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. ”

“ I seek only to ascertain whether the impulses that are concerned in the securing of a territory, and in the search for society, bear any relation to the problem as a whole. I hold that the origin of migration is not to be found merely in conditions peculiar to a remote past, but that the conditions inhere in the organic complex of the bird, and are thus handed down from generation to generation. ”

“ 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. ”

“ The cliff-breeding species—Guillemots, Razorbills, and Puffins—are difficult to investigate because individuals vary so little, and the sexes resemble one another so closely; yet, despite these difficulties, we can gain some idea of the general purport of their activities. But when the ledges are crowded and the air is filled with countless multitudes, how is it possible to keep a single bird in view for a sufficient length of time to understand its routine? ”

“ The former, by itself, is insufficient; in the dim light of the early dawn, when life is at its highest, and mating proceeds apace, what aid would it be for a male to perch on the topmost branch of a tree, how slender a guide in the depth of the forest? But whether in the twilight or in the dark, in the thicket or the jungle, on the mountain or on the moor, the voice can always be heard—and the voice is the principal medium143 through which the sexes are brought into contact. ”

“ Let us take the individual and see whether we can establish any relation between the hostility it displays towards members of other species and its general disposition to secure a territory. We must remember that a male can have no knowledge of the prospective value of its behaviour, nor is it likely that it has any ulterior purpose in ejecting other males, beyond the pleasure it derives from satisfying its impulse to do so. The proximate end of its behaviour is to attack, nothing more, and this, of course, it can only do just in so far as the intruder evokes the appropriate instinct. ”

“ If there had been no flock, if a few solitary individuals had appeared here and there and had established themselves in different parts of the meadow, one would have had no definite evidence of the strength of the impulse in the male to seek a position of its own, one could only have argued from the general fact of males flocking in the winter and isolating themselves in spring that something more than accident was required to explain so radical a change. ”

“ Warblers alter their mode of life, and in the osier bed, or amongst the elders, seek their food together in peace; Finches, Buntings, Pipits, and Wagtails, though food is everywhere abundant, gather themselves together respectively into bands which, as winter approaches, grow into flocks and even into composite flocks; and as the Warblers leave for the south, so their places are filled by flocks of Thrushes and Finches from the north. In whatever direction we turn, when the days begin to shorten, it is the community, not the individual, that thrusts itself upon our attention ”

“ 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 ”

“ 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. ”

“ An individual appears here, another there; then after a pause there is a further addition, and so on with increasing volume until the tide reaches its maximum, then activity wanes, and the slowly decreasing number of fresh arrivals passes unnoticed in the wealth of new life that everywhere forces itself upon our attention. If now, instead of surveying the migrants as a whole, our attention be directed to one species only, this gradual arrival of single individuals in their accustomed haunts will become even more apparent ”

“ 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. ”

“ If there were always sufficient breeding ground to support the offspring of all the individuals of each species, if the individuals were always so distributed that there was no possibility of overcrowding in any particular area, and if the conditions of existence of different species were so widely divergent that the presence of this one in no way affected the interests of that, no opportunity would be afforded for the development of so complex a system as is involved in the "territory" and all that appertains to it. But the available breeding ground is by no means unlimited. ”

“ 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. ”

“ What then is the stimulus? 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. ”

“ For, after all, what does each individual seek? There may be some immature birds which, though they have not reached the necessary stage of development, happen to fall in with others in whom the impulse is strong and are led by them—they know not where. But the majority seek neither continent nor country, neither district nor locality is their aim, but a place wherein the rearing of offspring can be safely accomplished; and the search for this place is the earliest visible manifestation in many species of the reawakening of the sexual instinct. ”

“ As far as my experience goes, the movement is less evident amongst the members of a flock than when an individual is resting alone, or even with a few companions, which may be due to the fact that since some members are always awake and watchful, a bird of prey would have more difficulty in approaching a flock unawares than it would have in approaching a single individual. With the greatest ease a Sparrow-Hawk can pick up a Thrush as it feeds on the meadow by itself, but if it attempts to seize one of a flock, the chances are that its approach is signalled and that its prey escapes. ”

“ 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. ”

“ I watch them from February to June, and observe the order in which they are mated—first a mature male; next, after a short interval, the immature male; and finally, after a still longer interval, the remaining bird gets a mate. As the season advances, still keeping watch on the development of the plumage and of the voice of the immature male, I observe that no very definite change takes place—that the colours remain dull, that there is a conspicuous absence in the song of certain phrases, and that the notes lack purity of tone. ”

