Territory in Bird Life

by Henry Eliot Howard

What, then, determines its value? Are there any qualities which, whether the bird is mature or immature, whether it is untrained or has acquired fuller expression by practice, whether it inhabits this district or that, are alike constant? Well, no matter how great the variation, no matter how much this voice falls below or exceeds the standard, judged from the human standpoint, attained by that, even we, with our duller perception, have no difficulty in recognising the species to which the owner of the voice belongs; in other words, the song is always specific, and this is the most noticeable, as it is the most remarkable, characteristic.

There is still, however, another quality to which I would draw attention—that of loudness. The sounds produced are on the whole alike penetrative, and the individuals of any given district, even though the climate by affecting their vocal muscles may have modified the character of the song, are at no disadvantage in this respect; neither are the females on the same account the less likely to hear the undeveloped voice of the immature male.

We have then the following considerations: firstly, there is the widespread and remarkable phenomenon of imitation, from which we can infer that the diversity of song is not due to structural differences but must be sought in some innate capacity to play one tune in preference to another; secondly, not all the individuals of the same species play a similar tune—we find that there is in certain directions a noticeable variation which nevertheless does not seem to affect the question of success or failure in the attainment of reproduction; in the third place, in contrast with this variation, we can observe a striking uniformity in two important particulars, namely in the specific character and penetrative power of the song—qualities which we know are essential for the purposes of "recognition" and "warning"; and finally, from the general course of our investigation, we can infer that if a male had no certain means of advertising its position, the territory would not be brought into useful relation in its life. Have we here sufficient ground on which to construct a theory of origin; in other words, has the evolution of song been incidental to, and contributory to, the evolution of the territory?

We have all along spoken of the song and of the call-notes as if they were manifestations of separate emotional states having their respective and well-defined spheres of usefulness; and while, speaking generally, this is a true statement of the case, there is much evidence to show that the relationship between them is nevertheless very close. There are, for example, quite a number of cases in which a particular call-note is uttered with unusual energy during sexual emotion, and is attached to the song, of which it may be said to form a part; but a still closer connection can be traced in many simple melodies which are merely compositions of social and family calls repeated many times in succession, and even in some of the more complex productions there will be found indications of a similar construction. And since this is so, since moreover, in the seasonal vocal development of such a bird as the Yellow Bunting, we can observe the gradual elaboration from simple to complex—from the repetition of single notes to phrases and from phrases to the complete melody—we have every reason to suppose that it is along these lines that the evolution of the voice has proceeded.

In all probability there was a time when vocal expression was limited to primitive social and family cries which would be called into play with special force during times of excitement, more particularly during the sexual season which is the period of maximum emotional excitement. But the excitement would express itself in all the congenital modes of behaviour peculiar to the season, and thus the repetition of these cries would become associated with combat, with extravagant feats of flight, and with other forms of motor response. Now the more emotional individuals would be the more pugnacious, and all the more likely therefore to secure territory and so to procreate their kind; and, being of an excitable disposition, they would at the same time be the more vociferous. Hence variations of the hereditary tendency to vocal expression, even though in themselves they were not of survival value, would be fostered and preserved, so long as they were not harmful, in virtue of their association with pugnacity. But if, instead of being neutral, they helped to further the biological end of combat, the relationship between the voice and pugnacity would be of a mutually beneficial kind; and those individuals in which variation in both directions happened to coincide, would have a better chance of success in the attainment of reproduction.

A territorial system, closely corresponding to that which we have discussed, forms part of the life behaviour of certain mammals, and of its existence much lower in the scale of life evidence is not wanting; from which we can infer that it is not of recent origin, but that the conditions in the external environment demanded such a system at a remote period of avian development. Now even in its incipient stages the system must have involved a separation of the sexes, and howsoever slight the degree of separation may have been in comparison with that which can be observed to-day, inasmuch as the power of locomotion was then less highly developed, mating could only have proceeded satisfactorily providing that males fit to breed had some adequate means of disclosing their positions. Thus there is reason to think that from the very commencement of the process variations of emotional disposition expressed through the voice would have been of survival value.

