Territory in Bird Life

by Henry Eliot Howard

This instinct controls the movements of many birds from early autumn to the commencement of the breeding season. And so powerful is the control that the individual is suppressed and its activities subordinated to the welfare of the community as a whole. Flocks of Waders roam about the tidal estuaries in search of food, and different kinds of Gulls assemble there and preen their feathers or sleep; Warblers alter their mode of life, and in the osier bed, or amongst the elders, seek their food together in peace; Finches, Buntings, Pipits, and Wagtails, though food is everywhere abundant, gather themselves together respectively into bands which, as winter approaches, grow into flocks and even into composite flocks; and as the Warblers leave for the south, so their places are filled by flocks of Thrushes and Finches from the north. In whatever direction we turn, when the days begin to shorten, it is the community, not the individual, that thrusts itself upon our attention; and throughout the winter continues to be the outstanding feature of bird life.

With the approach of the breeding season we witness that remarkable change which I have endeavoured to make clear in the previous chapters—the disintegration of the flock and the reinstatement of the individual. Instead of continuing with the flock, the individual now goes forth to seek the appropriate breeding ground; and having arrived there, is not only content to remain in isolation, but so behaves that isolation is insured. Intolerant of the approach of a stranger, intolerant even of the approach of the very members of the community whose companionship was previously welcomed, it not only fights to maintain the position it has selected, but fights indeed for the possession of ground already occupied, and, until reproduction is completed, asserts its individuality and exercises dominion over its territory. What, then, is the prospective value, biologically considered, of the changing interest that A displays in B, and to what will such changes lead? These are the questions to which we will now direct inquiry.

The annual life-history of a bird is in broad outline conditioned by two powerful and at first sight opposing impulses—the one to live in society, the other to live solitary. But, manifestly, a bird cannot be governed by opposing impulses. It has but one character, within which, according to the season and the circumstances, different impulses predominate. But these impulses, no matter how different they may appear to be, have their respective parts to play in furthering the life of the individual. Hence they cannot oppose, though they may conflict, if the resultant behaviour contributes towards survival.

The majority of birds live to-day in constant danger from predatory species, and that this danger was still greater in bygone ages there can be but little doubt. A curious mode of behaviour of the Curlew, Whimbrel, and Godwit demonstrates this, for it must be the outcome of the necessity for constant watchfulness. Whilst resting with its head turned back and its beak buried in the feathers of the mantle, the bird constantly moves the axis of its body, so that an observer, if placed in a direct line behind it, sees at one moment the right eye and at another the left. No movement of the feet or of the legs is perceptible, and the shifting of the body continues whether the eyes are open or closed. This body movement enables the bird to survey a much larger area of ground than it would otherwise be capable of doing, and thus adds to its security. As far as my experience goes, the movement is less evident amongst the members of a flock than when an individual is resting alone, or even with a few companions, which may be due to the fact that since some members are always awake and watchful, a bird of prey would have more difficulty in approaching a flock unawares than it would have in approaching a single individual. With the greatest ease a Sparrow-Hawk can pick up a Thrush as it feeds on the meadow by itself, but if it attempts to seize one of a flock, the chances are that its approach is signalled and that its prey escapes. And not only do the different members give warning one to another of the approach of danger, but they also combine to harass or even to drive away an enemy. So that there can be no doubt that the gregarious instinct is serviceable in promoting the welfare of the race, and has, as its end, the preservation of the individual in order that it may take its share at the appropriate time in procreating its kind.

In winter, then, the individual loses its individuality and is subordinated to the welfare of the community, whilst in spring it regains its individuality, and all its inherited instincts which then come into operation lead to its isolation from the flock. The impulse to seek isolation is dependent upon internal organic conditions which are peculiar to a certain season; whereas the gregarious impulse depends upon internal organic conditions which inhere at all times, though its functioning is inhibited by the functioning of the former impulse. The evidence which leads to this conclusion is to be found in the fact that a male often deserts its territory temporarily and joins the flock, where it remains at peace with its companions—an aspect of behaviour which we have discussed on various occasions. The former impulse becomes dominant in the spring owing to its innately superior strength; the latter becomes dominant in the autumn because the organic condition which determines the functioning of the former then subsides. The impulse to seek the appropriate breeding ground and to dwell there would seem to be the strongest of all the impulses save one—the sexual. When, however, I speak of the sexual, I refer to the actual discharge of the sexual function, which is the consummation of the whole process. But the territory and all that appertains to it is part of that process—the search for the breeding ground, the dwelling there, and the intolerance of intrusion are but different stages, each one of which must have an impulse peculiar to it; and since the completion of the sexual act can only be successfully accomplished providing that success is attained at every stage, the probability is that, of the impulses concerned, one is neither more powerful nor less powerful than another.

