Chapter III: Part 3

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.

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, 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.

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.]

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, 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.

Figure 27.--Distribution and migration of the red-eyed vireo. It is evident that the red-eyed vireo has only recently invaded Washington by an extension of its breeding range almost due west from the upper Missouri Valley. Like the bobolink, however (fig. 23), the western breeders do not take the short cut south or southeast from their nesting grounds, but migrate spring and fall along the route traversed in making this extension.]

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.

Of all species of North American birds, the Lapland longspur (_Calcarius lapponicus_) seems to be the most frequent victim of mass destruction from storms. These birds sometimes congregate in enormous numbers where grass or weed seed is abundant. Almost every winter brings in reports of their death by thousands somewhere in the Middle West. While migrating northward at night they have encountered blinding storms of wet, clinging snow, which have so bewildered them that they have flown into various obstructions, or have sunk to the ground and perished of exposure and exhaustion. In 1907 an experienced ornithologist estimated that 750,000 longspurs were lying dead on the ice of two lakes in Minnesota, each about 1 square mile in extent, and dead birds were reported in greater or less abundance on this occasion over an area of more than 1,500 square miles. The heaviest mortality occurred in towns, where, bewildered by the darkness and the heavy falling snow, some of the birds congregating in great numbers, flew against various obstacles and were killed or stunned, while many others fell to the ground exhausted. Similar catastrophes have been reported from eastern Colorado, Nebraska, and North Dakota.

During the early part of June 1927, a hailstorm of exceptional severity in and around Denver, Colo., killed large numbers of robins, meadow larks, sparrows, and others. The lawns of parks were strewn with the bodies of these birds, and many lay dead in their nests where they were covering their eggs or young when the storm broke.

AERIAL OBSTRUCTIONS

The destruction of migratory birds by their striking lighthouses, light ships, tall bridge piers, monuments, and other obstructions has been tremendous. Beams of the lanterns at light stations have a powerful attraction for nocturnal travelers of the air. It may be likened to the fascination for lights that is shown by many insects, particularly night-flying moths. The attraction is not so potent in clear weather, but when the atmosphere is moisture laden, as in a heavy fog, the rays have a dazzling effect that lures the birds to their death. They may fly straight up the beam and dash themselves headlong against the glass, or they may keep fluttering around the source of the light until exhausted, and then drop to the rocks or waves below. The fixed, white, stationary light located 180 feet above sea level at Ponce de Leon Inlet (formerly Mosquito Inlet), Fla., has caused great destruction of bird life even though the lens is shielded by wire netting. On one occasion an observer gathered up a bushel-basketful of warblers, sparrows, and other small passerine birds that had struck during the night. The birds apparently beat themselves to death against the wire or fell exhausted to the concrete pavement below, frequently to be destroyed there by cats or skunks. Two other lighthouses at the southern end of Florida, Sombrero Key and Fowey Rocks, have been the cause of a great number of bird tragedies, while heavy mortality has been noted also At some of the lights on the Great Lakes and on the coast of Quebec. It is the fixed white lights that cause such disasters to birds, as the stations equipped with flashing or red lights do not present such strong attractions. That it is not a mere case of geographical location has been demonstrated, for it is observed that when fixed white lights have been changed to red or flashing lights, the migrating birds are no longer endangered. At some of the light stations in England and elsewhere, shelves and perches have been placed below the lanterns to afford places where birds can rest until they have overcome their bewilderment.

For many years at the National Capital, the Washington Monument, although unilluminated, caused the destruction of large numbers of small birds, due apparently to their inability to see this obstacle in their path, towering more than 555 feet into the air. One morning in the spring of 1902 the bodies of nearly 150 warblers, sparrows, and other birds were found about its base. Then, as the illumination of the city was improved and the Monument became more visible at night, the loss became steadily less, until by 1920 only a few birds would be killed during an entire migration. On November 11, 1931, however, as part of the Armistice Day celebration, batteries of brilliant floodlights grouped on all four sides about the base of the Monument, were added to the two searchlights already trained on the apex, so that the lighted shaft probably corresponded in brilliancy to a very low magnitude lighthouse lantern. Airplane pilots have ventured opinions that on a clear night it could be seen for 40 miles. It is certain that there is an extensive area of illumination, and on clear, dark nights when the nocturnal travelers seem to fly at lower altitudes, many of them are attracted to the Monument as to a lighthouse beacon, and wind currents prevent a last-minute avoidance. During the fall migration of 1932 more than 500 warblers, vireos, thrushes, kinglets, sparrows, and others were killed. In 1933 the mortality was less, but the Monument at times still remains a serious menace to birds during migration.

When the torch on the Statue of Liberty in New York Harbor was kept lighted, it caused an enormous destruction of bird life, tabulations showing as many as 700 birds in a single month.

