Territory in Bird Life

by Henry Eliot Howard

If, then, there is in the call-notes an adequate means of communication and of recognition, why do I suggest that the song has also been evolved primarily for the purpose of recognition?

What, first of all, are the conditions in the life behaviour during the season of reproduction that make the intervention of the voice a consideration of such importance? The general result of our investigation might be summed up thus: we found that the male inherits a disposition to secure a territory, that at the proper season this disposition comes into functional activity and leads to its establishment in a definite place, and that it cannot search for a mate because its freedom of action in this respect is forbidden by law; that the female inherits no such disposition, that she is free to move from place to place, free to satisfy her predominant inclination, and to seek a mate where she wills; and, since the appropriate organic condition which leads to pairing must coincide with appropriate conditions in the environment, that the union of the sexes must be accomplished without undue delay. Furthermore we found that a territory is essential if the offspring are to be successfully reared; that, since the available breeding ground is limited, competition for it is severe, and that the male is precluded from leaving the ground which he has selected, and is obliged, in order to secure a mate, to make himself conspicuous. That was our general result. Now there are two ways by which the male can make himself conspicuous—by occupying such a position that he can be readily seen, or by producing some special sound which will be audible to the female and direct her to the spot. The former, by itself, is insufficient; in the dim light of the early dawn, when life is at its highest, and mating proceeds apace, what aid would it be for a male to perch on the topmost branch of a tree, how slender a guide in the depth of the forest? But whether in the twilight or in the dark, in the thicket or the jungle, on the mountain or on the moor, the voice can always be heard—and the voice is the principal medium through which the sexes are brought into contact.

Well now, we come back to the question, why, if all species have a serviceable recognition call, that call should not be sufficient for the purpose, just as, without a doubt, it is adequate for all purposes at other seasons? The answer is, I think, clear. The recognition call is not confined to one sex, nor only to breeding birds; it is the common property of all the individuals of the species, and if the female were to rely upon it as a guide she might at one moment pursue another female, at another a non-breeding male; she might even be guided to a paired female or to a paired male, and time would be wasted and much confusion arise. So that no matter how much a male might advertise himself by cries and calls which were common alike to all the individuals of the species, it would not assist the biological end which we have in view. Something else is therefore required to meet the peculiar circumstances, some special sound bearing a definite meaning by which the female can recognise, amongst the host of individuals of no consequence to her, just those particular males in a position to breed and ready to receive mates. Hence the vocal powers, the power of producing sounds instrumentally, and the power of flight, have been organised to subserve the biological end of "recognition."

And this view is strengthened, it seems to me, by the erratic behaviour of certain species, more particularly by one remarkable case, the case of the Cuckoo. The male, after having established himself, utters his call persistently from the day of arrival until approximately the middle of June; but, in contrast with the large majority of species, the female has a characteristic call which she, too, utters at frequent intervals. The female is polyandrous and has a sphere of influence embracing the territories of a number of males; she wanders from place to place, is often silent, and not unfrequently is engaged in dealing with her egg or in searching for a nest in which to deposit it, and therefore she is not always in touch with a male, still less with any particular one. Now there is much evidence to show that the discharge of the sexual function amongst birds is subject to control, and that this control operates through the female—through her physiological state becoming susceptible to stimulation only at certain periods. So that we have these considerations, that the female is polyandrous, that she has a territory distinct from that of the male, and that her sexual impulse is periodical; and the further consideration that the impulse, since it is periodical, is of limited duration and must receive immediate satisfaction. Such being the circumstances of the case, would the voice of the male serve to insure the union of the sexes at the appropriate moment? Well, the fact that she is polyandrous implies that every male in her sphere of influence is not always capable of satisfying her sexual instinct. Is, then, the male's call an indication of his readiness to yield to stimulation? Without a doubt it is an index of the general physiological state which generates the sexual impulse, without a doubt it denotes a general preparedness to breed, but there is no evidence to show that it denotes the degree of ardour of the male at any particular moment, and much that proves the contrary. So that only by the female producing some special sound which will attract the males that are eager and bring them rapidly to the spot where she happens to be, only thus is it possible to insure the consummation of the sexual act. This, it seems to me, is the purpose of the peculiar call of the female—a call which, so far as biological interpretation is concerned, is just as much a song as the melody of the Marsh-Warbler—and its interest for us just now lies in this, that here we have a special case in which the sexes have separate territories, the female is polyandrous, and the voice of the male is not sufficient by itself to bring to pass the union of the sexes; and in which, consequently, if the purpose of song be that of recognition, we should expect to find, as we do find, that the female had a distinct and penetrating call.

We now come to the question of "warning," by no means the least important purpose of song. I pointed out that one of the chief differences between the call-notes and the song was that the former were socially serviceable, whereas the latter was only serviceable to certain individuals; and in making this statement, I had in mind the direct benefits to the community which proceeded from an appreciation of sounds having a mutually beneficial meaning, not the indirect, though none the less beneficial, consequences to the species as a whole. Biologically considered, song, if it acts as a warning and thereby leads in one way or another to more complete success in the rearing of offspring, may be spoken of as socially serviceable; but it is legitimate to draw a distinction between the prospective value of remote relationships which we can foresee, and the mutual assistance which the individuals of a community derive from their close association.