So that we have two impulses operating at different seasons and guiding the behaviour into widely divergent channels. But though the proximate end to which the behaviour is directed is apparently different, there are not two biological ends in view, but one—the attainment of reproduction; and the changes that we witness are not contrary but complementary, and their prospective value lies in the circumstance that they contribute towards the preservation of the race.

If, then, every male is driven by inherited impulse to seek the appropriate breeding ground each recurring season; if, having arrived there, it is driven to seek a position of its own; if, in order to secure isolation it is obliged to attack other males or to ward off the attacks of intruders; if, in short, success can only be attained providing that the inherited nature is so adjusted that the bird can accomplish all that is here demanded—what will be the general result? That the individual will rear its offspring in safety and that they will inherit the peculiarities of their parents, enabling them, in their turn, to procreate their kind; all this will certainly follow. We are not concerned, however, at the moment, with the direct effect upon the individual, but with the consequences that will accrue to the species as a whole.

Now certain facts are presented to observation which enable us not only to understand the nature of the change that is wrought in the history of the species, but to foreshadow, with no small degree of certainty, the extent of that change. I suppose that it has come within the experience of most of us to observe, at one time or another, the ebb and flow of a given species in a given district. Some favourite haunt is deserted for a year, or for a term of years, and is then revisited; or, if it is always occupied, the number of inhabitants fluctuates—plenty of pairs in this season, only a few in that. Many intricate relationships, both external and internal, contribute towards this state of affairs. Fluctuation in a downward direction, or temporary extinction, is brought about by changes in the physical world, by changes in the available supply of food, by the increase of enemies, or by adverse climatic conditions; whilst fluctuation in an upward direction, though due indirectly to a combination of circumstances in the external world favourable to the survival of large numbers of individuals, is directly determined by the impulse to seek isolation. As individuals of different species establish themselves, and form kingdoms and lesser kingdoms, we can watch the gradual quickening into life of moorland and forest and we can observe the manner in which it all comes to pass. Males that for weeks or months have lived in society, drifting from locality to locality according to the abundance of food or its scarcity, now set forth alone and settle first here and then there in search of isolation. Lapwings settle in the water meadows, and, finding themselves forestalled, pass on in search of other ground; Blackbirds arrive in a coppice or in a hedgerow and, meeting with opposition, disappear; and the Curlew, wandering with no fixed abode but apparently with a fixity of purpose, searches out the moorland where it can find the particular environmental conditions to which its inherited nature will respond. In fact, wherever we choose to look, we can observe in a general way the gradual appropriation of breeding ground; and if we fix our attention upon particular males, we can watch the method by which success or failure is achieved.

On more than one occasion I have watched the efforts of Reed-Buntings to appropriate territories in a marsh that was already inhabited. Sometimes their efforts met with success, at other times with failure. In the former case, the males, whose ground was intruded upon, were severally forced to yield part of their holding and were thus left in possession of a smaller area. The success of the intruder seemed to depend upon persistent determination, rather than upon superior skill in battle. Recently I had an opportunity of observing the intrusion of a male Willow-Warbler upon ground already occupied. By persistent effort it succeeded in appropriating one half of the territory of its rival. The intruder occupied some trees on the outskirts of the territory it was invading, and used them as a base from which it made repeated efforts to enter the ground of its rival. These efforts were time after time frustrated. No sooner did it leave its base than it was seen and intercepted, or else attacked; and no matter from which direction it attempted to effect an entrance, its efforts, for a time, were all to no purpose. The fighting was of a determined character, and after each attack the owner of the territory showed signs of great excitement, and, sitting upright upon a branch, spread and waved its wings, which is the specific emotional manifestation during the period of sexual activity. Eventually the intruding male succeeded by persistent effort in appropriating part of the occupied ground.

Thus we can actually witness the efforts of the individual to isolate itself from members of its own kind, and can observe the immediate consequences that follow from success or from failure. And from these consequences we can infer that, within a certain range but in accordance with the relative abundance of the species that dwell in it, every corner of the available breeding ground will be explored and every situation that evokes the appropriate response will be occupied. Moreover, since the annual dispersion is not merely a repetition in this season of that which occurred in a previous one, a progressive increase in the area occupied will follow. Yet, if the majority of species desert their breeding ground so soon as reproduction is ended, how can this be? An answer to the question will be found in the fact that a bird has an innate capacity to return to the neighbourhood of its birthplace, or to the place wherein it had previously reared offspring—which means that the results of prior process persist as the basis and starting-point of subsequent process.