Henry Eliot Howard

“ The various steps by which the territory is not only established but made secure from invasion, imply an inherited nature nicely balanced in many directions—first of all the male must be so attuned as to be ready to search for a territory at the right moment; then it must be capable of selecting a suitable environment; and, having established itself, it must be prepared to defend its area from a rival, and to resist encroachment by its neighbours—and if it failed in any one of these respects, it would run the risk of failure in the attainment of reproduction. ”

“ But what is the condition under which the pugnacious instinct of the female is rendered susceptible to stimulation? It cannot be merely the presence of a male ready to breed, for then there would be endless commotion amongst the flocks of Chaffinches or of Lapwings which in March are composed of both sexes, including even males that have secured territories. There must be some other circumstance; and, judging by experience, it is to be found in the territory—a male, that is to say, in occupation of one, is the condition under which the inherited nature of the female is allowed free play. ”

“ We must remember, however, that fighting continues throughout the whole period of reproduction. Even after the discharge of the sexual function has ceased, and the female is engaged, in incubation or in tending her young, the male is still intolerant of intruders; and it is difficult to believe that, at so late a stage in the process, a female could be any attraction sexually to an unpaired male. But apart from any theoretical objection, there remains the fact—namely that there is no evidence that a male, after having once paired, is liable to be robbed of its mate. ”

“ A male intrudes, and the intrusion evokes an immediate display of irritation on the part of the owner of the territory, who, rapidly uttering its song and jerking its wings, begins hostilities. Flying towards the intruder, it attacks viciously, and there follows much fluttering of wings and snapping or clicking of bills. At one moment the birds are in the tree-tops, at another in the air, and sometimes even on the ground, and fighting thus they gradually approach and pass beyond the limits of the territory. ”

“ 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. ”

“ The fighting is thus one link in a chain of events whose end is the attainment of reproduction; it is a relationship in an inter-related process, and to speak of it as being even directly related to the territory is scarcely sufficient, for it is intimately associated with the disposition which is manifested in the isolation of the male from its companions, and forms therewith an imperium in imperio from which our concept of breeding territory is taken. ”

“ Or suppose that different pairs of Kittiwake Gulls on the crowded ledges, or different pairs of Puffins in the crowded burrows, varied in like manner, would they all have equal chances of rearing their offspring? The struggle for reproduction is nowhere more severe than amongst the cliff-breeding sea birds; it is not for nothing that one sees Kittiwake Gulls, locked together, fall into the water hundreds of feet below and struggle to the point of exhaustion, or, as has been reported, to the point of death; it is not for nothing that Puffins fight with such desperation. ”

“ But the warfare extends in a variety of directions, it is not confined to one sex, nor to unpaired individuals, nor need the opponents necessarily be of the same sex; it involves both sexes alike singly or combined. Now the view that the biological end of battle is, in its primary aspect, related to the female, cannot, as we have seen, apply to the conflicts between different pairs, and only by much stretching of the imagination can it be held responsible for the hostility that males frequently display towards females or vice versa. It is valid only for a certain form of warfare. ”

“ A feature which marks all the fighting, and which we cannot afford to disregard, is the conative aspect of the behaviour of the owner of the territory. The bird attacks with apparent deliberation as if it were striving to attain some definite end. ”

“ I suppose, that they are vestigial—fragments of warfare that determined the survival of the species in bygone ages. But if the conclusion at which we have just arrived be correct, if we can recognise a single aim passing through the whole of the warfare—and that one the removal of an intruder from a certain position, then we need no longer concern ourselves as to the degree of severity of the battles—we see it all in true perspective. Neither exhaustion nor physical inability are the sole factors which determine the nature and extent of the fighting ”

“ In the previous chapter I endeavoured to show that each male establishes a territory at the commencement of the breeding season, and there isolates itself from members of its own sex. And further I gave my reasons for believing that this particular mode of behaviour is determined by the inherited nature of the bird, and that we are justified in speaking of it as "a disposition to secure a territory" because we can perceive its prospective value. But the act of establishment is only one step towards "securing." By itself it can achieve nothing ”

“ Just as, in the securing of a territory, the ultimate appeal is to the physical strength of the male, so, in the course of her search for a mate, the female may be called upon to challenge, or may be challenged by a rival, and the issue is decided by force. ”

“ 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. ”

