Part 2

Experimental work has abundantly demonstrated the effect of increased light upon the growth, flowering, and fruiting of plants. Similarly, experiments with the common junco, or snowbird (_Junco hyemalis_), reported by Rowan (_42, p. 121_), resulted in increased development of the sexual organs by the end of December, although the birds were confined in outdoor aviaries in Canada and had been exposed to temperatures as low as -44° F. From the first of November until early in January, the juncos were subjected to ever-increasing light, supplied in the aviaries by electric bulbs. As regards illumination, they were thus artificially provided with conditions approximating those of spring. At the close of this period, it was found that the sexual organs of the birds had attained the maximum development normally associated with spring. With gradual reduction of the lighting over a period of little more than 1 month, the organs returned to their normal winter condition.

After a consideration of all evidence, including the fact that no ultraviolet rays were used, it was concluded that the explanation lay in the increased exercise taken during the periods of increased light. A simple test whereby certain birds were forced by mechanical means to take more exercise, the light being so reduced that there was merely sufficient glow for them to see the advancing mechanism that forced them into movement, showed that the rate of development of the sexual organs exactly paralleled that in the birds that were exposed to extended periods of illumination in the outdoor aviaries. Other features in this experiment--such as the behavior of the birds themselves--also indicated that more activity due to increased light is the governing cause of the spring development of the sexual organs. If this development be accepted as a controlling cause of migration, then this experiment must be recognized as of great importance.

Upon closer analysis, however, it is found that this theory, like those before discussed, is open to serious objections. First, some of our summer residents that migrate south for the winter do not stop in equatorial regions, where they might find the periods of day and night about equally divided, but push on beyond, some penetrating as far south as Patagonia. Also it might be asked: If the lengthening day is the stimulating factor, why should our summer birds wintering in the Tropics ever start northward, as in their winter quarters the variation in the length of day from winter to summer is imperceptible. Like all the other theories advanced, this also, as at present understood, is subject to unanswered criticism.

=WHEN BIRDS MIGRATE=

It is known that at any given point many species leave in fall and return in spring. Since bird banding has had such wide application as a method of study, it is known also that in some species one of the parent birds (rarely both) frequently returns and nests in the same tree, bush, or box that held its nest in the previous season (fig. 1). One ordinarily thinks of the world of birds as quiescent during two seasons each year, at nesting time and in winter. For individual species this is obviously the case, but when the entire avifauna of the continent is considered it is found that there are at almost all periods some latitudinal movements.

MOVEMENTS OF SPECIES AND GROUPS

Some species begin their fall migrations early in July, and in some parts of the country distinct southward movements can be detected from then until the beginning or middle of winter. For example, many shore birds start south in the early part of July, while the goshawks (_Astur atricapillus_), snowy owls (_Nyctea nyctea_), redpolls (_Acanthis linaria_), Bohemian waxwings (_Bombycilia garrula_), and many others do not leave the North until forced to do so by the advent of severe winter weather or by lack of the customary food. Thus, an observer in the northern part of the United States may record an almost unbroken southward procession of birds from midsummer to winter, and note some of the returning migrants as early as the middle of February. Purple martins (_Progne subis_) have been known to arrive in Florida late in January on their way north, and the northern movement may continue among late arrivals into the first week of June. In some species the migration is so prolonged that the first arrivals in the southern part of the breeding range will have performed their parental duties while others of that species are still on their way north.

A study of these facts indicates that sometimes there exists a very definite relationship between what we may term northern and southern groups of individuals of the same species. A supposition, on which additional banding work is expected later to give definite facts, is that in the case of some species that have an extensive latitudinal breeding range and a normal migration, the individuals that nest farthest south migrate first in fall and proceed to the southern part of the winter range; those that occupy the central parts of the breeding range migrate next, and travel to regions in the winter range north of those occupied by the first group; and finally the individuals breeding farthest north are the last to begin their fall migration and these remain farthest north during the winter. In other words, this theory supposes that the southward movement of the species is normally such that the different groups maintain their relative latitudinal positions, both spring and fall.

