Territory in Bird Life Part 8

Though it be true that every instinct requires for its functioning a stimulus of an appropriate kind, yet it is also true that the condition which will render it responsive must be present. What we have then to consider is whether the phenomena which we have explored give us any clue as to the particular nature of that condition. In the first place, we have the general fact that the hostility is not confined to a few species belonging to a few families, but that it is of wide application--birds of prey, Warblers, Woodp.e.c.k.e.rs, all supply us with evidence which serves to show, in greater or less degree, its nature and extent. Next, we found that the hostility was peculiar to a certain season--and that one the season of reproduction. And if the question were asked: What condition would then be most likely to render the instinct susceptible, the answer that would most certainly be given would be--the presence of a female. And in reply to a further question as to the particular nature of the stimulus to which the instinct would respond, we should be told--the presence of another male of the same species. Now the possible influence of the female on the course of the male's behaviour was the subject of inquiry in the second chapter, wherein we endeavoured to explain the hostility between males of the same species, and we came to the conclusion that it was not alone sufficient to account for the facts disclosed. Still less likely, therefore, is it that her presence can bear any direct relation to the hostility between different species, the more so since the biological end of securing a mate is definitely excluded. And we have something in the nature of proof of the correctness of this view in the fact that she accompanies her mate when he joins the flock, and that there his instinct is not susceptible to stimulation. We then proceeded to examine certain cases in which all the indications pointed to the fact that the "will to fight" was present in only one of the opponents; and we attached considerable importance to this circ.u.mstance, because we knew from experience that the same bird which seemed to lack courage, could at other times and in other situations be most aggressive. If then we ask what condition was present on the one occasion that was absent on the other, we have no difficulty in finding a reply--on every occasion on which the opponents appeared to be unevenly matched, one was in occupation of a territory and the other was not. And if we inquire further as to which of the two was the aggressor, the answer is again clear, namely, the bird that occupied a territory. Finally we considered some particular instances in which the "will to fight" was present alike in both opponents, and in which the battles were protracted and severe.

But the fact that a bird has established a territory is not in itself sufficient to render its hostile nature susceptible; it must be actually in occupation if a response is to be elicited. We reach this conclusion step by step: the behaviour of the migrant, that lacks the "will to fight" when on migration but is pugnacious when it has secured a territory, shows it; the behaviour of the resident, which temporarily joins the flock and is there sociable, shows it; and it is shown also by the determination with which both opponents fight when the question of ownership of a station is in dispute. And of all the facts we have reviewed, this is perhaps the most important in relation to our present subject, for it demonstrates that the change from sociability to hostility is not merely an incident of the s.e.xual season, not merely an indirect result of the functioning of the general disposition which leads to the establishment of a territory, but that it is intimately a.s.sociated with the whole process, and that the particular part of the bird's nature which is concerned is so nicely balanced that it will respond under one condition and one only.

Thus we are led to the only conclusion which seems consistent with the facts, namely that there is a relationship between the "territory" and the hostility.

If we are satisfied that all this warfare is not merely an expression of an instinct which is serviceable in another direction, what part does it play in the whole scheme of reproduction?

The young of many birds are delicate at birth and unable to withstand exposure to cold, and in the previous chapter we came to the conclusion that the territory was serviceable in that it provided an adequate supply of food in the vicinity of the nest, and thus obviated the necessity of the parents being absent from them for long. But manifestly no matter how active a male may be in driving away members of its own s.e.x and kind, it will neither make its position secure, nor insure a supply of food for its young, so long as any number of individuals of different kinds are allowed to establish themselves in the same s.p.a.ce of ground. On the one hand, then, we have the fact that there is constant strife between males of close affinity, whilst on the other, we know that many species require like conditions of existence and are bound to a.s.semble wherever these conditions are suitable; and we can infer that the territory would fail to serve its purpose if no restriction were imposed upon the measure of such a.s.semblies.