But expressed in what direction, in loudness and persistency of utterance, these are the qualities which, I imagine, would have been more likely to have facilitated the search of the female? Yet if she were uncertain as to the owner of the voice, neither loudness nor persistent repetition would avail much; and as species multiplied and the competition for the means of living became increasingly severe, so the necessity of a territory would have become intensified, and so, too, with the extension of range, would the separation of the sexes have been an ever-widening one; and as with their multiplication, irregularities and delays in mating, arising from the similarity of the calls, would have increased in frequency, so a distinctive call, which would have tended to minimise these risks, would have come to possess biological value.

Here we have a theory of origin, but origin of what? Of certain characteristics of song—nothing more; and therefore to suppose that it furnishes a complete explanation, which satisfies all the requirements of scientific logic, of so wonderful an intonation as that, for example, of the Marsh-Warbler, or that no other relationships, except that of the territory, enter into the total emotional complex, simplifying here or elaborating there to meet the exigencies of diverse circumstances—to suppose this would be foolish. That there are many relationships which even to-day are leading to modifications in important particulars, but which at the present time are beyond our cognisance, of this there can be no doubt.

There is one process by which song may have attained a fuller development, and which would account in some measure for the elaboration, inexplicable merely in terms of "recognition." It is this: the effect of the sexual call upon the female cannot well be neutral, it must be either pleasurable or the reverse—it must, that is to say, be accompanied by some suggestiveness, and by suggestion I mean the arousing of some emotion akin to that of the male; and if there are degrees of suggestiveness, which well there may be, some males will mate sooner than others and some will remain mateless—this is the theory of sexual selection. The question to be decided here is whether the biological emphasis is on loudness, or specific distinctness, or pitch, or modulation, or the manner in which the phrases are combined—that is, on some qualities in preference to others—or whether the emphasis is on the whole. We have already seen, and it is well known, that there is much variation in the voices of different individuals of the same species, and thus the first condition of the theory is fulfilled. Now the conditions which lead to variation are threefold—immaturity, seasonal sexual development, and isolation. Of the three, the variation in the case of the immature bird is the most instructive; the tone is not so pure, the combination of phrases is incomplete, and elaboration is imperfect, and yet, notwithstanding all these imperfections, we can observe that the bird pairs as readily as does the adult. But even if we lacked this demonstrative evidence, we should still be justified in assuming that such must be the case, for we know from experience in the preservation of game, where there is no surer way of reducing the stock than by leaving too high a percentage of old cocks, that for the young bird to be at any disadvantage in competition with the adult is detrimental, if not disastrous, to the species. So that while there is plenty of evidence of variation in those particular qualities which appeal to our æsthetic faculties, there is at the same time evidence which demonstrates that such variations exercise no influence on the course of mating; and inasmuch as it is difficult to conceive of any voice departing more from the normal type in these particular qualities than the immature does from the adult, if there be degrees of suggestive influence, we must seek it in some other direction. There remain the two other characteristics which we found to be constant under all circumstances, namely, loudness and specific distinctness; and if, in addition to serving the purpose of disclosing the positions of the males, they serve to evoke some emotion in the female, which helps to further the biological end of mating, so much the more reason is there for their survival.

There can be no question that this ingenious and attractive theory, if it were true in its special application to song, would immensely simplify interpretation, and moreover that preferential mating would contribute not a little to the success of the whole territorial system. No one can deny the strength of the argument: that the sexual instinct, like all other instincts, must require a stimulus of an appropriate kind; that the effect of the sexual call upon the female cannot be neutral; and hence the probability that stimulation varies too; no one, I say, can question the strength of this evidence, and, one might add, of the evidence derived from the analogy of the human voice. But when we have said this, we have said all; and our acceptance of the hypothesis, so far as song is concerned, must remain provisional so long as the evidence remains but secondary evidence.