Territory in Bird Life

by Henry Eliot Howard

Here, however, I must refer to a view which is held by some psychologists, namely, that amongst the higher animals, even on the occasion of the first performance of an instinctive act, there is some vague awareness of the proximate end to be attained. Discussing the nature of instincts, Dr M'Dougall[3] says, "Nor does our definition insist, as some do, that the instinctive action is performed without awareness of the end towards which it tends, for this, too, is not essential; it may be, and in the case of the lower animals no doubt often is, so performed, as also by the very young child, but in the case of the higher animals some prevision of the immediate end, however vague, probably accompanies an instinctive action that has often been repeated." A similar view seems to be held by Dr Stout.[4] "As I have already shown," he says, "animals in their instinctive actions do actually behave from the outset as if they were continuously interested in the development of what is for them one and the same situation or course of events; they actually behave as if they were continuously attentive, looking forward beyond the immediately present experience in preparation for what is to come. They apparently watch, wait, search, are on the alert. They also behave exactly as if they appreciated a difference between relative success and failure, trying again when a certain perceptible result is not attained and varying their procedure in so far as it has been unsuccessful. All these characters are found in the first nest-building of birds as well as in the second; they are found also in courses of conduct which occur only once in the lifetime of the animal." Both these writers would, I imagine, contend that, even when a female is absent, the idea of the female, as the end in view throughout, is present; and they would argue that the fact of her absence during the fighting in no way disposes of the belief that she is the condition under which the pugnacious instinct of the male is rendered susceptible to stimulation. What reason is there to think that this interpretation is applicable to the case under consideration? When a female is present, we observe that the males are pugnacious, and, when she is absent, that they still continue to be hostile—that is to say, they behave as if she were present. Now, as far as I can ascertain, the "as if" is the only ground there is for supposing that the female is represented in imaginal form—there is no evidence of the fact, if fact it be. On the contrary, the behaviour of the male affords some fairly conclusive evidence that no such image is the primary factor in exciting the instinctive reaction. For if it be the actual presence of the female, or, in the absence of such, a mental image, that renders the pugnacious nature of the male responsive; provided the usual stimulus were present, the instinct ought surely to respond, not only under one particular circumstance, but under all circumstances. Yet, as we shall presently see, a male is by no means consistently intolerant of other males. It may be sociable at one moment or pugnacious at another, but the pugnacity is always peculiar to a certain occasion—the occupation of a territory. What shall we say then—that a mental image is a situational item only when the territory is occupied? It may be so; it may be that the fact of occupation gives rise to the mental image which, in its turn, renders the fighting instinct explosive, which again renders the possession of the territory secure. That such an interpretation is possible we must all admit. But if it were true, though it would not affect the main consideration, namely, whether the fighting has reference to the possession of a particular female, or to the protection of the territory, it would make further discussion as to which of these is the condition of the fighting unprofitable, for each would have its part to play in the process, the territory remaining, however, the principal factor in the situation.

Now the difference in the times of arrival of the male and female migrant varies in different species from a few days to a fortnight or even more. It is most marked in those that return to their breeding grounds early in the season, and the greater the margin of difference the greater scope is there for observation. In my records for the past twelve years, there are frequent references to these initial male contests in the life of the Willow-Warbler and of the Chiffchaff; and in the district which I have in mind, these two species arrive early in the season, the males preceding the females by a week or even as much as a fortnight. Suppose, then, that two Chiffchaffs establish themselves in adjoining territories; or suppose that a male settles in a territory already occupied; what is the result? Well, scenes of hostility soon become apparent; as the birds approach one another they become more and more restive, their song ceases, they no longer search for food in the usual methodical manner, but instead their movements are hurried and their call-notes are uttered rapidly—all of which betrays a heightened emotional tone. Then the climax is reached, there is a momentary fluttering of tiny wings, a clicking of bills, and for the time being that may be all. But unless one or other of the combatants retires, this scene may be repeated many times in the course of a few hours, and repeated with varying degrees of severity. Yet the fighting, even in the most extreme form, when the birds locked together fall slowly to the ground, is seldom of an impressive kind, and one has to bear in mind the capabilities of the actors, remembering that the most severe struggle might readily be interpreted as a game if it were not for certain symptoms which reveal its inner nature.