The question then arises: Does all this warfare contribute towards the attainment of reproduction? Not far from my house there is a small water meadow, three acres in extent, which for some years has been derelict and is now overgrown with the common rush (_Juncus communis_) and small alder trees. For three successive seasons I watched the bird life of this meadow, and more especially the Reed-Buntings whose behaviour I was studying at the time. In every respect the meadow was suitable for this bird; there was an abundance of food and numberless situations in which nests could be placed. Each year all the pairs were successful in rearing one, if not two broods, yet the number of pairs never exceeded five--the first year there were three; the second year five; and the third year four. In addition to the four pairs of Reed-Buntings, there were in the spring of 1915, six pairs of Whitethroats, one pair of Lesser Whitethroats, four pairs of Willow-Warblers, one pair of Sedge-Warblers, two pairs of Gra.s.shopper-Warblers, one pair of Chiffchaffs, three pairs of Hedge-Sparrows, two pairs of Tree-Pipits, one pair of Skylarks, one pair of Whinchats, one pair of Flycatchers, two pairs of Song-Thrushes, one pair of Blackbirds, one pair of Redstarts, three pairs of Chaffinches, and one pair of Wrens--in all, thirty-five pairs, whose young were mainly dependent for their living upon the insect life of that meadow and the ground immediately surrounding it. If we allow three young to each pair--and this would take no account of second broods--we arrive at the following result, namely, that one hundred and five young and seventy adults had to be supplied with food from that locality, which would mean, if the search for food were strictly limited to that meadow, that 83 square yards would be allotted to each individual.

Suppose now that the four male Reed-Buntings had each admitted one other male, and that they had secured mates, what would have been the effect upon the whole community? The four additional pairs with their young would have represented twenty individuals, which would have represented a decrease of 8.5 square yards in the s.p.a.ce allotted to each individual.

The pressure of the bird population upon the means of support would then have been materially increased; and not only the Buntings, but the Warblers, Pipits, and all the rest would have suffered. But the result would have been the same if, instead of the four additional male Reed-Buntings, four males of other kinds had been allowed to enter the marsh, and we can multiply the number four until we arrive at a point when the means of subsistence would no longer have been adequate for the adults, still less for the young. If, then, there were nothing to prevent this happening, many of the birds in that marsh would have no chance of rearing their young successfully. Hence, if the territory is adequately to serve the purpose for which we believe it has been evolved, some provision must have been included in the system to meet the difficulty.

There are three ways by which this may have been accomplished--indirectly, by increasing the size of the area occupied by each individual, and thereby reducing the relative number of each species; or directly, by rendering the fighting instinct of the bird susceptible to stimulation by individuals of other species; or, possibly, by a combination of the two. There were four pairs of Reed-Buntings in the marsh, and their territories covered the whole of it. But inasmuch as other insectivorous birds were established there also, and found sufficient food to maintain both themselves and their families, it is clear that the area these Reed-Buntings occupied was in excess of that which they would have required if they had been the sole inhabitants. And such often appears to be the case. Many a Warbler allocates to itself a s.p.a.ce 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 though a reduction in the numerical standing of the different species would certainly follow from any increase in the area occupied by the respective individuals, and with even greater certainty would place them in a more secure position as regards their supply of food, yet, when we remember how large a number are dependent upon a supply of insect life for their young, we can understand that it would not alone be a sufficient safeguard against the dangers attendant upon overcrowding. It is here, I believe, that we shall find the true explanation of the hostility; it roughly insures that the number of pairs in any given area does not exceed the available means of support, and indeed it is difficult to imagine how such uniformity of distribution as would free the young from the risk of exposure could be obtained without some such control.

Some birds, however, have no difficulty in finding the necessary food for their young, yet have great difficulty in finding a station where they can rear their young in safety; and the area each one occupies has been reduced to the smallest proportions in order that the maximum number can be accommodated. Here, any increase in the size of the territory would inevitably lead to the extinction of the race, so that nothing stands between failure and success except the ability of the bird to defend its territory. If we study the bird population at one of the breeding stations on the coast, we find, generally speaking, that each kind of bird inhabits a particular portion of the cliff; on the lower ledges are the Guillemots and Kittiwake Gulls; higher up are Razorbills and Fulmars, and at the top, where the cliff is broken and the face of the rock covered with turf and soil, the Puffin finds shelter for its egg. At the same time there is much overlapping; the kind of ledge that suits a Razorbill is equally suitable for a Guillemot or a Fulmar, and so, no matter how successful the Razorbill may be in establishing a territory and preventing intrusion upon it by other Razorbills, it will be all to no purpose if it allows itself to be jostled out of its position by a Fulmar. Hence, inasmuch as breeding stations are limited and compet.i.tion for territory so severe, only those forms in which the fighting instinct responds freely to a wide range of stimuli will be in a position to maintain a footing upon the cliff.

