Part 3

It seems more likely that migrations are performed in a leisurely manner, and that after a flight of a few hours the birds pause to feed and rest for one or several days, particularly if they find themselves in congenial surroundings. Some indication of this is found in the records of banded birds, particularly waterfowl. Considering only the shortest intervals that have elapsed between banding in the North and recovery in southern regions, it is found that usually a month or more is taken to cover an air-line distance of a thousand miles. For example, a black duck banded at Lake Scugog, Ontario, was killed 12 days later at Vicksburg, Miss. If the bird was taken shortly after its arrival, the record would indicate an average daily flight of only 83 miles, a distance that could have been covered in about 2 hours' flying time. Among the thousands of banding records obtained in recent years, evidences of such rapid flight are decidedly scarce, for with few exceptions all thousand-mile flights have required 2 to 4 weeks or more. Among sportsmen, the blue-winged teal is well known as a fast-flying duck and quite a few of these banded on Canadian breeding grounds have covered 2,300 to 3,000 miles in a 30-day period. Nevertheless, the majority of those that have traveled to South America were not recovered in that region until 2 or 3 months after they were banded. Probably the fastest flight over a long distance for one of these little ducks was one made by a young male which traveled 3,800 miles from the delta of the Athabaska River, in northern Alberta, Canada, to Maracaibo, Venezuela, in exactly 1 month. This flight was at an average speed of 125 miles per day. The greatest migration speed thus far recorded for any banded bird is that of a lesser yellowlegs banded at North Eastham, Cape Cod, Mass., on August 28, 1935, and killed 6 days later, 1,900 miles away, at Lamentin, Martinique, French West Indies. This bird traveled an average daily distance of more than 316 miles.

It seems certain that migratory journeys are performed at the normal rate of flight, as this would best conserve the strength of the birds and eliminate the fatigue that would result from effort required for great speed. Migrating birds passing lightships and lighthouses, or crossing the face of the moon, have been observed to fly without hurry or evidence of straining to attain high speed.

The speed of migration also is demonstrated by the dates of arrival, particularly during the spring movement. The Canada goose affords a typical example of regular, but slow migration. Its advance northward at this season is at the same rate as the advance of the season (fig. 4). In fact, the isotherm of 35° F. appears to be a governing factor in the speed at which these geese move north, and over their entire trip the vanguard follows closely the advance of this isotherm.

Few species perform such regular migrations, many waiting in their winter homes until spring is well advanced and then moving rapidly to their breeding grounds. Sometimes this advance is so rapid that the later migrants actually catch up with species that for a month or more may have been pressing slowly but steadily northward.

One of the best examples of rapid migration is found in the gray-cheeked thrush. 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. 5). 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 temperatures take 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, the crow, the red-winged blackbird, and the robin.

The blue goose 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 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 blackpolled warbler furnishes an excellent example (fig. 6). 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 birds. 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. 3), 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.

=Altitude 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 rarified 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 the rarified 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. 28), 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 crossing the Gulf of Mexico to Louisiana and Florida flew 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 a height of 3,000 feet above the earth. 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, and northern phalaropes, 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 Wars broad areas in the air were under constant 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 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 so much as the unerring certainty with which birds 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 from birds marked with numbered bands offer 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 other mammals as well as by some insects and fishes, the well-known migrations of the salmon and the eel 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 well known 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 by 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 coastlines and rivers are just those that are most likely to be seen in the faintest light, particularly by the acute vision of the 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, 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 decks, and carried northward to distances varying from 400 to 800 miles before being released. Landmarks of all kinds 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-o'-war birds 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 should 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."

Some students have leaned strongly toward the possible existence of a "magnetic sense" as being the important factor in the power of geographical orientation. The theory that migratory birds might be responsive to the magnetic field of the earth was conceived as early as 1855, when some experimental work was done in Russia, and nearly 60 years later in France. Recently investigations in this field have been conducted by Yeagley (1947) and by Gordon (1948) with diametrically opposite results. The idea carries with it the implication that contained in the bird's body is an organ that is sensitive to the effect of its motion through the vertical component of the magnetic field and to other related factors. In the tests by Dr. Yeagley, 20 young homing pigeons were given training flights to their home loft from distances up to 100 miles. Permanent magnets were then affixed to the under side of the manus part of the wings of half of the birds while copper plates of equivalent weight were attached to the wings of the other half. All birds were released singly at an air-line distance of about 65 miles from the loft. The results were most suggestive, as only two of the birds carrying magnets returned to the loft, whereas eight of the controls returned.

With certain minor modifications, this experiment was repeated by Gordon. In this case 60 pigeons were used and releases were made from points up to 58 miles, where the direction of flight was such that the birds had to navigate across the gradient of the magnetic field. Every bird returned to its loft on the day of release regardless of whether it carried magnets or unmagnetized bars of the same weight.

Attempts to demonstrate the effect of radio waves on the navigational ability of birds also have produced contradictory results. In some of these tests, homing pigeons released near broadcasting stations have appeared to be hopelessly confused, whereas in others, apparently conducted in the same manner, no effects could be discerned. It is obvious that before the electromagnetic theory can be accepted or rejected, much additional experimental work is necessary.