“ We were impressed, it may be remembered, with the contrast between the general behaviour of the birds at the opposite ends of the meadow. But now it appears as if the contrast were not between this individual and that, but between the behaviour of the same one under different circumstances. The male, that is to say, which, while in its territory, tolerates the approach of no other male, flies to the flock and is there welcomed by the very individuals with whom a short time previously it had been engaged in serious conflict. ”

“ Thus two pairs approach one another, and, when they meet, throw themselves upon their backs, each bird striking at its adversary with its feet or seizing hold of it with its beak; and though, in the commotion that ensues, it is almost impossible to determine what exactly is happening, there is reason to believe that the sexes attack one another indiscriminately. ”

“ Each individual has its own particular territory, embracing a piece of open water as well as a part of the rush-covered fringe, within which it moves and lives. But in the early part of the season, when the territories are still in process of being established, and definiteness has still to be acquired, trespassing is of frequent occurrence, and the conflicts are often conspicuous for their severity. ”

“ And so, by the time the young were hatched, the parents would be in occupation of an area too limited in extent to insure the necessarily rapid supply of food, and would be compelled to intrude upon the adjoining ground. But knowing how routine becomes ingrained in the life of the individual, knowing that for weeks this pair had submitted to their neighbours, can we believe that they would be capable of asserting their authority and that the young would be properly cared for? ”

“ And bearing in mind that these two females had already been with their respective mates for some considerable time, and that there was reason to believe that coition had actually taken place, is it likely that any counter-attraction would have proved successful in tempting either of them away from its mate, or probable, if they were the sole attraction, that the intruding male would have been so persistent in remaining? How very much simpler it is to fit the pieces together, if for the time being we ignore the female and fix our attention upon the territory. ”

“ Moreover, in some species, the male rejoins the flock at intervals during the early part of the season and for a time leads a double existence, passing backwards and forwards between its territory and the neutral ground. Its behaviour under these circumstances affords some valuable evidence, for the bird displays little if any hostility when accompanying the flock, yet when it returns to the ground over which it exercises dominion, no male can approach without being attacked. ”

“ If then we single out certain pairs and day by day observe their actions and their attitude towards intruders, we shall notice that, instead of their routine of existence consisting, as a casual acquaintance with the pool and its inmates might lead us to believe, of an endless series of meaningless disputes, the behaviour of each individual is directed towards a similar goal—the increasing of the security of its possession ”

“ And in support of this fact I have only to state that I have met with no single instance of failure to obtain and hold a mate when once a territory had been secured. Bearing in mind then that both sexes participate in the fighting, and that individuals of the opposite sex frequently attack one another; that all such conflicts are characterised by persistent effort, and that they are not limited to just the particular period when the sexual instinct is dominant but continue throughout the breeding season ”

“ There are, it is true, occasional displays of pugnacity which never seem to develop into anything very serious; for instance, one bird will fly at another, and a momentary scuffle is followed by a short pursuit but nothing more—nothing, that is to say, in the least comparable with the battle previously described. Of what is the flock composed? Of members of both sexes. There is no difficulty in assuring oneself that this is so. But is it entirely composed of individuals in whom development has not reached a stage adequate for the functioning of the primary dispositions? ”

“ 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. ”

“ It requires the presence of a female and, as we have seen, a female is by no means always present. Then it attributes to the one side of the inherited nature an influence which is not borne out by the facts, for in the ordinary routine of existence, without the incentive of battle, every individual is liable to wander occasionally beyond its boundary and to intrude temporarily upon its neighbours ”

“ 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. ”

“ Therefore, if it be true that the female raises the emotional tone of the male, the result will be an increased flow of energy into all the specific modes of behaviour connected with reproduction, amongst which those directly concerned in the securing and defence of the territory will receive their share ”

“ 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. ”

“ How then does the whole matter stand? If it were males only that engaged in serious conflict, and if they fought only in the presence of a female, the problem would resolve itself into one simply of obtaining mates. ”

“ In withdrawing from its companions in the spring, the male is breaking with the past, and this action marks a definite change in its routine of existence. But the change does not end in attempted isolation; it is carried farther and extends to the innermost life and affects what, humanly speaking, we should term its emotional nature, so that the bird becomes openly hostile towards other males with whom previously it had lived on amicable terms. ”

“ Let us take some common species, the Willow-Warbler being our first example; and, having found three adjoining territories occupied by unpaired males, let us study the conflicts at each stage in the sexual life of the three individuals, observing them before females have arrived upon the scene, again when one or two of the three males have secured mates, and yet again when all three have paired. ”