In trying to estimate the importance of the hostility in its relation to the territory, we must bear in mind that compet.i.tion varies in different seasons and in different localities. The surface of the land is constantly undergoing modification, partly owing to human and partly to physical agency--forests are cleared; marshes are drained; the face of the sea-cliffs is altered by the erosion of the waves; here the coast may be locally elevated, there locally depressed; and so forth. Many of these changes are slow and imperceptible, many can be observed in our own lifetime. The timber is felled and the undergrowth cleared in some wood, and the following spring we notice a change in the character of the bird population. Migrants which formerly found in it no suitable accommodation now begin to appear, and as the seasons pa.s.s 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 pa.s.s 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.

Or, as an ill.u.s.tration of the effect produced by natural agency, let me describe a change which has taken place in a corner of Co. Donegal. The promontory of Horn Head is bounded on the west by extensive sand-hills, 100 ft. or more in height. On the southern side it is divided from the mainland by a channel, which narrows down to 100 yards or so in width where it fringes the sand-hills, and then widens out again, covering an area of approximately 270 acres. As far as is known in the memory of man, this area has always been tidal. But in recent years a change has taken place, and the blown sand has silted up the channel, with the result that this tidal area has been transformed into a brackish lake.

What has brought about the change is not easy to determine. There is evidence, however, of a slow alteration of the level of the sh.o.r.e-line; for in the midst of the sand-hills, situated 150 yards or so from the present sea-margin, and running parallel with it, there is an acc.u.mulation of pebbles some 3 feet high by 4 feet deep. This raised beach is now separated from the Atlantic by sand-drifts of considerable height, and consequently there are some grounds for believing that secular elevation is taking place, which, if it be the case, will account for the change in progress. Now the effect on the bird population can be seen even now, and will doubtless become more apparent as the years pa.s.s by. Sand-Martins used to find plenty of places to breed amongst the sand-drifts, and moreover do so still. But their nesting sites are constantly changing and disappearing, and the breeding-place of one colony, that was situated in the bank of a stream twelve years ago, is now buried 10 feet or more below the surface of the sand. The area that was once tidal, but is now a brackish lake, is fed by mountain streams, and as the fresh water predominates, so in course of time will it become fringed with vegetation; and instead of the flocks of Curlew, Dunlin, and other waders that, at low water, resorted there to feed, Coots will fight with one another for the possession of territories, and the Wild Duck will teach her young to seek their food.

In whatever direction we turn, we find that many breeding grounds are subject to incessant change. Ancient haunts disappear, new ones come into being, a change which makes life impossible for this bird, as likely as not benefits that one, and so on. There is no stability. Hence in any given district each recurring season there must needs be a large number of individuals which are obliged to seek new stations, and if there were no control over their distribution, if each one were free to establish itself wherever it chanced to alight, this locality might be overcrowded and that one deserted; and, bearing in mind how many species there are that require similar conditions of existence, we can infer that the successful attainment of reproduction would become impossible for many of those individuals so long as each species was indifferent to the presence of the others. On the other hand, if there were no control over the range of the intolerance, the smaller bird would have no chance in compet.i.tion with the larger, and it is doubtful whether the larger would gain an advantage commensurate with the energy it would expend in ridding its area of the smaller. I have described battles in which the opponents were only distantly related; for instance, the Moor-Hen will attack almost any bird--Partridge, Lapwing, or Starling--that approaches its territory even temporarily. Nevertheless the antagonism between kindred forms is more prevalent, and, as a rule, characterised by more persistent effort; and thus it seems as if the susceptibility of the fighting instinct has its limitations, the degree of the responsiveness being dependent upon the affinity of the opponents.