EXHAUSTION

Although it would seem that the exertion incident to the long flights of many species of migratory birds would result in their arrival at their destination in a state bordering on exhaustion, this is contrary to the truth. Both the soaring and the sailing of birds show them to be proficient in the use of factors employed in aerial transportation that only recently have become understood and imitated by aeronautical engineers. The use of ascending currents of air, employed by all soaring birds, and easily demonstrated by observing the gulls that glide hour after hour along the windward side of a ship, are now utilized by man in his operation of gliders. Moreover, the whole structure of a bird renders it the most perfect machine for extensive flight that the world has ever known. Hollow, air-filled bones, making an ideal combination of strength and lightness, and the lightest and toughest material possible for flight in the form of feathers, combine to produce a perfect flying machine. Mere consideration of a bird's economy of fuel or energy also is enlightening. The golden plover, traveling over the oceanic route, makes the entire distance of 2,400 miles from Nova Scotia to South America without stop, probably requiring about 48 hours of continuous flight. This is accomplished with the consumption of less than 2 ounces of fuel in the form of body fat. To be as economical in operation, a 1,000-pound airplane would consume in a 20-mile flight not the gallon of fuel usually required, but only a single pint.

The sora, or Carolina rail (_Porzana Carolina_), which is such a notoriously weak flyer that at least one writer was led to infer that most of its migration was made on foot, has one of the longest migration routes of any member of its family, and easily crosses the wide reaches of the Caribbean Sea. The tiny ruby-throated hummingbird (_Archilochus colubris_) crosses the Gulf of Mexico in a single flight of more than 500 miles.

While birds that have recently arrived from a protracted flight over land or sea sometimes show evidences of being tired, their condition is far from a state of exhaustion, unless unusual conditions have been encountered, and with a few hours' rest and a crop well filled with proper food they exhibit eagerness to resume their journey. The popular notion that birds find the long ocean flights excessively wearisome and that they sink exhausted when terra firma is reached does not agree with the facts. The truth lies in the opposite direction, as even small land birds are so little averse to ocean voyages that they not only cross the Gulf of Mexico at its widest point, but may even pass without pause over the low, swampy coastal plain to the higher regions beyond. Under favorable conditions birds can fly when, where, and how they please. Consequently the distance 4 covered in a single flight is governed chiefly by the food supply. Exhaustion, except as the result of unusual factors, cannot be said to be an important peril of migration.

=INFLUENCE OF THE WEATHER ON MIGRATION=

The state of the weather at any point has little if anything to do with the time of arrival of migratory birds. This is contrary to the belief of observers who have thought that they could foretell the appearance of various species by a study of the weather conditions. Though the insistent crescendo note of the ovenbird is ordinarily associated with the full verdure of May woods, this bird has been known to reach its breeding grounds in a snowstorm and the records of its arrival in southern Minnesota show a temperature variation from near freezing to full summer warmth. Temperatures recorded at the time of arrival of several other common birds show variations of 14° to 37° F., the average variation being about 24°.

It should be remembered that North American species spending the winter months in tropical latitudes experience no marked changes in climatic conditions from November to March or April, yet frequently they will start the northward movement in January or February. This is in obedience to physiological promptings and has no relation to the prevailing weather conditions. For migratory birds the winter season is a period of rest, a time when they have no cares other than those associated with the daily search for food or escape from their natural enemies. Their migrations, however, are a vital part of their life cycles, which have become so well adjusted that the seasons of travel correspond in general with the major seasonal changes on their breeding grounds. With the approach of spring, therefore, the reproductive impulse awakens, and each individual bird is irresistibly impelled to start the journey that ends in its summer home.

Through their influence upon the food supply, weather conditions in the breeding areas do, of course, figure prominently in the picture. In fact, they are the major factors in determining the average dates of arrival of the different species. The word "average" must be emphasized, for the migrations of birds have so evolved that in general they synchronize with average climatic conditions. In other words, migration is so ingrained that each species moves north in spring when the average weather that will be encountered is not unendurable. The hardy birds travel early, fearless of the blasts of retreating winter. The more delicate species come later, when there is less danger of encountering prolonged periods of inclement weather. Some of these hardy birds pause and allow the spring season to advance, and then by rapid travel again overtake it, or in some cases actually outstrip it. At times this results in some hardship and occasionally in the destruction of large numbers of individuals, as has happened when early migrating bluebirds have been overwhelmed by a late winter storm. Unless such conditions are prolonged, however, no serious effect on the species is noted. Nevertheless, the soundness of the bird's instincts is evidenced by the fact that natural catastrophes, great though they may be, do not permanently diminish the avian population.