The black-and-white warbler (_Mniotilta varia_) furnishes an example. The breeding range of this bird extends west and northwest from South Carolina and New Brunswick as far as Great Bear Lake in northwestern Canada (fig. 2). The bird spends the winter in southern Florida, the West Indies, central Mexico, Central America, and northwestern South America. In the southern part of its breeding range it is nesting in April, but the summer residents of New Brunswick do not reach their breeding grounds before the middle of May. Therefore, about 50 days are required for these northbound birds to cross the breeding range, and if 60 days be allowed for nest building, egg laying, incubation, care of the young, and molt, they would not be ready to start southward before the middle of July (fig. 3). Then another 50-day trip south, and the earliest migrants from the northern areas would reach the Gulf coast in September. But both adults and young have been observed at Key West, Fla., by the middle of July, and on the northern coast of South America by August 21. Since the birds at Key West were fully 500 miles south of the breeding-range, it is evident that they must have come from the southern part of the nesting area.

Many similar cases might be mentioned, such as the black-throated blue warblers (_Dendroica caerulescens_), which are still observed in the mountains of Haiti in the middle of May when others of the species are en route through North Carolina to breeding territory in New England or have even reached that region. Redstarts (_Setophaga ruticilla_) and yellow warblers (_D. aestiva_), evidently the more southern breeders in each case, are seen returning southward on the northern coast of South America just about the time that the earliest of those breeding in the North reach Florida on their way to winter quarters.

[Illustartion: B2282M

Figure 4.--The Kentucky warbler, a night migrant, in traveling to its winter quarters in Central America and northwestern South America, uses route no. 5, figure 20.]

NOCTURNAL AND DIUNAL MIGRATION

When one recalls that most birds appear to be more or less helpless in the dark, it seems remarkable that many should select the night hours for extended travel. Among those that do, however, are the great hosts of shore birds, rails, flycatchers, orioles, most of the great family of sparrows, the warblers (fig. 4), vireos, and thrushes, and, in fact, the majority of small birds. That it is common to find woods and fields on one day almost barren of bird life and on the following day filled with sparrows, warblers, and thrushes, would indicate the arrival of migrants during the night. The passage of flocks of ducks and geese is frequently observed by sportsmen sitting in their blinds, but great numbers of these birds also pass through at night, the clarion call of the Canada goose (_Branta canadensis_) or the conversational gabbling of a flock of ducks being common night sounds in spring and fall in many parts of the country. The sibilant, nocturnal calls of the upland plover, or Bartramian sandpiper (_Bartramia longicauda_), and other shore birds during their spring and fall flights form vivid memories in the minds of many students of migration. Observations made with telescopes focused on the full moon have shown processions of birds. The estimate of one observer that birds passed his point of observation at the rate of 9,000 an hour gives some indication of the numbers of birds that are in the air during some of the nights when migration is at its height. While the passage of migratory birds has thus been recorded throughout the night, the bulk of the flocks pass during the earlier hours of the evening and toward daylight in the morning, the periods from 8 o'clock to midnight and from 4 to 6 a. m. seeming to be favorite times for nocturnal flight.

It has been claimed, with some reason, that small birds migrate by night the better to avoid their enemies, and that most of the nocturnal travelers are those that are naturally timid, sedentary, or feeble-winged. Included in this group are not only small song and insectivorous birds, but also such weak fliers as the rails, as well as the wrens, the small woodland flycatchers, and other species, which, living habitually more or less in concealment, are probably much safer making their long flights under the protecting cloak of darkness. This cannot fully account for the nocturnal habit, however, since among the night migrants are the snipe, sandpipers, and plovers, birds that are generally found in the open and are among the more powerful fliers, some of them making flights of more than 2,000 miles across the ocean. Such exceptionally long flights, of course, require both day and night flying.

