Part 3

One of the best examples of rapid migration is found in the gray-cheeked thrush (_Hylocichla minima aliciae_). This bird winters in Colombia, Ecuador, Peru, Venezuela, and British Guiana and does not start its northward journey until many other species are well on their way. It does not appear in the United States until the last of April--April 25, near the mouth of the Mississippi, and April 30 in northern Florida (fig. 9). A month later, or by the last week in May, the bird is seen in northwestern Alaska, the 4,000-mile trip from Louisiana having been made at an average speed of about 130 miles a day.

Another example of rapid migration is furnished by the yellow, or summer, warbler. Coming from the Tropics, the birds reach New Orleans about April 5, when the average temperature is 65° F. Traveling north much faster than does the season, they reach their breeding grounds in Manitoba the latter part of May, when the average temperature is only 47°. Encountering progressively colder weather over their entire route, they cross a strip of country in the 15 days from May 11 to 25 that spring takes 35 days to cross. This "catching up" with spring is habitual in species that winter south of the United States and in most of the northern species that winter in the Gulf States. To this rule there appear to be only six exceptions--the Canada goose, the mallard, the pintail (_Dafila acuta_), the crow (_Corvus brachyrhynchos_), the red-winged blackbird (_Agelaius phoeniceus_), and the robin.

The blue goose (_Chen caerulescens_) (fig. 10) presents a striking example of a late but very rapid spring migration. Practically all members of the species winter in the great coastal marshes of Louisiana, where 50,000 or more may be seen grazing in the "pastures" or flying overhead in flocks of various sizes. Their breeding grounds are chiefly on Baffin Island and on Southampton Island in the northern part of Hudson Bay, in a region where conditions of severe cold prevail except for a few weeks each year. The birds seem to realize that even though the season in their winter quarters is advancing rapidly, their nesting grounds are still covered with a heavy blanket of ice and snow. Accordingly they remain in the coastal marshes until the last of March or the first of April, when the local birds are already busily engaged with the duties of reproduction. The flight northward is rapid, almost nonstop, so far as the United States is concerned, for although the birds are sometimes recorded in large numbers in the Mississippi Valley, including eastern South Dakota, and in southeastern Manitoba, there are few records anywhere along the route of such great flocks as are known to winter in Louisiana. When the birds arrive in the James Bay region of Canada they apparently enjoy a prolonged period of rest, as they are not noted in the vicinity of their breeding grounds until the first of June. During the first 2 weeks of that month they pour into the tundra country by the thousands, and each pair immediately sets about the business of rearing a brood.

The robin has been mentioned as a slow migrant, and as a species it takes 78 days to make the 3,000-mile trip from Iowa to Alaska, a stretch of country that is crossed by advancing spring in 68 days. In this case, however, it does not mean that individual robins are necessarily slow, for probably the northward movement of the species depends upon the continual advance of birds from the rear, the first individuals arriving in a suitable locality remaining to nest, while the responsibility of the northward movement of the species is continued by those still to come.

Special interest attaches to the great variation in the speed at which birds travel in different sections of the broad flyway extending from the Gulf of Mexico to the Arctic Ocean by way of the Mississippi and Mackenzie Valleys. The blackpoll warbler furnishes an excellent example (fig. 11). This species winters in north-central South America and migrates in April across the West Indies to Florida. From this point some individuals fly northwest to the Mississippi Valley, north to Manitoba, northwest to the Mackenzie River, and thence almost due west to western Alaska. In tracing the long route of these birds it is found that a fairly uniform average speed of 30 to 35 miles a day is maintained from the Gulf to Minnesota. Then comes a spurt, for a week later the blackpolls have reached the central part of the Mackenzie Valley, and by the following week they are observed in northwestern Alaska. During the latter part of the journey, therefore, many individuals must average more than 200 miles a day. They use 30 days in traveling from Florida to southern Minnesota, a distance of about 1,000 miles, and scarcely half that time to cover the remaining 2,500 miles to Alaska. It should be noted that the increased speed is directly associated with the change in direction, the north-and-south course in the Mississippi Valley being accomplished slowly, while the northwesterly course across Canada is made at a much greater speed. Increased speed across western Canada to Alaska is also shown by many other species. A study of all species traveling up the Mississippi Valley indicates an average speed of about 23 miles a day. From southern Minnesota to southern Manitoba 16 species maintain an average speed of about 40 miles a day. From that point to Lake Athabaska, 12 species travel at an average speed of 72 miles a day; while 5 others travel to Great Slave Lake at 116 miles a day; and another 5 species cover 150 miles a day to reach Alaska. This change is in correlation with a corresponding variation in the isothermal lines, which turn northwestward west of the Great Lakes.