As has been pointed out, the advance of average temperature lines, known as isotherms, is found to correspond closely with the northward movements of certain species. For example, the northward travels of the Canada goose are found to coincide with the advance of the isotherm of 35° F. (fig. 8).

The spring flight of migrants, if interrupted for any reason, is resumed when weather conditions again become favorable, and it is probable that all instances of arrival of birds in stormy weather can be explained on the theory that the flight was begun while the weather was auspicious. The state of the weather when a flight starts at any southern point, the relation of that place to the average position of the bird under normal weather conditions on that date, and the average rate of migratory flight, are data basic to any reasonably accurate prediction of the time arrival may be expected in northern areas.

Head winds are as unfavorable to migration as is rain or snow, as they greatly increase the labor of flight and cut down the speed of cross-country travel. If such winds have a particularly high velocity they may force down the weaker travelers, and should this happen over water areas, large numbers of birds would be lost. Even strong winds that blow in the direction of aerial travel are unfavorable for the birds, as they interfere with their balance and disarrange their feathers. Moderate tail winds and cross or quartering breezes appear to offer the best conditions for the passage of the migrants.

Figure 28.--A banded belted kingfisher, one of about 2,000,000 birds that have been marked with Biological Survey bands.]

=PROBLEMS OF MIGRATION=

BANDING STUDIES

The study of living birds by the banding method, whereby great numbers of individuals are marked (fig. 28) with numbered aluminum leg rings, has come to be recognized as a most accurate means of ornithological research. Since 1920, banding work in North America has been under the direction of the Bureau of Biological Survey, in cooperation with the National Parks Branch of Canada. Every year volunteer cooperators, working under permit, place bands on thousands of birds, game and nongame, large and small, migratory and nonmigratory, each band carrying a serial number and the legend, NOTIFY BIOLOGICAL SURVEY, WASHINGTON, D. C., or on the smaller sizes, NOTIFY BIOL. SURV., WASH., D. C. When a banded bird is reported from a second locality, a delinite fact relative to its movements becomes known, and a study of many cases of this nature develops more and more complete knowledge of the details of migration.

The records of banded birds are also yielding other pertinent information relative to their migrations, such as the exact dates of arrival and departure of individuals, the length of time that different birds pause on their migratory journeys to feed and rest, the relation between weather conditions and the starting times for migration, the rates of travel of individual birds, the degree of regularity with which birds return to the exact summer or winter quarters used in former years, and many other details that could be learned in no other manner. Banding stations that are operated systematically throughout the year, therefore, are supplying much information concerning the movements of migratory birds that heretofore could only be surmised.

MOVEMENTS OF RESIDENTS

Typical migration consists of definite movements that are repeated regularly year after year, and it is to these that the term is generally restricted. It is desirable, however, if only for purposes of comparison, that some account be taken of the movements of some other birds, which, while not typical, do possess some of the characteristics of true migration. Data on this subject are being collected through bird banding.

There are several species that are customarily grouped under the heading "permanent residents", the term implying that these birds do not travel but remain throughout the year in one locality. Among these are the cardinal, the tufted titmouse (_Baeolophus bicolor_), the wren tit (_Chamaea jasciata_), the Carolina wren, the house finch (_Carpodacus mexicanus frontalis_), the bobwhite (_Colinus virginianus_), the California quail (_Lophortyx californica_), and the ruffed grouse (_Bonasa umbellus_). Each species may be present constantly throughout the year, although in the northern part of the range there is probably a slight withdrawal of the breeding birds in winter. The individuals to be seen at that season, therefore, may not always be the same as those observed during the summer. It is certain, however, that these species do not regularly perform extensive journeys.

While the blue jay is disposed to be secretive, it is such a showy and noisy bird that is not likely to escape notice. In the vicinity of Washington, D. C., as in many other places, it is present the year round, but at the end of September or early in October when the weather is becoming cooler, troops of jays are sometimes seen working southward through the trees. A corresponding northward movement occurs again in May. This is unquestionably a migration to and from some winter range, but its extent or significance is not now known. Some light is being shed on the matter, however, through the records of banded birds, and these eventually will fill in a more perfect picture of the movements of this species. One jay, banded on September 14, 1923, at Waukegan, Ill., was killed at Peruque, Mo., on November 15 of the same year; another, banded at Winnetka, Ill., on June 16, 1925, was retaken at Sulphur Kock, Ark., the following December 10; a third, banded on May 6, 1925, at Whitten, Iowa, was recaptured at Decatur, Ark., on January 22, 1926. These three birds unquestionably had made a flight that had every appearance of being a true migration to winter quarters in Missouri and Arkansas.

The black-capped chickadee is apparently resident in many places, but occasionally in winter it invades the range of the southern Carolina chickadee (_Penthestes carolinensis_) and in northern Canada it is regularly a migrant.