Suppose now that we take an area inhabited by a number of different species requiring like conditions of existence, divide it into three sections, and imagine that in one they were all sociable, that in another they were all hostile, and that in a third those which were closely related were intolerant of one another. Let us suppose further that each one of them was represented by the full number of individuals that the law of territory would allow. In the first section an individual would establish itself, and, becoming intolerant of its own kind, would exercise dominion over an area roughly sufficient, providing conditions were normal, to insure an adequate supply of food for its young. But it would take no account of other species, and since any number might occupy the same ground, the fact of its having established a territory would not alone suffice to render its supply of food secure. Success in the attainment of reproduction would then become largely a matter of chance, depending upon the number of individuals that happened to settle in this place or in that. In the second section there would be perpetual warfare; for whereas the appropriate organic condition which leads to pairing arises in different species at different times, fresh claimants to occupied ground would constantly be appearing, and the efforts of the inhabitants to preserve their boundaries intact would have to be maintained throughout the whole period of reproduction; and while the stronger or more persistent forms would be more likely to breed, they would do so at the expense of their young, to which they would be unable to devote proper attention, and with an expenditure of energy that would reflect itself upon the future of the race. But the conditions of life in the third section would be such as would be more likely to yield good results. The relations of the different members of the community would be more evenly balanced, for a male would only be called upon to compete with those of its own size and strength. Thus, on the one hand, accommodation would be so divided as to secure the breeding of the maximum number of individuals with the minimum of expenditure of energy, whilst on the other, any undue pressure upon the available means of subsistence would be prevented.

There can be no question that in the latter section a higher percentage of individuals would succeed in rearing offspring. And so, by reason of the fighting instinct being more susceptible or less susceptible according to the affinity of the opponents, a control is established which, while preventing unnecessary extension of warfare, allows for sufficient extension to render the biological end secure.

These, then, are the facts--this the conclusion which can be drawn from them. It may, however, be said of these facts, as it has been said, with even less justification, of the battles between individuals of the same species, that they do not afford evidence of genuine hostility. No doubt there are many naturalists who could supplement these facts with others in which the conflicts resulted in bodily injury, or terminated fatally, or at least were of a more determined kind. But I have already drawn attention to the fact that, so long as a definite result is attained, the severity of the struggle and the amount of injury inflicted are matters of small moment. Let us, however, run over the substance of the argument, and then briefly refer again to this point of view.

After enumerating instances of hostility, sufficient in number, so it seemed, to const.i.tute reasonable ground for the belief that they had a part to play in the life-history of the individual, the two questions we set ourselves to examine in this chapter were: Is there any circ.u.mstance in the life behaviour of the individual with which the hostility can be definitely related; and, will the hostility lead to the securing of a greater measure of success in the attainment of reproduction?

Many different species a.s.semble together in winter and roam from place to place in search of food. But in spring their behaviour undergoes a remarkable transformation; they avoid one another and become quarrelsome, so much so that whereas the outstanding feature of the winter is sociability, that of the spring is hostility. With this general fact before us, we proceeded to investigate this change of behaviour still further. First of all we took the case of a migrant, and, comparing its behaviour, as it journeyed, with that when finally it reached its destination, we found that the bird which was notoriously pugnacious when in occupation of a territory betrayed no interest in other species as it travelled to the accustomed breeding ground. Not only so, but even though it was attacked, we found that its pugnacious instinct still failed to respond. Here, however, it may be contended, and with reasonable justification, that in the interval which elapses before the ultimate destination is reached, some change in the organic condition of the bird may occur which will account for its altered behaviour; or, it may be urged, with no less justification, that whereas on migration the bird is unpaired, when the destination is reached it is probably in possession of a mate and is therefore quarrelsome. Now, at the most, the interval can only be a matter of a few days, and it is unlikely that organic changes sufficient to bring about so important an alteration of behaviour could occur in so short a time, still less likely that they could be timed to come into functional activity just at the moment when the bird reaches its breeding ground. And with regard to the suggestion that the change can be accounted for by the presence of a mate, we shall do well to remember not only that males as a rule precede the females by some days, but that a male may even remain in its territory, mateless, for some weeks, and yet display hostility.

