breeder in the great mountain parks and irrigated valleys of Colorado and elsewhere almost to the Pacific coast. In retracing their course to reach the western edge of the route followed by the bulk of the bobolinks that breed in the northern United States and southern Canada, these western pioneers must fly long distances along a line that runs almost due east and west.
Similarly it is possible to sketch what seems to be the logical evolution of the remarkable routes of the golden plover (fig. 22). It may be assumed that the eastern birds of this species first followed an all-land route from the South American winter quarters through Central America, Mexico, and Texas to the western parts of the Mississippi Valley. As the migration route lengthened northward with the retreat of the ice and the bird's powers of flight developed, it would have a tendency to straighten the line and to shorten it by cutting off some of the great curve through Mexico and Texas. First a short flight across the western part of the Gulf of Mexico was probably essayed. Proving successful, this was followed by flight lines that moved farther east, until finally the roundabout curve through Texas was entirely discarded and the flight made directly across the Gulf to southern Louisiana.
As the great areas in Canada were gradually added to the bird's domain, other important factors arose, the chief being the attractiveness of the vast stretches of coast and plain of the Labrador Peninsula, which in fall offered a bountiful store of berries. The fall route therefore worked eastward to the Gulf of St. Lawrence thence southwest through the interior to the coast of Florida and across the Gulf of Mexico to the Central American mainland. A series of shortening flights followed to take out the great curve of the New England coast. A relatively short ocean flight was probably attempted, say from Cape Cod to the Bahama Islands, Cuba, and Jamaica, followed eventually by the long direct oceanic route as it is now known.
As the Labrador Peninsula in spring is bound by frost and shrouded by fog while the season advances rapidly through the interior, the oceanic route proved useful only in fall, and the spring flight continued through the Mississippi Valley. The above outline gives a probable and fairly plausible explanation of the origin of this wonderful route, particularly when it is remembered that migration routes as now known are evolutions--age-long modifications of other routes.
The evolution of the migration of the Pacific golden plover may be explained in a similar fashion. At first the route probably followed the Asiatic coast, through the Malay Peninsula and Oceania, thence east in a great curve to the Low Archipelago, with individuals and flocks dropping out to winter at many points along the way. The Siberian birds probably continue to follow this ancient flyway, but those nesting in Alaska began a long evolutionary series of flights that cut down the length of their journey by shortening the curve, until finally the transoceanic route of the present day was developed.
=VERTICAL MIGRATION=
In the effort to find winter quarters furnishing satisfactory living conditions, many North American birds fly hundreds of miles across land and sea. Others, however, are able to attain their objective merely by moving down the sides of a mountain. In such cases a few hundred feet of altitude corresponds to hundreds of miles of latitude. Movements of this kind, known as "vertical migrations", are found wherever there are large mountain ranges. In the Rocky Mountain region they are particularly notable, as chickadees, rosy finches (_Leucosticte_), juncos, pine grosbeaks (_Pinicola_), and some other species that nest in the Alpine Zone move down to the lower levels to spend the winter. It has been noted that such species as Williamson's sapsucker (Sphyrapicus thyroideus), and the western wood pewee (_Myiochanes richardsoni_), which nest in the higher mountains, move down to the lower regions in August following the breeding season. At this time there is a distinct tendency also among the young of mountain-breeding birds to work down to the lower levels as soon as the nesting season is over. The sudden increases among birds in the edges of the foothills are particularly noticeable when cold spells with snow or frost occur at the higher altitudes.
Some species that normally breed in the Hudsonian or Arctic Zones find suitable breeding areas on the higher levels of the mountains, as for example the pipit, or titlark (_Anthus spinoletta rubescens_), which breeds on the tundras of Alaska and northern Canada and also south as far as Colorado on the summits of many peaks in the Rocky Mountains. On the other hand a few species, as the Clark's crow, or nutcracker (_Nucifraga columbiana_), nest at relatively low altitudes in the mountains and as the summer advances move higher up, thus performing a vertical migration that in a sense is comparable with the post-breeding movements of herons on the Atlantic coast. These illustrations show that the length of a migration route may depend upon factors other than latitude.