If there were always sufficient breeding ground to support the offspring of all the individuals of each species, if the individuals were always so distributed that there was no possibility of overcrowding in any particular area, and if the conditions of existence of different species were so widely divergent that the presence of this one in no way affected the interests of that, no opportunity would be afforded for the development of so complex a system as is involved in the "territory" and all that appertains to it. But the available breeding ground is by no means unlimited. The supply of food, which is a determining factor in the environment, is always fluctuating according to the climate and according to the changes in the earth's surface; and so the distribution of the bird population in any given area, though it may be suitably adjusted for one year or even for a period of years, is bound in the course of time to require readjustment. Now there cannot be readjustment without competition, nor competition without combat. But the appeal to physical force is only a means to an end, and, since no male can endure incessant warfare and the perpetual strain of always being on the alert, without experiencing such physical exhaustion as might affect his power of reproduction, its direct effect upon the combatants cannot be otherwise than harmful—in fact it is a necessary evil which for the good of the species must be kept strictly within bounds. Bearing in mind, then, these two facts, namely that the distribution of the males is never stable and that overmuch fighting may defeat the end in view, we can appreciate the importance of any factor which will lead to a more uniform distribution and at the same time insure security by peaceable means.

The proximate end of the male's behaviour is isolation—how is it to be obtained? If, after having occupied a territory, the bird were to remain silent, it would run the risk of being approached by rivals; if, on the other hand, it were merely to utter the recognition call of the species, it would but attract them. In neither case would the end in view be furthered, and isolation would solely depend upon alertness and the capacity to eject intruders. Supposing, however, that the song, just as it serves to attract the females, serves to repel other males, a new element is introduced deserving of recognition; for those males that had established themselves would not only be spared the necessity of many a conflict, but they would be spared also the necessity of constant watchfulness, and so, being free to pursue their normal routine—to seek food, to rest, and, if migrants, to recover from the fatigue of the journey, they would be better fitted to withstand the strain of reproduction; and those that were still seeking isolation in an appropriate environment, instead of settling first here and then there only to find themselves forestalled, would avoid and pass by positions that were occupied, establishing themselves without loss of time in those that were vacant. Without the aid of something beyond mere physical encounter to regulate dispersal, it is difficult to imagine how in the short time at disposal anything approaching uniformity of distribution could be obtained. Hence, both in the direction of limiting combat, of insuring accommodation for the maximum number of pairs in the minimum area, and of conserving energy, the song, by conveying a warning, plays an important part in the whole scheme.

And if this be so, if the song repels instead of attracting, it follows that the more distinct the sounds, the less likelihood will there be of confusion; for supposing that different species were to develop similar songs, whole areas might be left without their complement of pairs just because this male mistook the voice of that, and avoided it when there was no necessity for doing so. So that just as from the point of view of "recognition" each female must be able to distinguish the voice of its own kind, so likewise the warning can only be adequate providing that the sounds are specifically distinct. A point, however, arises here in regard to closely related forms. Some species require similar food and live under similar conditions of existence; they meet in competition and fight with one another; and, if they did not do so, the food-supply of a given area would be inadequate to support the offspring of all the pairs inhabiting that area. Generally speaking, the more closely related the forms happen to be, the more severe the competition tends to become; and it may be argued that in such cases a similar song would contribute to more effective distribution and in some measure provide against the necessity of physical encounter; that, in fact, it would stand in like relation to the success of all the individuals concerned, as does the song to the individuals of the same species. But we must bear in mind that the primary purpose of song is to direct the females to those males that are in a position to breed; and to risk the possibility of prompt recognition in order that the males of closely related species should fight the less, would be to sacrifice that which is indispensable for a more remote and less important advantage.

What meaning does the song convey to a male that is unestablished? Does the bird recognise that it is forestalled; does it foresee and fear the possibility of a conflict, and conclude that the attempt to settle is not worth while? I do not imagine that it thinks about it at all. How then does the warning warn? We will endeavour to answer this question, but, in order to do so, we must review the stages by which a territory is secured.

We take as our starting point the internal organic changes which are known to occur. These changes are correlated with other changes, manifested by a conspicuous alteration in behaviour—to wit, the disappearance of sociability and its replacement by isolation. Having found a station which meets the requirements of its racial characteristics, the male establishes itself for a season, becomes vociferous, displays hostility towards others of its kind, and in due course is discovered by a female. The whole is thus an inter-related whole, a chain of activities which follow one another in ordered sequence. Now we have seen that it is neither pugnacious nor vociferous until the territory is actually occupied; we have seen that the fact of occupation is the condition under which the instincts of pugnacity and of song are rendered susceptible to appropriate stimulation; we have discussed the nature of the stimulus in each case, and we wish to know the sort of meaning that the song conveys to an individual which is still in the preliminary stage of seeking a station. In sequential order we have the following: (1) internal organic changes which lead to isolation, (2) the appropriate environment which gives rise to an impulse to remain in it, (3) the occupation of a territory which is the condition under which the instincts are rendered susceptible to stimulation, (4) the various stimuli. Each is dependent upon that which precedes it, and no part can be subtracted without failure of the biological end in view, neither can the different stages be combined in different order. So that, in considering the significance of song to an unestablished male, we are dealing with the situation at a point at which all the latent activities have not been fully felt, for all that so far has occurred is the change from sociability to isolation determined by internal organic changes. The bird has not established a territory because it has not come into contact with the appropriate environment, and it is not pugnacious because the condition which renders its instinct susceptible is absent; and so, as it wanders from place to place and hears the voices of males here or males there, it merely behaves in accordance with that part of its nature which predominates just at that particular moment—the impulse to avoid them.