Nevertheless the case of the migrant did not, by itself, afford sufficient evidence upon which to base any conclusion. We therefore inquired into the behaviour of some of the residents at a corresponding period. The Bunting served as an ill.u.s.tration. Early in the season it establishes a territory, and because food is then scarce it is forced to seek it elsewhere than on the small plot of ground which it has acquired; and so it makes its way to some spot where the supply is abundant, and there, meeting with other species bent on a similar errand, forms with them a flock. Part of its time is then spent in the territory and part on the feeding ground, and between these two points a highway is formed by the bird pa.s.sing constantly to and fro. But the attention which it pays to other species is very different on these two occasions--when in the territory it is intolerant of strangers, but when it accompanies the flock it displays no interest in their movements.

From hour to hour its nature seems to change. But, as we saw, the inborn const.i.tution of the bird cannot change, and therefore we came to the conclusion that an explanation of the altered behaviour was to be found in the fact that the pugnacious instinct is only rendered susceptible under a certain condition. So that all the evidence tended to confirm the impression which we had gained from the course of events in the life of the migrant, namely, that the hostility bears a direct relation to the occupation of a territory.

Finally we were led to inquire whether the hostility was serviceable in promoting the welfare of the individuals. 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 a.s.semble 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. Hence if the territory is adequately to serve its purpose, some control over the local distribution of species is of paramount importance. Nevertheless, if all the different forms that require similar conditions of existence were intolerant of one another in a like degree, the smaller bird would have no chance in compet.i.tion with the larger. This, however, is not the case. Some, as we saw, arouse little or no animosity in others, in fact the more closely related the rivals, the more responsive their pugnacious nature seems to become.

To return now to the view that the fighting is not really serious, but, on the contrary, that it is either vestigial and has no longer any part to play in furthering the life of the individual, or that it is a by-product of the seasonal s.e.xual condition to which no meaning can be attached. First, there is the relationship with the territory, and this, it seems to me, is a fact of some importance; for if the fighting were merely an exuberant manifestation of s.e.xual emotion, one would expect to find it occurring under all conditions, and not merely under one particular condition in the life of the bird. The hostility is too widespread, however, and too uniform in occurrence for us to suppose that it has no root in the inherited const.i.tution of the bird; and if it served some useful purpose in the past, the instinct might still persist, so long as it were not harmful. Thus the view that the behaviour is vestigial is not perhaps unreasonable. But manifestly it makes no difference whether it be vestigial or a by-product of s.e.xual emotion, whether the battle be fierce or so trivial as to appear to us to be more in the nature of "play," so long as some change in the relative prospects of the opponents is the result.

For us, then, the main consideration lies in the question: Is the behaviour serviceable now in furthering the life of the individual?

Whether the evidence which we have examined affords sufficient ground for the belief that the hostility is genuine and has a part to play in the whole scheme of reproduction, each must judge for himself.

CHAPTER VII

THE RELATION OF THE TERRITORY TO MIGRATION

Coincident in time with the growth of appropriate conditions in the environment, organic changes take place rendering certain instincts susceptible to stimulation; and the stimulus being applied, the Warbler leaves the country wherein it had pa.s.sed the winter and finds its way back, with apparently little difficulty, to the district in which it was reared or had previously reared offspring. What is the nature of these changes and of the impulse which is first brought into functional activity; whence comes the stimulus; and what directs the bird on its journey--these are all different aspects of one great problem, the problem of migration. I do not propose to discuss all these various aspects, for indeed I have no suggestions to offer which are in the least likely to be helpful, but I seek rather to ascertain whether the phenomena which we have explored bear any relation to the problem as a whole; whether, that is to say, the compet.i.tion for territory and all that appertains to it can have supplied the conditions under which, in the process of time, this complex and definite mode of behaviour has evolved.