Night travel is probably best for the majority of birds, chiefly from the standpoint of feeding. Digestion is rapid in birds, and yet the stomach of a bird killed during the day almost always contains food. To supply the energy required for long flight, it is essential that food be obtained at comparatively short intervals, the longest of which in most species is during the hours of darkness. If the smaller migrants were to make protracted flights by day they would be likely to arrive at their destination at nightfall almost exhausted, but unable to obtain food until the following morning, since they are entirely daylight feeders. This would delay resumption of flight and result in great exhaustion or possibly even death were they so unfortunate as to have their evening arrival coincident with unusually cold or stormy weather. Traveling at night, they pause at daybreak and devote the entire period of daylight to alternate feeding and resting. This permits complete recuperation and resumption of the journey at nightfall.

Many species of wading and swimming birds migrate indifferently by day or by night, as they are able to feed at all hours and are not accustomed to seek safety in concealment. Some diving birds, including ducks that submerge when in danger, sometimes travel over water by day and over 1 and at night. The day migrants include, in addition to some of the ducks and geese, the loons, cranes, gulls, pelicans, hawks (fig. 7), swallows, nighthawks, and the swifts (fig. 6), all strong-winged birds. The swifts, swallows, and nighthawks (sometimes called bullbats) feed entirely on flying insects, and use their short, weak feet and legs only for grasping a perch during periods of rest or sleep. Thus they feed as they travel, the circling flocks being frequently seen late in summer working gradually southward. Years ago, before birds of prey were so thoughtlessly slaughtered, great flocks of red-tailed hawks (_Buteo borealis_), Swainson's hawks (_B. swainsoni_), and rough-legged hawks (_B. lagopus_ and _B. regalis_) might be seen wheeling majestically across the sky in the Plains States, and in the East the flights of broad-winged hawks (_B. platypterus_), Cooper's hawks (_Accipiter cooperi_), and sharp-shinned hawks (_A. velox_) are still occasionally seen, although these birds do not actually travel in flocks. To the birds of prey and possibly to the gulls also, a day's fasting now and then is no hardship, particularly since they frequently gorge themselves to repletion when opportunity is afforded.

The spring migrations of the blackpoll warbler (_Dendroica striata_) and the cliff swallow (_Petrochelidon albifrons_) afford an interesting comparison of the flights of day and night migrants. Both spend the winter in South America, at which season they are neighbors. But when the impulse comes to start northward toward their respective breeding grounds, the warblers strike straight across the Caribbean Sea to Florida, while the swallows begin their journey by a westward flight of several hundred miles to Panama (fig. 5). Thence they move leisurely along the western shore of the Caribbean Sea to Mexico, and continuing to avoid a long trip over water, they go completely around the western end of the Gulf of Mexico. This circuitous route adds more than 2,000 miles to the journey of the swallows that nest in Nova Scotia. The question may be asked, Why should the swallow select a route so much longer and more roundabout than that taken by the blackpoll warbler? The simple explanation is that the swallow is a day migrant while the warbler travels at night. The migration of the warbler is made up of a series of long, nocturnal flights, alternated with days of rest and feeding in favorable localities. The swallow, on the other hand, starts its migration several weeks earlier and catches each day's ration of flying insects during a few hours of aerial evolutions, which at the same time carry it slowly in the proper direction. Flying along the insect-teeming shores of the Gulf of Mexico, the 2,000 extra miles that are added to the migration route are but a fraction of the distance that these birds cover in pursuit of their food.

Although most of our smaller birds make their longest flights at night, close observation shows that travel is continued to some extent by day. This is particularly the case during the latter half of a migratory season, when the birds manifest a desire to hasten to their breeding grounds. At this time flocks of birds while feeding maintain a movement in the general direction of the seasonal journey. Sometimes they travel hurriedly, and while their flights may be short, they cover considerable distances in the course of a day.

=HOW BIRDS MIGRATE=

SPEED OF FLIGHT AND SPEED OF MIGRATION

There is wide-spread misconception concerning the speed at which birds normally fly, and even regarding the speed they can attain when occasion demands, as when closely pursued by an enemy. It is not unusual to hear accounts of birds flying "a mile a minute." While undoubtedly some birds can and do attain a speed even greater than this, such cases are exceptional, and it is safe to say that even when pressed, few can develop an air speed of 60 miles an hour. They do, however, have two speeds, one being the normal rate for everyday purposes and also for migration, and an accelerated speed for escape or pursuit; this in some cases may be nearly double the normal rate of movement. Nevertheless, the effort required for the high speeds could not be long sustained, certainly not for the long-distance migratory journeys that are regularly made by most birds.

