Before presenting some of the present theories concerning the origin of bird migration, it seems well to consider briefly the ends served by this annual round trip between breeding grounds and winter quarters. It is apparent that the migratory habit enables a species to enjoy the summers of northern latitudes while avoiding the severity of the winters. In other words, migration makes it possible for some species to inhabit two different areas at seasons when each presents favorable conditions. In the performance of its reproductive duties every pair of birds requires a certain domain, the extent of which varies greatly in different species. Generally, however, this territory must be large enough to provide adequate food, not only for the parent birds but also for the lusty appetites that come into being with the hatching of the eggs. Thus, if all birds were to remain constantly either in tropical or in temperate regions, there might be intolerable overcrowding during the breeding season. By the spring withdrawal to regions uninhabitable earlier in the year, the migrants are assured of adequate space and ample food upon their arrival in the winter-freed North, and it may be assumed that nonmigrant species resident in the South are benefited by the withdrawal of the migrants.
Nevertheless it cannot be said that the winter or summer areas are entirely unsuited to the requirements of every migrating species at other seasons, for some individuals pass the winter in areas that are frequented only in summer by other individuals of their species. The extensive breeding ranges of such species present wide climatic variations, so that some individuals may actually be resident in a region where others of their kind are present only in winter.
The tendency in many species to move southward at the approach of winter is not always due to the seasonal low temperatures, since experiments have demonstrated that many summer insect feeders, when confined in outdoor aviaries, comfortably withstand temperatures far below zero. The main consideration is the depletion of the food supply, caused either by disappearance or hibernation of insects, or by the mantle of snow or ice that prevents access to the seeds and other forms of food found on or close to the ground or submerged in water. Possibly also the shortened hours of daylight materially restrict the ability of the birds to obtain sufficient food at a time when the cold requires an increased supply to maintain body heat. It is noteworthy that chickadees (Penthestes atricapillus) and some other of our smaller birds have no fear of Arctic weather, as their food supplies are mainly arboreal and so are always available. Also, when there is a good supply of food in the form of pine seeds in Canadian woods, nuthatches (Sitta carolinensis and S. canadensis) and crossbills (Loxia curvirostra and L. leucoptera) will remain through the winter. When these birds appear abundantly in winter at points in southern latitudes, it may be concluded that there is a shortage of their food in the North, or that they have been lured farther south by the greater abundance of this food there.
THEORIES OF THE CAUSES OF MIGRATION
Migration has long since become a definite hereditary habit that recurs in annual cycles, probably because of physiological stimuli associated with the reproductive period. In seeking its origin it is necessary to study the history of the birds' occupation of their present ranges, and from the information available to consider what appear to be reasonable theories. The two now most commonly accepted are diametrically opposed to each other.
NORTHERN ANCESTRAL HOME THEORY
According to one of these, nonmigratory birds swarmed over the entire Northern Hemisphere in earlier ages, the conditions of food and habitat being such as to permit them to remain in their haunts throughout the year. The entire northern area then afforded the two important avian requirements—suitable breeding conditions and a year-long food supply. This is the condition today in the Tropics, and it is noteworthy that many tropical birds are nonmigratory. Gradually, however, in the Northern Hemisphere the glacial ice fields advanced southward, forcing the birds before them, until finally all bird life was concentrated in southern latitudes. As the ages passed and the ice cap gradually retreated, each spring the birds endeavored to return to their ancestral homes in the North, only to be again driven south at the approach of winter. As the size of the ice-covered area diminished, the journeys made became ever longer, until eventually the habits of migration were fixed to accord with the climatic conditions of the present age.
Thus, this theory supposes that today migratory birds follow the path of a great racial movement that took place in a distant past, associated with advances and recessions of the ice. The actions of the birds themselves lend some support to this theory, as every bird student has noted the feverish impatience with which certain species push northward in spring, sometimes advancing so rapidly upon the heels of winter as to perish in great numbers when overtaken by late storms. It is probable, however, that at that season the reproductive impulse urges the birds on to their northern breeding grounds.
SOUTHERN ANCESTRAL HOME THEORY
The opposing theory is simpler in some respects and supposes that the ancestral home of all birds was in the Tropics and that as all bird life tends to overpopulation there was a constant effort to seek breeding grounds where competition would be less keen. Species that strove for more northern latitudes would be kept in check by the ice and forced to return southward with the recurrence of winter conditions. As the ice retreated, vast areas of virgin country became successively suitable for summer occupancy, but the winter habitat remained the home to which the birds returned after the nesting season. It is a fact that some species spend very little time on their breeding grounds; the orchard oriole (Icterus spurius), for example, spends only 2½ months in its summer home, arriving in southern Pennsylvania about the first week in May and leaving by the middle of July.