As has been previously indicated, the advance of spring in the northern interior is much more rapid than in the Mississippi Valley and on the Gulf coast. In other words, in the North, spring comes with a rush, and during the height of the migration season in Saskatchewan the temperature in the southern part of the Mackenzie Valley just about equals that in the Lake Superior area, which is 700 miles farther south. Such conditions, coupled with the diagonal course of the birds across this region of fast-moving spring, exert a great influence on migration and are the chief factors in the acceleration of speed of travel.

Variations in speed of migration in different parts of the country are illustrated also by the movements of the cliff swallow (fig. 5), which breeds from Mexico to Alaska and winters in Brazil and Argentina. It would be expected in spring to appear in the United States first in Florida and Texas, then in the southern Rocky Mountain region, and finally on the Pacific coast. As a matter of fact, however, the earliest spring records come from north-central California, where the bird usually is common before the first arrivals are observed in Texas or Florida. The route taken, for many years a migration problem, was solved when it was found that these swallows went around the Gulf of Mexico rather than across it. The isochronal lines on the map show the more rapid advance along the Pacific coast. By March 20, when the vanguard has not quite reached the lower Rio Grande in Texas, the species is already north of San Francisco in California.

ALTITUDES AT WHICH BIRDS TRAVEL

At one time students of bird migration held firmly to the theory that normal migration takes place at heights above 15,000 feet, reasoning (somewhat uncertainly) that flying becomes easier as altitude is gained. Since the development of the airplane, however, and with it man's exploration of the upper regions of the air, it has become common knowledge that rarefied atmosphere adds greatly to the difficulties of flight. This is due not only to the reduction in oxygen (whether for gasoline engine or the lungs of a bird) but also to the lack of buoyancy of rarefied air. Such birds as vultures, pelicans, cranes, and some of the hawks feel this the least, since compared with body weight the supporting surface of their wings is very great, but for the smaller and shorter winged birds lack of buoyancy at high altitudes presents a difficult obstacle in flight. Even when flying close to the earth, small birds have to keep their wings in rapid motion.

Another postulate favoring the high-altitude flying theory was that the wonderful vision of birds was their sole guidance during migratory flights; and to keep landmarks in view the birds were obliged to fly high, particularly when crossing wide areas of water. This will be considered in greater detail under Orientation (p. 23), so here it will be sufficient to say that birds rely only in part upon vision to guide them on migration. Also, it is to be remembered that there are definite physical limitations to the range of visibility even under perfect atmospheric conditions. Chief of these is the curvature of the earth's surface. Thus, if birds flew over the Gulf of Mexico to Louisiana and Florida at a height of 5 miles, they would still be unable to see a third of the way across. And yet this trip is made twice each year by thousands of thrushes, warblers, and others.

Actual knowledge of the altitude of migratory flight is scanty, though estimates obtained by means of the telescope, and still more accurate data resulting from altimeter observation from airplanes, are slowly accumulating. It is, of course, obvious that some birds that cross mountain ranges during migration must attain a great altitude. Observers at an altitude of 14,000 feet in the Himalayas have recorded storks and cranes flying so high that they could be seen only through field glasses. Being beyond the range of unaided vision they must have been at least 6,000 feet above the observers, or at an actual altitude of 20,000 feet above sea level. Such cases, however, are exceptional, as aviators have reported that they rarely meet birds above an altitude of 5,000 feet.

It is now known that migration in general is performed below an altitude of 3,000 feet. Some proof of this statement is available. Observations made from lighthouses and other points of vantage indicate that migrants commonly travel at altitudes of a very few feet to a few hundred feet above sea or land. Sandpipers, sanderlings (_Crocethia alba_), and northern phalaropes (_Lobipes lobatus_), observed in migration on the Pacific oceanic route, have been noted to fly so low that they were visible only as they topped a wave. Observers stationed at lighthouses and lightships off the English coast have similarly recorded the passage of land birds, which sometimes flew just above the surface of the water, and rarely above 200 feet. During the World War broad areas in the air were under constant close surveillance, and among the airplane pilots and observers many took more than a casual interest in birds. Of the several hundred records resulting from their observations only 36 were of birds flying above 5,000 feet, and only 7 above 8,500 feet. Cranes were once recorded at an altitude of 15,000 feet, while the lapwing (_Vanellus vanellus_) was the bird most frequently seen at high levels, 8,500 feet being its greatest recorded altitude.