=VAGRANT MIGRATION=
The most striking feature of the migrations of some of the herons is a northward movement after the nesting season. The young of some species commonly wander late in summer and in fall, sometimes traveling several hundred miles north of the district in which they were hatched. The little blue heron (_Florida caerulea caerulea_) breeds commonly north to South Carolina, and by the last of July the young birds begin to appear along the Potomac, Patuxent, and Susquehanna Rivers, tributary to Chesapeake Bay. Although almost all are immature individuals, as shown by their white plumage, an occasional adult may be noted. With them come snowy herons (_Egretta thula thula_) and egrets (_Casmerodius albus egretta_), and on occasion all three species will travel in the East as far north as New England, and in the Mississippi Valley to southeastern Kansas and Illinois. In September most of them disappear, probably returning south by the same route.
The black-crowned night heron (_Nycticorax nycticorax hoactli_) has similar wandering habits, and young birds banded in a large colony at Barnstable, Mass., have been recaptured the same season north to Maine and Quebec and west to New York. This habit seems to be shared by some of the gulls also, although here the evidence is not so conclusive. Herring gulls (_Larus argentatus smithsonianus_) banded as chicks at colonies in the Great Lakes have scattered in all directions after the breeding season, some having been recovered well north in Canada.
These movements may be considered as migration governed only by the availability of food, and they are counteracted in fall by a directive migratory impulse that carries back to their normal winter homes in the south such birds as after the mating period have attained more northern latitudes. They are not to be compared with the great invasions of certain birds from the North. Classic examples of the latter in the eastern part of the country are the periodic flights of crossbills. Sometimes these migrations will extend well south into the Carolinian Zone.
Snowy owls are noted for occasional invasions that probably are caused by a shortage of the lemmings and rabbits that constitute their normal food in the North. At least nine notable flights of these birds occurred during the period 1876 to 1927. In the great flight of 1926-27 they were noted as far south as Iowa, Ohio, West Virginia, and North Carolina.
In the Rocky Mountain region great flights of the beautiful Bohemian waxwing (_Bombycilla garrula pallidiceps_), are occasionally recorded. The greatest invasion in the history of Colorado ornithology occurred in February 1917, at which time the writer estimated that at least 10,000 were within the corporate limits of the city of Denver. The last previous occurrence of the species in large numbers in that section was in 1908.
Evening grosbeaks (_Hesperiphona vespertina_) likewise are given to performing more or less wandering journeys, and curiously enough, in addition to occasional trips south of their regular range, they travel east and west, sometimes covering long distances. For example, grosbeaks banded at Sault Ste. Marie, Mich., have been recaptured on Cape Cod, Mass., and in the following season have been re trapped at the banding station. Banding records demonstrate that this east-and-west trip across the northeastern part of the country is sometimes made also by purple finches (_Carpodacus purpureus_).
=PERILS OF MIGRATION=
The period of migration is a season full of peril for birds. Untold thousands of the smaller migrants are destroyed each year by storms, in unfamiliar habitats, and through attacks of predatory birds, mammals, and reptiles. If each pair of adult birds should succeed in raising two fledglings to maturity, the population of migratory birds would have a potential annual increase of 100 percent and the world would soon be heavily overpopulated with them. It is evident, therefore, that there is no such increase, and that the annual mortality from natural causes is heavy enough to keep it in check.
STORMS
Of the various factors limiting the abundance of birds, particularly the smaller species, storms are the most potent. Special sufferers are those birds that in crossing broad stretches of water are forced by a storm down within reach of the waves. Such a catastrophe was once seen from the deck of a vessel in the Gulf of Mexico, 30 miles off the mouth of the Mississippi River. Great numbers of migrating birds, chiefly warblers, had accomplished nearly 95 percent of their long flight and were nearing land when, caught by a norther, against which they were unable to contend, hundreds were forced into the waters of the Gulf and drowned. On another occasion, on Lake Michigan, a severe storm, coming up at a time when large numbers of migratory birds were crossing, forced numerous victims into the waves. During the fall migration of 1906, when thousands of birds were crossing Lake Huron, a sudden drop in temperature, accompanied by a heavy snowfall, resulted in the death of incredible numbers. Literally thousands were forced into the water and subsequently cast up along the beaches, where in places their bodies were piled in windrows. On one section of the beach the dead birds were estimated at 1,000 per mile, and at another point at five times that number. Most of them were species that rank among our most desirable birds as destroyers of insects and weed seeds, including slate-colored juncos, tree sparrows, white-throated sparrows, swamp sparrows, winter wrens, and golden-crowned kinglets, together with many brown creepers, hermit thrushes, warblers, vireos, and others.