In the coastal plain between Washington, D. C., and the Atlantic Ocean, the white-breasted nuthatch is usually absent during the summer, nesting at that season in the higher, or piedmont, country. Late in fall, however, it appears in fair abundance in the wooded bottoms, remaining at the lower levels until the following March or April.

Some birds, including the screech owl (_Otus asio_), bobwhite, Carolina wren, and mockingbird (_Mimus polyglottos_), seem to be actually sedentary, but even these are sometimes given to post-breeding wanderings. Ordinarily bob whites that are marked with numbered bands are seldom retaken far from the area where banded, but sometimes they will travel 10 miles or more. A screech owl banded at Glenwood, Minn., in March, was recovered the following December at Emmetsburg, Iowa, 180 miles south. Such flights, however, are probably more in the nature of a search for new feeding areas, or to escape from a winged enemy, than a true migratory journey.

MIGRATION OF THE WHITE-THROATED SPARROW

The white-throated sparrow (_Zonotrichia albicollis_) (fig. 29), one of the most abundant members of its family, breeds from northern Mackenzie and the southern part of the Ungava Peninsula south to southern Montana, northern Pennsylvania, and Massachusetts. The winter range extends from the southern part of the breeding range south to the Gulf coast and northeastern Mexico. It is therefore a common migrant in many sections. Since it is a ground-feeding bird and is readily attracted to the vicinity of dwellings, it has been banded in large numbers, the total to February 1, 1935, being nearly 100,000. It would be expected that these would yield a comparable number of return records, and that the facts would furnish basic data relative to the migrations of the species. Such, however, is not the case. Banded white-throated sparrows are rarely recaptured at stations between the breeding and the wintering grounds. Operators of stations in the winter area, as Thomasville, Ga., and Summerville, S. C., have obtained return records showing that these birds do come back to the exact winter quarters occupied in previous seasons. The fact that they do not again visit banding stations on their migration routes indicates some unusual aspects of their travels, which it is hoped will eventually be discovered by banding studies. Problems of this type constitute definite challenges to the student of bird migration.

MIGRATION OF THE YELLOW-BILLED LOON

The semiannual movements of the yellow-billed loon present an unusual problem in migration. It breeds along the Arctic coast, probably from Cape Prince of Wales eastward to Franklin Bay, and also in the interior of northern Canada south to Clinton-Colden, Aylmer, and Artillery Lakes, where it is rather common. It has been reported as already present by May 25 at the mouth of the Liard River, in southwestern Mackenzie. This coincides with the time that first arrivals are noted fully 700 miles north, at Point Barrow, Alaska. The problem has been to ascertain the route used by these birds to then principal nesting grounds in the interior.

For a long time it was believed that this big diver did not winter in large numbers anywhere on the Pacific coast, and it had been supposed that the spring route extended 2,000 miles northeastward from a wintering ground somewhere in eastern Asia to Bering Strait, then 500 miles still northeast to round Point Barrow, then 500 miles east to the coast of Mackenzie, and finally 700 miles south--in spring--to the region near the eastern end of Great Slave Lake.

The yellow-billed loon is a powerful flier, and it is probable that this suggested route is correct for those birds that breed in the northern coastal regions. A reasonable doubt may be entertained, however, whether the breeding birds of Great Slave Lake and contiguous areas reach their breeding grounds by the 700-mile flight south from the Arctic coast. Within recent years it has been found that these birds are fairly common in the maze of channels and islands off the coast of southeastern Alaska as late as the last of October and in February. Possibly they are present there during the period from November through January also, or they may at that time move farther offshore and so escape detection. If this region is an important wintering ground, as seems probable, then it is likely that the breeding birds of the interior reach their nesting grounds by a flight eastward across the mountains, a trip that is well within their flying ability, rather than by a circuitous route around the northern coast. The air-line distance from southeastern Alaska to the mouth of the Liard River is in fact less than the distance to that point from the mouth of the Mackenzie.

Figure 29.--White-throated sparrow, a bird that apparently breeds and winters each year in the same areas, but either travels by different routes or, at least, does not make the same stops, while on migration.]

Differing routes to various parts of a large breeding or wintering ground, and used by large groups of individuals of other species, are not unknown. For example, the redhead duck is one of the common breeding ducks of the Bear River marshes of Utah, where a great many have been banded each summer. The recovery records of banded redheads show that while many travel westward to California, others start their fall migration in the opposite direction and, flying eastward across the Rocky Mountains, either turn southeast across the plains to the Gulf of Mexico, or deliberately proceed in a northeasterly direction to join the flocks of this species moving toward the Atlantic coast from the prairie regions of southern Canada.

=CONCLUSIONS=