These observations naturally relate only to daytime travelers, but there is no reason to believe that nocturnal migration is performed at higher altitudes. The fact that many birds are killed each year by striking the lanterns at lighthouses, or other man-made obstructions, does not, however, furnish conclusive proof that low altitudes are generally used during nocturnal flight, for it should be recalled that these accidents occur chiefly in foggy or unsettled weather, and also, that powerful lights have a great attraction for many species of birds. The altitude at which birds travel is affected by other weather conditions also. For example, flight at the higher elevations is facilitated on clear, warm days by the currents of warm air that ascend from broad areas.

ORIENTATION

There probably is no single aspect of the entire subject of bird migration that challenges our admiration for birds so much as the unerring certainty with which they cover thousands of miles of land and water to come to rest in exactly the same spot where they spent the previous summer or winter. The records of birds marked with numbered bands afford abundant proof that the same individuals of many species will return again and again to their identical nesting sites. These data show also that many individuals migrate in fall over the same route, year after year, making the same stops, and finally arriving at the precise thicket that served them in previous winters.

The faculty that enables these birds to point their course accurately over vast expanses of land and water may for want of a better term be called a "sense of direction." Man recognizes this sense in himself, though usually it is imperfect and frequently at fault. Nevertheless the facility with which experienced hunters and woodsmen locate tiny camps or other points in forested or mountainous country, frequently cloaked by darkness or fog, with all recognizable landmarks obliterated seems due to this faculty. Ability to travel with precision over unmarked trails is not limited either to birds or to man. It is likewise possessed by many mammals as well as by some insects and fishes, the well-known migrations of the salmon (_Oncorhynchus_) and the eel (_Anguilla_) being notable examples.

Ability to follow a more or less definite course to a definite goal is evidently part of an inherited faculty. Both the path and the goal must have been determined either when the habit originated or in the course of its subsequent evolution. The theory is sometimes advanced that the older and more experienced birds lead the way, showing the route to their younger companions. This explanation may be acceptable for some species, but not for those in which adults and the young migrate at different times. The young cowbird that is reared by foster parents flocks with others of its kind when grown and in many cases can hardly be said to have adult guidance in migration. An inherited migratory instinct with a definite sense of the goal to be reached and the route to be followed must be attributed to these birds.

It is known, however, that birds possess wonderful vision. If they also have retentive memories, subsequent trips over the route may well be steered in part by recognizable landmarks. The arguments against the theory of vision and memory are chiefly that much migration takes place at night and that great stretches of the open sea are crossed without hesitation. Nevertheless, the nights are rarely so dark that all terrestrial objects are totally obscured, and such features as coast lines and rivers are just those that are most likely to be seen in the faintest light, particularly by the acute vision of a bird and from its aerial points of observation. But some birds fly unerringly through the densest fog. Members of the Biological Survey, proceeding by steamer from the island of Unalaska to Bogoslof Island in Bering Sea through a fog that was so heavy as to make invisible every object beyond a hundred yards, recorded the fact that flocks of murres, returning to Bogoslof, after quests for food, broke through the wall of fog astern, flew by the vessel, and disappeared into the mists ahead. The ship was heading direct for the island by the use of compass and chart, but its course was no more sure than that of the birds.

Some investigators have asserted that the sense of direction has its seat in the ears or nasal passages and thus that the bird is enabled to identify air currents and other phenomena. It has been found that disturbance of the columella, or the semicircular canals of the inner ear, will destroy the homing instinct of the racing pigeon, but experiments in the form of delicate operations, or closing the ears with wax, prove such a serious shock to the sensitive nervous system of the bird that they cannot be considered as affording conclusive evidence. Several years ago careful studies were made of the homing instinct of the sooty and noddy terns (_Sterna fuscata_ and _Anoüs stolidus_), tropical species that in the Atlantic region reach their most northern breeding point on the Dry Tortugas Islands, off the southwest coast of Florida. They are not known to wander regularly any appreciable distance farther north. It was found that some were able to return to their nests on the Tortugas after they had been taken on board ship, confined in cages below deck, and carried northward distances varying from 400 to 800 miles before being released. Landmarks of all lands were entirely lacking, and the birds certainly were liberated in a region in which they had had no previous experience.