We are sometimes told that we must seek the origin of migration in the physical changes that have occurred in the ancient history of the earth--in glacial conditions which gradually forced birds to the south, or in the "stability of the water and mobility of the land" which brought about a gradual separation of the feeding area from the breeding area--and which continued for a sufficient length of time to lead to the formation of an instinct, and that the instinct persists because it is serviceable in promoting the welfare of the race. 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 a.s.suredly no less strong, and the volume of migration, if not greater, is a.s.suredly no less; in short, that the whole phenomenon is progressive rather than retrogressive, we shall find the view that the instinct owes its origin to conditions which no longer exist, receives but little encouragement.

I doubt not that, throughout the ages, geological changes have been an important factor in directing or limiting the scope of migration, and moreover are so still; just as climatic changes and the relative abundance or scarcity of enemies have influenced the course of its evolution. These are all contributory factors operating in the external environment. But there are, besides, internal factors which form part of the inherited const.i.tution of the bird, and, being pa.s.sed on from generation to generation, afford the conditions under which migration is constantly being renewed. It is, I believe, in this field of organic change and relationship that the conditions of origin must be sought.

Just as the moth in pa.s.sing from the rudimentary to the perfect condition runs through a series of changes, each one of which is marked by a typical behaviour response adjusted to meet some particular circ.u.mstance in the external environment, so the annual history of a bird displays an ordered routine, each phase of which can be observed to correspond with one of the successive changes in the environment. In almost every direction, we find that this routine is characterised, in broad outline, by great uniformity; so much so that, providing we know the history of one species, we can forecast with no small degree of certainty the general course of behaviour of other members of the family. But only the _general_ course. There is endless variation in just the particular way in which the behaviour is adapted to meet the needs of particular species--the major details may be said to be specific, the minor details varietal.

Now it is that part of the behaviour routine which has reference to the relationship between one bird and another upon which, for the time being, I wish to dwell; for the interest that A displays in B is by no means always the same--it changes according to the season, and this change can be observed to be uniform throughout a wide range of species.

In winter, in whatsoever direction we turn, we observe not only that different individuals but that different species also collect together in flocks. And since food at that season is not always easy to obtain, and, moreover, is only to be found in certain situations, which are limited both in number and extent, it would seem that such a.s.semblages are in the main determined by accident. No doubt the abundance or the scarcity of food does determine the movements of birds, and hence to that extent may be held to account for the flocks. But we shall but deceive ourselves if we think that it is the sole or even the princ.i.p.al reason, or that the situation is in no wise affected by internal factors. The behaviour of the individual in relation to the flock bears ample testimony to the presence of a gregarious impulse which derives satisfaction from the fact of close a.s.sociation.

As an ill.u.s.tration, let us take a bird whose movements are easily watched, and in whose hereditary const.i.tution the impulse to which I allude seems to be strongly implanted--the Curlew. When the breeding season is over, Curlew leave the mountain and the moor and return to the coast or tidal estuaries for the remainder of the year. Here, at low water, they find an abundant supply of food--crustaceans amongst the sea-weed upon the rocks, and lobworms (_Arenicola piscatorum_) in the mud as the tide advances or recedes. But when the tide is full, they retire to those parts of the sh.o.r.e that remain uncovered--to isolated rocks, or to sand-dunes, or it may even be to pasture-land in the neighbourhood. During this period of repose large numbers of individuals gather together on a comparatively small s.p.a.ce of ground. They are not constrained to do so by any shortage of accommodation, nor by any question relative to food, nor, for the matter of that, by any circ.u.mstance in the external environment; they are brought together solely, this at least is the impression that one gains, by some inherited impulse working towards that end. And their subsequent course of behaviour tends to confirm that impression. For if we watch the gathering together of the different units of which the flock is composed, and study more particularly the emotional manifestation which accompanies their arrival and departure, we shall find that the coming of a companion arouses some emotion which is expressed by a vocal outburst that sweeps through the flock.

Now each call, and the Curlew has a great variety, is not only peculiar, generally speaking, to certain occasions, but is accompanied by a specific type of behaviour, whence we can infer in broad outline the type of emotion which is aroused. Thus we come to recognise fear, anger, or s.e.xual emotion, by just the particular sound which is emitted. But even if we are going too far in referring particular calls to particular emotions, we can, without a doubt, divide them into two broad categories--those which are pleasurable and those which are the reverse.

