Henry Eliot Howard

“ The Curlew, when surprised, utters a cry with which most of us. I suppose, are familiar; but when with lowered head it drives away another individual from the feeding ground, it gives expression to its feelings by a low, raucous sound, which again is different from its cry when a Common Gull steals the arenicola that has been drawn out of the mud with such labour. ”

“ And so, all species, or at least a large majority of them, have, in greater or lesser variety, cries and calls which are peculiar to certain seasons and certain situations; and since on many occasions we have indisputable evidence of the utility of the sound produced—as when, upon the alarm being given by one individual, the flock of Lapwing rises, or when, in response to a particular note of the parent, the nestling Blackcap ceases to call—so are we bound to infer that all the cries are, in one way or another, serviceable in furthering the life of the individual. ”

“ Now a male may leave its territory for three reasons—to pursue an intruder, to join the flock on neutral ground, or to find the necessary means of subsistence on other feeding grounds. On each of these occasions it hears the song of, and is in close contact with, other males; and if the relationship of which we are speaking be really exclusive of cross-correlation, its instinct ought to respond with the customary freedom. But what happens? A male pursues its rival, betraying much emotion and singing extravagantly, until the boundary is passed, when emotion subsides and it is silent ”

“ We have already seen that it belongs to the nature of the male during the season of reproduction to establish itself in a definite place, and this action is just as much a part of its hereditary nature as the building of the nest is of that of the female, and it is just as necessary for the successful attainment of reproduction. What exactly the stimulus is to this mode of behaviour we do not know; we can go no further back than the internal organic changes which are known to occur and which we assume, not without some reason, are responsible for its initiation. ”

“ 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 nonbreeding 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. ”

“ Clearly then we are confronted with a relationship of a different kind from that which we have been discussing; for not only is anything in the nature of stimulation absent, but, and this is a remarkable fact, the other items in the environment which formerly evoked response no longer do so in quite the same way. Is there any awareness on the part of the male of the relation between his voice and the mate that is to be, or is it merely that as the sexual situation increases in complexity some inhibiting influence comes into play? ”

“ Through awareness of something in the environment the male responds to stimulation, and the only reply we can give is that the headquarters, or a distant song, or the proximity of another male—with all of which, as we have seen, the instinct is definitely related—are the specific factors which normally evoke response—and experience teaches us that the periods of quiescence are just those when life is at its lowest ebb and these stimulating factors less in evidence. ”

“ We have already had occasion to refer to the fact that the males of some species desert their territories temporarily and join together on ground which is regarded by the birds that associate there as neutral, and that they do so not merely for the purpose of securing food but because they derive some special pleasure from the act of association, and we shall find that the altered behaviour of the male when it leaves its territory to seek food or to join the flock is an important point for us just now. ”

“ After the manner of the race the male makes temporary excursions from its territory accompanied by his mate, and it is noteworthy that when he returns and she is absent he sings, but that the moment she joins him, or even comes into sight, he is silent. In fact, in greater or less degree, a change is noticeable in the song of many resident and migratory species under similar circumstances, a deterioration so marked that we learn by experience to regard it as a certain indication of the arrival of a mate. ”

“ In fact the males of most species, when they are finally established on the breeding grounds, make themselves as conspicuous as possible by sight and by sound. And since the sounds produced by no two species are exactly alike, the females are able to recognise their prospective mates, and the males that are still in search of ground have ample warning if that upon which they are treading is already occupied. So that you see, from the remarkable development of the vocal powers in the male, there follow two important results—"recognition" and "warning." ”

“ But, in truth, the presence of a second male makes the situation, so far as the relationship between the song and the female is concerned, all the more perplexing; for, as we have already seen, the instinct of pugnacity, when aroused by the appearance of an intruder, is also liable to be accompanied by a similarly extravagant song. On each occasion the vocal effort is infused with much feelling tone, and it would be impossible to point to any one feature which is peculiar to only one occasion. ”

“ 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. ”

“ The male Grasshopper-Warbler, when it first reaches us, sings persistently, but when it is joined by a female a change becomes apparent; instead of the incessant trill, there are spasmodic outbursts of short duration, and in the course of a few days the bird lapses into a silence which may be broken for a short while at dawn, or late in the evening, but is often complete. ”

“ Now when different individuals collect in flocks at certain seasons, they assist one another in finding food, and afford mutual protection by giving timely warning of the approach of a common enemy, and the gregarious instinct is thus of great advantage to the species; but no matter how powerful the impulse to flock might be, if there were no adequate means of communication, the different units would frequently fail to discover their neighbours. ”

“ If, in the latter part of May, we take up a position at dawn in some osier bed, we listen to songs which have reached a high degree of specialisation, songs, moreover, which appeal to us on account of their beauty; if, on the other hand, we climb down the face of the sea cliff, we hear an entirely different class of songs—harsh, guttural, weird, monotonous sounds, which, appeal to us though they may, lack the music of the voices in the osier bed. ”

“ 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. ”

“ 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. ”

“ When the normal course of the song is not interrupted by the arrival of a female, when, that is to say, the male still pursues the routine to which he has all along been accustomed, and still sings at stated intervals in stated places with a voice that betrays no heightened emotional tone, even though the song may convey some meaning to the delicate perceptual powers of the female, we have nothing to lay hold upon which can be construed as an indication of direct relationship between the song and the presence of the female. ”

“ Thus it will be seen that, even after the internal organic changes have taken place, the instinct of song is not susceptible to stimulation at all times and under all circumstances, but only at certain specified times and under special circumstances which can be observed to correspond with the occupation of the territory.
In many species each male singles out within its territory some prominent position to which it resorts with growing frequency. This position is an important feature of the territory, and exercises a dominating influence on the life of the bird. ”

“ Further evidence is afforded in the behaviour of those species that make temporary excursions from their, territories. The male, on its return, flies as a rule direct to its special tree and sings. Sometimes, however, it settles upon the ground, not unfrequently accompanied by the female, and while there remains silent; but presently rising from the ground and deserting its mate, it flies to the headquarters and sings. Again, nearly every male at one time or another in the course of the season is aroused to action by the intrusion of a rival. ”

“ 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. ”

“ 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? ”

“ 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. ”

“ 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 ”

“ When the Wood-Warbler secures a territory it repeats its sibilant trill with unwearying zeal, yet no sooner does a mate appear than its emotion is manifested in other directions. The Reed-Bunting is vociferous during February and March; but when a female arrives, periods of silence are frequent and the instinct of the bird becomes progressively less susceptible to stimulation. ”

“ 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 ”

“ 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. ”

“ 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. ”

“ 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. ”