Possibly the "homing instinct" as shown by these terns, by the man-of-war birds (_Fregata minor_), that are trained and used as message carriers in the Tuamotu, Gilbert, and Marshall Islands, and by the homing pigeon, is not identical with the sense of perceptive orientation that figures in the flights of migratory birds. Nevertheless, it seems closely akin and is probably caused by the same impulses, whatever they may be and however they may be received. It is to be remembered, however, that while homing may involve flight from a point that the bird has never before visited, the flight is always to a known point — that is, the bird's nest— while, on the other hand, the first migratory flight is always from the region of the bird's birth to a region it has never before visited. The spring migration might, of course, be more nearly considered as true "homing."

At the present time some students lean strongly toward the possible existence of a "magnetic sense" as being the important factor in the power of geographical orientation. No direct evidence in support of this has been obtained, but it is not impossible that there may exist some form of physiological sensibility to the phenomena of terrestrial magnetism. The theory as laid down (chiefly by European investigators) is highly complex, but briefly stated it is based on a supposed sensitiveness of birds to the magnetic influences that cause variations in the declination and dip of magnetic needles. Some experimental work already done lends a little support to the theory but it is still far from established.

In concluding this discussion of orientation it is pertinent to point out that the migratory instinct appears to be more or less transitory, that it is not persistent over an extended period. Migratory birds may be arrested en route, either by natural conditions, such as unusual food supplies, or forcibly by the act of man, and detained until the end or nearly the end of the migratory season, and then may not attempt to finish the journey, apparently having lost the migratory impulse. In the fall and early winter of 1929, abundant food and an open season caused an unusual number of mallard ducks to arrest their migration and remain in western Montana and northern Idaho. Later, however, a heavy snowfall with subzero temperatures suddenly cut off the food supply, with the result that great numbers of the birds starved to death, when a flight of a few hours would have carried them to a region of open water and abundant food.

SEGREGATION DURING MIGRATION

During the height of the northward movement in spring the woods and thickets may be suddenly filled with several species of wood warblers, thrushes, sparrows, flycatchers, and others, which it is natural to conclude have traveled together and arrived simultaneously. Probably they did, but such combined migration is by no means the rule for all species.

As a group the wood warblers (Compsothlypidae) probably travel more in mixed companies than do any other single family of North American birds. The flocks are likely to be made up of several species, spring and fall, with both adults and young. Sometimes swallows, sparrows, blackbirds, and some of the shore birds also migrate in mixed flocks. In fall, great flocks of blackbirds frequently sweep south across the Plains States, and occasionally one flock will contain bronzed grackles (_Quiscalus quiscula_), red-winged blackbirds, yellow-headed blackbirds (_Xanthocephalus xanthocephalus_), and Brewer's blackbirds (_Euphagus cyanocephalus_).

On the other hand, many species keep strictly to themselves. It would be difficult for any other kind of bird to keep in company with a bird of such rapid movements as the chimney swift, which is rarely found associated with any other species at any season. Nighthawks, or bullbats, also fly in separate companies, as usually do crows, waxwings, crossbills, bobolinks, and kingbirds. Occasionally, a flock of ducks will be observed to contain several species, but generally when they are actually on migration the individuals of each species separate and travel with others of their own kind. The flocks of blue geese, previously mentioned in connection with speed of flight (p. 18), frequently have with them a few of the closely related snow geese (_Chen hyperborea_), particularly in the eastern part of their winter range. The proportion here is usually about 10 to 1, but farther west the numbers of snow geese increase until they outnumber their blue relatives.

The adults of most perching birds drive the young away when they are grown, probably to be relieved of the necessity of providing for them, and also in order that the parents may have opportunity to rest and renew their plumage before starting for winter quarters. The young birds are therefore likely to drift together and, having no further responsibility, may start south ahead of their parents. In contrast with this indifference on the part of the adults of perching birds, Canada geese and some others, remain in family groups, the parent birds undergoing the wing molt that renders them flightless during the period of growth of their young, so that old and young acquire their full plumage at the same time and thus are able to start south together. The large flocks, therefore, are composed of many families that band together, and when they separate into V-shaped units it is probably correct to assume that it is an old bird that leads the group. Where there is segregation of the sexes, the young birds usually accompany their mothers, as is the case with some of the ducks. After the females start to incubate their eggs, the males of most species of ducks flock by themselves and remain together until fall.