Numerous instances have since come under my notice. Hen Chaffinches become so absorbed that they fall to the ground and there continue the struggle. Seizing hold of one another by the feathers of the head, they roll from side to side, and then, without relaxing their grip, lie exhausted—the quickened heart-beat, altered respiration, tightly compressed feathers and partially expanded wings betraying the intensity of the conflict.
As the breeding season approaches, hen Blackbirds grow more pugnacious. Individuals that early in the year have frequented the same spot daily and have even shown every sign of friendship, become openly hostile. For two years in succession I had an opportunity of observing females under such conditions, and of studying the gradual change in their relationship. Each morning at break of day and for some hours afterwards they could be seen in the same place, one following the other as they searched for food first in this direction and then in that, as if they derived some special pleasure from the fact of their companionship. Then a change began to manifest itself. Indications of animosity became apparent; one would run towards the other in a threatening attitude and, in a half-hearted manner, peck at it; and gradually the hostility grew, until the tentative pecking developed into a scuffle and the scuffle into a conflict.
Much fighting also occurs between the females of the Reed-Bunting, and likewise between those of the Moor-Hen, and because these two species are not only common but inhabit respectively open stretches of marshy ground or large sheets of water, the fighting can be readily observed.
Why do the females fight before they are definitely paired? To obtain mates? This certainly seems to be the obvious explanation because any question of securing territory can be excluded; yet if it be true that their sex is numerically inferior, it is difficult to understand the necessity for such strenuous competition. 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 bear in mind, however, that the competition between the males is very severe, that large numbers probably fail to pass even this preliminary test, and that only a proportion are in a position to offer to the female the condition under which her process can successfully run its course; so that the presumption is—though it is incapable of demonstration—that there is a competition for such males each recurring season, and that, on the average, the weaker females fail to procreate their kind.
But apart from any direct assistance she may give in driving away intruders, does she in any way help to further the biological end of reproduction? This is a difficult question to answer, and the suggestion I have to make can only apply in those cases in which the territory is occupied throughout the breeding season. Much of the fighting between the males occurs in her presence, and it must be admitted—though it is difficult to speak with any degree of certainty—that such fighting, taken as a whole, bears the stamp of exceptional determination. Let us then grant that the excitement of a male does, under these circumstances, reach a higher level of intensity, and let us see how this will add to the security of the territory. The fact that the male has established itself and obtained a mate is not alone sufficient to accomplish the end for which the territory has been evolved. During the period between the initial discharge of the sexual function and the time when incubation draws to a close, much may happen to prejudice the future of the offspring; there is always the possibility of invasion by an individual whose development is backward or which has been unsuccessful in making good the first step, and, as we saw in the case of the Reed-Bunting, a portion of the ground won may be lost; there is always the danger of gradual encroachment by neighbouring owners; and there is even a possibility that a pair may be so persistently harassed by more virile neighbours as to forsake the locality permanently. If then a male is to attain a full measure of success it must be capable of keeping its boundaries intact up to the time when the young are able to fend for themselves, and consequently it is important that its intolerant nature should remain susceptible to stimulation throughout the greater part of the season.
Does the presence of a female serve to promote this end? Now we know very little of the influence exerted by one sex upon the other. Professor Lloyd Morgan has suggested that the male raises the emotional tone of the female, a suggestion which seems to me in accordance with the facts. There is reason to believe, however, that the converse is also true—namely that the excitement of the male reaches a higher level of intensity when a female is present. Granting then that his emotional tone is raised, how will this affect the question? So great is the difference of opinion as to the part that the emotions play in furthering the life of the individual that one hesitates to accept any particular one. But it seems to be generally admitted that emotion adds to the efficacy of behaviour, and this is the view of Professor Lloyd Morgan. "Whatever may be the exact psychological nature of the emotions, it may be regarded," he says, "as certain that they introduce into the conscious situation elements which contribute not a little to the energy of behaviour. They are important conditions to vigorous and sustained conation." 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; so that instead of a progressive weakening of just those elements in the situation which make for success, the level of their efficiency will be maintained as a result of such reinforcement. But the female becomes intolerant of her own sex when she has discovered a male ready to breed, and, later, assists her mate in resisting intrusion; and by raising her emotional tone, he may be the means of furthering more strenuous behaviour on her part. Each member of the pair would in this way contribute towards the energy of behaviour of its mate, and hence add indirectly to the security of the territory.
It may be well to illustrate the foregoing remarks. Suppose that there is a small piece of woodland barely sufficient to hold three pairs of Willow-Warblers, and suppose that the male and female in the middle territory did not respond to one another's influence quite as readily as the adjoining males and females, what would be the result? The emotional tone of the central pair would stand at a lower level of intensity; and, since their congenital dispositions would lack the necessary reinforcement, the birds would tend to become less and less punctilious in keeping their boundaries intact, whereas the adjoining pairs, always on the alert and meeting with little opposition, would encroach more and more and gradually extend their dominion. 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? 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. And surely success will be attained by that pair whose emotional tone stands high and whose impulse to fight is therefore strong, rather than to the ill-assorted couple.
The argument, then, is briefly this. In the spring, a marked change takes place in the character of the males of very many species; instead of being gregarious they either avoid one another and become hostile, or, if their conditions of existence require that they shall still live together, they become irritable and pugnacious. This change is made known to us by the battles of varying degrees of severity which are such a feature of bird life in the spring; and since a female can commonly be observed to accompany the combatants, the possession of a mate appears at first sight to be the proximate end for which the males are contending. But when the circumstances which lead up to the quarrels are investigated closely, the problem becomes more difficult; for it is not merely a question of males fighting in the presence of a female, as is generally supposed to be the case, but on the contrary there is a complexity of strife which is bewildering—males attack females or vice versa; female fights with female; or a pair combine to drive away another pair, or even a solitary individual no matter of which sex. This complexity of strife makes against the view that the possession of a mate is the reason of the fighting. But an even stronger objection is to be found in the fact that males are hostile when no female is present—and hence we must seek elsewhere for the true explanation.
Now if the behaviour of a male be closely observed, it will be found that its pugnacious instinct gains or loses susceptibility according to the position which it happens to occupy—when its ground is trespassed upon, the impulse to fight is strong; but when it crosses the boundary it seems to lose all interest in the intruder. 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. The conclusion, therefore, seems to be inevitable, namely that the actual occupation of a territory is the condition under which the pugnacious nature of the male is rendered susceptible to appropriate stimulation.