And we need have no hesitation in placing the particular call to which I allude in the first of these two categories, not only on account of the nature of the sound produced, but because the activities which are aroused are not such as normally accompany irritation. This is well seen if the behaviour of different individuals be closely observed. After resting on one leg for some time, first one and then another is seized with cramp, and running a few yards in an ungainly way, b.u.mps up against its companions as if it had not full control over its movements. Its behaviour produces irritation which is expressed by a vocal outburst, and followed by actions the meaning of which is clear. Moreover, the call is taken up by other individuals and sweeps over part of the flock as does the greeting. But the nature of the cry is entirely different from that which greets the arrival of a companion--humanly speaking it is a pa.s.sionate and impatient utterance, the height of displeasure. The arrival, then, acts as a stimulus to something in the inherited const.i.tution which is expressed in, and presumably is satisfied by, this vocal outburst; and, since the bird that arrives joins also in the chorus, there is reason to think that the impulse which determines its movements is similar to that which is temporarily aroused in the flock.

Apart, however, from the evidence derived from the affective aspect of the operation of the instinct, the general course of behaviour lends support to the view that the a.s.semblies are determined by internal factors, and are not merely the outcome of circ.u.mstances in the external environment. Observe, for example, the manner in which the flock is built up. Single individuals are content to rest alone so long as no a.s.sembly is in sight, but they are drawn towards their companions directly the opportunity arises, just as surely as the smaller aggregation is drawn towards the flock; and so, as the flock increases, it gradually absorbs all the lesser flocks and smaller parties, for the greater the flock the greater the attraction seems to be; and different individuals appear to gain some satisfaction from being in close bodily contact with one another.

When the Curlew flies to that part of the mud-flat which is first exposed by the receding tide, and there a.s.sociates with others, it does not then do so because it has any interest in its fellows, nor because they serve as an attraction, but because it is constrained by hunger--in other words, the a.s.sociation is determined by accident. But when, during periods of repose, it sees a flock, flies to it, and takes up a position in the midst of it, it does so not because suitable accommodation is lacking--not therefore because of external constraint--but because it derives some pleasure from satisfying something in its organic complex.

We speak of this behaviour and of the emotion which characterises it as the _gregarious instinct_: by which we mean that the inherited nature of the Curlew, as a tribe, is so const.i.tuted that, given the appropriate internal conditions and adequate external stimulation, every individual will respond in a similar manner--that is, the behaviour is primarily determined by racial preparation. This is what we mean by the _gregarious instinct_ biologically considered. We may resolve our own experience in relation to the crowd into its simplest const.i.tuents, project our own primitive feelings into the Curlew, and say that the bird feels uneasiness in isolation and satisfaction in being one of the flock. But in truth we know nothing, save by a.n.a.logy, of the correlated psychical state. All the knowledge we possess is derived from a study of the objective aspect of the behaviour, which in simple terms may be expressed thus: the individual is drawn towards its companions; there is a relation between the size of the flock and the strength of the attraction; and all Curlew behave similarly under similar circ.u.mstances.

This instinct controls the movements of many birds from early autumn to the commencement of the breeding season. And so powerful is the control that the individual is suppressed and its activities subordinated to the welfare of the community as a whole. Flocks of Waders roam about the tidal estuaries in search of food, and different kinds of Gulls a.s.semble there and preen their feathers or sleep; 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; and throughout the winter continues to be the outstanding feature of bird life.

With the approach of the breeding season we witness that remarkable change which I have endeavoured to make clear in the previous chapters--the disintegration of the flock and the reinstatement of the individual. Instead of continuing with the flock, the individual now goes forth to seek the appropriate breeding ground; and having arrived there, is not only content to remain in isolation, but so behaves that isolation is insured. Intolerant of the approach of a stranger, intolerant even of the approach of the very members of the community whose companionship was previously welcomed, it not only fights to maintain the position it has selected, but fights indeed for the possession of ground already occupied, and, until reproduction is completed, a.s.serts its individuality and exercises dominion over its territory. What, then, is the prospective value, biologically considered, of the changing interest that A displays in B, and to what will such changes lead? These are the questions to which we will now direct inquiry.