“ This behaviour is expressed in a peculiar flight. The bird rises high in the air, circles round with slowly beating wings above the marsh, and utters a call which, as far as my experience goes, is characteristic of the performance. The air is often full of individuals circling thus even beyond the confines of the marsh, for a male does not limit its flight to a space immediately above its territory; but nevertheless careful observation will show how unerringly each one returns to its own position on the breeding ground, no matter how extensive the aerial excursion may have been. ”

“ The Blackcap and the Garden-Warbler are constant rivals, and the scenes which can be witnessed when the two meet in competition are interesting from many points of view. The birds not only pursue and fight with one another, but their emotional behaviour reaches a high level of intensity—excitable outbursts of song are indulged in, tails are outspread, wings are slowly flapped, and feathers raised—in fact the attitudes assumed are similar in all respects to those which occur during the contests which are so frequent between the respective individuals of each species ”

“ 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. ”

“ Hence it is clear that the nature of a bird when on migration is not quite the same as it is when its destination is reached; that the positions occupied from time to time during the journey carry no meaning, or, rather, are not brought into relation with its life in quite the same way as is the position which it finally occupies; and further, it is clear that the interest it displays in other species undergoes a somewhat remarkable transformation when at length its destination is reached. ”

“ But the system of reproduction does not consist merely of a search for the breeding ground, and of the discharge of the sexual function; it is a much more complex business, yet withal more complete. Nothing is left to chance; the end is attained step by step; and each successive stage marks the appearance of some specific factor which contributes towards the success of the whole. We start with the appropriate organic condition under which, when adequate stimulation is provided, the disposition to secure a territory comes into functional activity. ”

“ We saw that many different species require similar conditions of existence, that ancient breeding haunts disappear and that new ones come into being, and that in the ordinary course of events such species must often assemble in the same area for the purpose of reproduction. So that even though a male might be successful in protecting its ground from intruders of its own kind, yet it might still fail to rear offspring, just because it happened to choose a position in which other kindred forms had gathered. ”

“ Migrants which formerly found in it no suitable accommodation now begin to appear, and as the seasons pass by and the undergrowth affords more and more shelter for the nests and an increasing supply of insect life, so their numbers increase until the wood becomes an important breeding station, resonant with the song of many individuals. But slowly the growth increases; the bushes pass into saplings and the saplings into trees, and the undergrowth then disappears just as surely as do the migrants which can no longer find there the conditions which they require. ”

“ Many a Warbler allocates to itself a space of ground more than sufficient to supply it with all that it needs; so, too, does the Finch, or the Pipit, or the Falcon—if we take no account of kindred species. Thus there is reason to believe that, by limiting the number of individuals in a given locality, this apparently wasteful expanse of territory is serviceable in that it provides against the pressure of the bird population upon the available means of support becoming too great. ”

“ But the significance for us just now lies in the fact that the definiteness, which accompanies the initial behaviour of the migratory male in relation to the territory, cannot have been acquired by repetition; for this reason, that when the male occupies its space of ground at the end of its long and arduous journey, it does so without preparation or experiment, even without hesitation, as if aware that it was making good the first step in the process of reproduction. ”

“ One marvels that it ever does survive; one marvels at the evolution of a constitution sufficiently elastic to withstand such changes. But, however much the constitution may give us cause to wonder, it is clear that much depends upon the parents. A slight inefficiency of the instinctive response which the presence of the young evokes, a little slowness in searching for food or sluggishness in returning to the nest, might lead to exposure and prove fatal. And, however much is demanded of the parents, it is clear that much also depends upon the relationships in the external environment ”

“ Yet a Whinchat, when it has established itself, is most pugnacious; it not only attacks every bird of a similar size that approaches its position, but its behaviour under such circumstances bears the impress of unusual determination; and if we were to take a male and place it in the position of the Stonechat, we should find that its nature would change, that the presence of the Stonechat would evoke a hostile response, and, conversely, that the instinct of the Stonechat would not be susceptible to stimulation. ”

“ 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. ”

“ But when we consider the lapse of time, and the changes that must have occurred in the character of the bird population—the appearance of new forms and the disappearance of the old, the ebb and flow of a given species in a given area—and bear in mind that, notwithstanding this, the migratory instinct, if not stronger, is assuredly no less strong, and the volume of migration, if not greater, is assuredly no less ”