It has been dogmatically asserted repeatedly that birds invariably breed in the most northerly part of their range, and winter in the most southerly. Winter, when speaking of Holarctic birds, only applies to the season in the northern hemisphere; the birds which pass south of the equator winter in summer. Whilst accepting this as a rule, two reservations must be made. First, that it only applies to birds of the northern hemisphere, and secondly that it is a rule with exceptions. It seems probable that the breeding area of some of the birds which reach the British Islands in autumn by the so-called east and west route is in more southerly latitudes than our islands, and certainly it seems evident that the temperature of the winter refuge has more effect upon the birds than its geographical position. Perhaps the statement that a bird always nests in the coldest part of its range is more universally correct. Even this may not be invariably the habit, but in acknowledging it as a rule we must clearly understand that this cold district is resorted to at the period of the year when its temperature is at its highest. There are certain birds which breed in Australia and winter in Oceanic islands where the temperature is cooler than in their breeding area.
When considering the migration of birds which summer in the extreme north or breed in the extreme south—alas, but little is known about the migratory habits of many southern breeders—it is comparatively simple to offer an explanation; in the long winter months this home, so desirable in the short weeks of daylight, is dark, ice-bound, and foodless; it is wholly unsuited to the requirements of birds, which, in spite of many assertions to the contrary, have never been proved to hibernate, the only way in which animals can survive for any lengthened period when food supply is entirely cut off.
Birds are structurally provided with the means of escaping from the disastrous effects of adverse circumstances; the power of flight, though not the only way in which animals can migrate, is at the root of the migration of birds. The advantages of the power of flight, to which also it owes its development, include the ability to avoid active and passive enemies, and to remove from one feeding ground to another undeterred by the barriers which restrict the terrestrial animal. A natural sequence of this ability to take advantage of aerial locomotion is the habit of wandering in search of food, more or less noticeable in all birds. The habit of wandering led to the discovery of feeding grounds and suitable nesting places; where these nesting places, probably at first, only removed a short distance from the parents' nesting site, were suitable, dispersal and an extension of the distributional area or range of the species followed; but where the feeding area was unsuited or not so well suited to the needs of the species, hereditary attachment to the original home and memory of the direction of this home, or even in some cases accidental wandering back to the more suitable locality, would originate a migration. Coupled with this are two important factors which would tend to make the habit periodical and regular both as regards time and locality. The memory of the bird, call it instinctive memory if we like, would limit the wanderings in search of food to a certain number of places where food was most abundantly found, and the passage between feeding area and breeding area become regular journeys, at the seasons of the year when an increasing number of young birds in the breeding area drove the overgrown population to seek food further from the base, and again when the sexual impulses urged the birds to seek secure nesting sites. The other factor is the weeding-out influence of mistaken effort, the natural selection which leads to the survival of the fittest. The young wanderer which reached unsuitable lands must either wander further or perish. Judging by the juvenile mortality amongst young birds the failures would be many, and only the successful competitors would return to leave progeny.
Great stress has been laid on the attachment of birds to certain nesting sites, an undoubted fact, and it has been argued that because, in some cases, for hundreds of years certain sites have been occupied by the same species, it is evident that after the death of parents the young will return to and occupy the home. This has even been put forward as evidence that birds do not wander in search of fresh nesting sites. The argument is not sound. It is improbable that in most cases both parents perish in the same year. Birds of prey, and many of the cited instances of long tenancy refer to raptorial birds, have a wonderful power of finding a mate, male or female, to complete the hatching and rearing of the young, when one of a pair has been destroyed. The survivor of any pair might have the home attachment and by bringing a fresh mate create an attachment which would be passed on from mate to mate indefinitely. Again it must not be overlooked that certain sites present advantages to particular species which must be evident to all in search of those advantages; it by no means follows that the occupiers of a nesting site are in any way related, except specifically, to those which occupied it in previous years.
The answer to the argument that birds do not seek fresh nesting places and thus extend their distributional area, is evident when we consider those species which, at the present time, are extending their range. Within the last few years, for instance, the turtle dove and tufted duck have begun to nest regularly in many parts of England in which they were entirely unknown twenty or thirty years ago. The starling has spread and in some parts is spreading still, and many other similar cases might be cited.
In this manner migration, as we know it to-day, may have originated, and as Mr P. A. Taverner expressed it, "however instinctive their habit may now be, there must have been a time when migrations were intelligent movements, intended to escape some danger or secure some advantage" [(51)]. Granting this, however, as the first cause, we are only on the threshold; the question still remains unanswered, what actually impels the birds to seek fresh food supplies or to look for safe nesting places? The natural answer, the cravings of nature and sexual impulses fails to give satisfaction in every case. Wanderings in search of food might lead in any direction, and probably did in the first place, but now birds in the main travel south in search of food and north in search of home, and many of them perform immense journeys, passing over or through lands which are capable of supporting a wealth of bird-life even in the winter months.
The majority of Arctic birds or those nesting in high latitudes leave before the great harvest of autumn fruits, and even our common swift begins to depart—for all do not go at once—towards the end of July, when insects are more abundant than at any other time of the year. Food supply has not failed when most birds start their journey in search of food! Again in spring, when it is claimed that the powerful sexual impulses are sufficient reason to account for the northward journey, hosts of sexually immature birds and of others which are apparently mature but do not breed that spring, migrate northwards, some even arriving before the mature birds of their own species.
The earlier students of migration insisted that temperature was the sole cause of change of abode; that the northern lands became unsuitable through their falling temperature, and that the birds deserted them for warmer climes, returning when the lands they wintered in became too hot. As a variant of this notion, which cannot be lightly cast aside, the suggestion was mooted that it was not cold but the lack of food during the cold months which drove them south, and that in the Tropics, where at one time it was thought that all migratory birds wintered, food was scarce during the months of extreme heat. Dr. Wallace went further and stated that the incentive to northern migration was the inability to find sufficient soft bodied insects suitable for the nestlings in the Tropics during summer [(54)]. Yet there are birds which do find food enough for their young, and some of them are insect eaters.
Seebohm, arguing with reason that the first home of the Charadriidae, was the Polar Basin [(44)], suggests that the desire for light originated the idea or the action, and though this was only applied by him to Arctic birds, others have striven to show that the longer hours of daylight would be an advantage to all birds, even though the difference of dark and light in the zone retired from and in that arrived at might be inconsiderable [(41)]. Against this must be taken into consideration the fact that many waders and ducks, northern breeders, feed by night or day, according to the state of the tide. Light is not an absolute necessity to them.