The theory that migrating birds attain high speeds received encouragement from the German ornithologist Gätke, who for many years, made observations on birds at the island of Heligoland and published thereon in 1891 (_21_). He postulated that the blue-throat (_Cyanosylvia suecica_), a species of thrush smaller than the American hermit thrush, would leave African winter quarters at dusk and reach Heligoland at dawn, which would mean a sustained speed of 200 miles an hour; and that the American golden plover (_Pluvialis dominion_) flew from the coast of Labrador to Brazil in 15 hours, or at the tremendous speed of 250 miles per hour. These conclusions are now considered unwarranted by most ornithologists.

Sportsmen also often greatly overestimate the speed at which ducks and geese fly and sometimes attempt to substantiate their estimates by mathematical calculations, based upon the known velocity of a charge of shot, the estimated distance, and the estimated "lead" that was necessary to hit the bird. If all three elements of the equation were known with certainty, the speed of the bird could be determined with a fair degree of accuracy. The majority of the ducks that are reported as killed at 40, 50, or even 60 yards, however, actually are shot at distances much less than estimated. To sight along a gun barrel and estimate correctly the distance of a moving object against the sky is so nearly impossible for the average gunner as to make such calculations of little value.

During the past few years reliable data on the speed of birds have accumulated slowly. It has been found that the common flying speed of ducks and geese is between 40 and 50 miles an hour, and that it is much less among smaller birds. Herons, hawks, horned larks, ravens, and shrikes, timed with the speedometer of an automobile, have been found to fly 22 to 28 miles an hour, while some of the flycatchers are such slow fliers that they attain only 1 to 17 miles an hour. Even such fast-flying birds as the mourning dove rarely exceed 35 miles an hour. All these birds can fly faster, but it is to be remembered that at training camps during the World War, airplanes having a maximum speed of about 80 miles an hour easily overtook flocks of ducks that it may be supposed were making every effort to escape. Aviators have claimed that at 65 miles an hour they can overtake the fastest ducks, though cases are on record of ducks passing airplanes that were making 55 miles an hour.

The greatest bird speeds that have been reliably recorded are of the swifts (fig. 6) and the duck hawk, or peregrine falcon (_Falco peregrinus_) (fig. 7). An observer in an airplane in Mesopotamia reported that swifts easily circled his ship when it was traveling at 68 miles an hour. To do this, the birds certainly were flying at a speed as high as 100 miles an hour. Once a hunting duck hawk, timed with a stop watch, was calculated to have attained a speed between 165 and 180 miles an hour.

The speed of migration, however, is quite different from that attained in forced flights for short distances. A sustained flight of 10 hours a day would carry herons, hawks, crows, and smaller birds from 100 to 250 miles, while ducks and geese might travel as much as 400 to 500 miles in the same period. Measured as air-line distances, these journeys are impressive and indicate that birds could cover the ordinary migration route from the northern United States or even from northern Canada to winter quarters in the West Indies or in Central America or South America in a relatively short time. It is probable that individual birds do make flights of the length indicated and that barn swallows (_Hirundo erythrogaster_) seen in May on Beata Island, off the southern coast of the Dominican Republic, may have reached that point after a nonstop flight of 350 miles across the Caribbean Sea from the coast of Venezuela. Nevertheless, whether they continue such journeys day after day is doubtful.

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 1 or several days, particularly if they find themselves in congenial surroundings. Some indication of this is found in the records of banded birds. 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 (_Anas rubripes_) 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. The greatest speed thus far recorded for a banded bird is that of a mallard (_A. platyrhynchos_) banded on November 23, 1930, in Green Bay, Wis., and shot 5 days later, 900 miles away, near Georgetown, S. C. This bird doubtless flew at least 1,000 miles in the 5 days, as its route probably was not in a direct air-line, but, even so, the average daily distance was only 200 miles, which could easily have been covered in 5 hours.

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. 8). 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.