The males of many other migrants can frequently be observed to fight when there was every reason to believe that females had still to arrive. The Blackcap is notoriously pugnacious, but not more so than the Marsh-Warbler or the Whinchat. Here in Worcestershire, the Arundo phragmites grows mainly on certain sheets of water which are comparatively few and far between, and the Reed-Warbler is consequently restricted to isolated and more or less confined areas. The males arrive early in May before the new growth of reeds has attained any considerable height, and each one has its own position in the reed-bed, sings there, and throughout the whole period of reproduction actively resists intrusion on the part of other males. I have kept watch upon a small area of reeds daily from the date of the first arrival; each individual was known to me, and as the growing reeds were only a few inches in height, a female could scarcely have escaped detection. Yet time and again disputes arose, and males pursued and pecked one another, striving to attain that isolation for which racial preparation had fitted them.

But on account of their violence, or their novelty, or because the absence of a female was beyond question, some battles stand out in one's memory more prominently than others. An instance of this was a struggle between two Whitethroats which happened in the latter part of April and lasted for three successive days. The scene of its occurrence was more or less the same on each occasion, and the area over which the birds wandered was comparatively small. The fighting was characterised by persistent effort and was of a most determined kind, and so engrossed did the assailants become that they even fluttered to the ground at my feet. No trace of a female was to be seen at any time during these three days, nor, during the pauses in the conflict, was the emotional behaviour of a kind which led me to suppose that a female was anywhere in the vicinity. And, if she had been near, she must have made her presence known, for the belief that she is a timid creature, skulking on such occasions in the undergrowth, is by no means borne out by experience.

Even more impressive was a battle between two male Cuckoos. It occurred high up in the air above the tops of some tall elm-trees which roughly marked the boundary line between their respective areas, and the actions of the birds were plainly visible. At the moment of actual collision the opponents were generally in a vertical position, and wings, feet, and beaks were made use of in turn; one could plainly see them strike at one another with their feet, and one could observe the open bill which generally denotes exhaustion, but may of course have been due to anger, or used as a means of producing terror. Yet no female appeared in the locality until six days after the occurrence of this struggle—and she certainly is not easily overlooked, for her note is unmistakable even when the behaviour of the male does not betray her arrival.

That the actual presence of the respective females exercised any influence on the course of these struggles is more than doubtful. Not only did one fail to detect them, but one's failure to do so was confirmed by the knowledge that they had not yet arrived in those particular localities. Hence the fact of the male preceding the female is a valuable aid to the interpretation of subsequent behaviour; and one appreciates it the more after having experienced the difficulty of deciding whether she is present during the conflicts between resident males, for no matter how carefully we may observe the conditions which lead up to, and which accompany, such conflicts, or how closely we may scrutinise the surrounding trees, undergrowth, or ground, there always remains the possibility that she may, after all, have been overlooked. But this must not be taken to imply that in such cases direct observation alone can lead to no serviceable result, or that the evidence gained therefrom is worthless. Far from it. Failure to detect a female is so very common an occurrence that, even if we lacked the corroborative evidence supplied in the life of the migratory male, it would still be unreasonable to suppose that it were solely due to mistaken observation. We mark her absence during the conflicts between the respective males of many common species—the Finches, Buntings, and Thrushes that occupy their territories early in the season when the hedgerows and trees are still bare; but more frequently amongst those that inhabit open ground, because the movements of the birds are there more accessible to observation. For instance, half a dozen or more Lapwings can be kept in view at the same time, and as they stand at dawn in solitary state, keeping watch upon their respective territories, they are conspicuous objects on the short, frosted grass; no stranger can enter the arena without the observer being aware of it, no commotion can occur but one detects it, no movement however small need be missed. And so they fight, in a manner which leaves no doubt as to the reality of the struggle, when their prospective mates are absent not only from the particular territories in which the conflicts take place, but absent too from those adjoining.

If the fact that males fight before they are paired and in the absence of a female could be placed beyond all question, it would no longer be possible to regard her possession as the end for which they are contending, and consequently there would be no need to produce further evidence. But the examples which I have given refer, of course, to only a few migrants and a few residents—and moreover it must be admitted that a female is often conspicuous during the battles—so that by themselves they must be regarded, and rightly so, as inconclusive. We must therefore pass on to consider evidence of a somewhat different character.

I spoke of the complexity of the strife. By this I mean that it is not merely a matter of disputes between adjoining males, but that it is a far more comprehensive business involving both sexes. Thus female fights with female and pair with pair, or a male will attack a female, or, again, a pair will combine against a single male or a single female. And from all this complexity of strife we gain much valuable evidence in regard to the question immediately before us. For when one pair attacks another, or males that are definitely paired fight with one another, or an unpaired male attacks either sex of a neighbouring pair indiscriminately, there is surely little ground for supposing that the possession of a mate is the reason of it all.