The annual life-history of a bird is in broad outline conditioned by two powerful and at first sight opposing impulses--the one to live in society, the other to live solitary. But, manifestly, a bird cannot be governed by opposing impulses. It has but one character, within which, according to the season and the circ.u.mstances, different impulses predominate. But these impulses, no matter how different they may appear to be, have their respective parts to play in furthering the life of the individual. Hence they cannot oppose, though they may conflict, if the resultant behaviour contributes towards survival.

The majority of birds live to-day in constant danger from predatory species, and that this danger was still greater in bygone ages there can be but little doubt. A curious mode of behaviour of the Curlew, Whimbrel, and G.o.dwit demonstrates this, for it must be the outcome of the necessity for constant watchfulness. Whilst resting with its head turned back and its beak buried in the feathers of the mantle, the bird constantly moves the axis of its body, so that an observer, if placed in a direct line behind it, sees at one moment the right eye and at another the left. No movement of the feet or of the legs is perceptible, and the shifting of the body continues whether the eyes are open or closed. This body movement enables the bird to survey a much larger area of ground than it would otherwise be capable of doing, and thus adds to its security. 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. And not only do the different members give warning one to another of the approach of danger, but they also combine to hara.s.s or even to drive away an enemy. So that there can be no doubt that the gregarious instinct is serviceable in promoting the welfare of the race, and has, as its end, the preservation of the individual in order that it may take its share at the appropriate time in procreating its kind.

In winter, then, the individual loses its individuality and is subordinated to the welfare of the community, whilst in spring it regains its individuality, and all its inherited instincts which then come into operation lead to its isolation from the flock. The impulse to seek isolation is dependent upon internal organic conditions which are peculiar to a certain season; whereas the gregarious impulse depends upon internal organic conditions which inhere at all times, though its functioning is inhibited by the functioning of the former impulse. The evidence which leads to this conclusion is to be found in the fact that a male often deserts its territory temporarily and joins the flock, where it remains at peace with its companions--an aspect of behaviour which we have discussed on various occasions. The former impulse becomes dominant in the spring owing to its innately superior strength; the latter becomes dominant in the autumn because the organic condition which determines the functioning of the former then subsides. The impulse to seek the appropriate breeding ground and to dwell there would seem to be the strongest of all the impulses save one--the s.e.xual.

When, however, I speak of the s.e.xual, I refer to the actual discharge of the s.e.xual function, which is the consummation of the whole process. But the territory and all that appertains to it is part of that process--the search for the breeding ground, the dwelling there, and the intolerance of intrusion are but different stages, each one of which must have an impulse peculiar to it; and since the completion of the s.e.xual act can only be successfully accomplished providing that success is attained at every stage, the probability is that, of the impulses concerned, one is neither more powerful nor less powerful than another.

So that we have two impulses operating at different seasons and guiding the behaviour into widely divergent channels. But though the proximate end to which the behaviour is directed is apparently different, there are not two biological ends in view, but one--the attainment of reproduction; and the changes that we witness are not contrary but complementary, and their prospective value lies in the circ.u.mstance that they contribute towards the preservation of the race.

If, then, every male is driven by inherited impulse to seek the appropriate breeding ground each recurring season; if, having arrived there, it is driven to seek a position of its own; if, in order to secure isolation it is obliged to attack other males or to ward off the attacks of intruders; if, in short, success can only be attained providing that the inherited nature is so adjusted that the bird can accomplish all that is here demanded--what will be the general result?

That the individual will rear its offspring in safety and that they will inherit the peculiarities of their parents, enabling them, in their turn, to procreate their kind; all this will certainly follow. We are not concerned, however, at the moment, with the direct effect upon the individual, but with the consequences that will accrue to the species as a whole.

Now certain facts are presented to observation which enable us not only to understand the nature of the change that is wrought in the history of the species, but to foreshadow, with no small degree of certainty, the extent of that change. I suppose that it has come within the experience of most of us to observe, at one time or another, the ebb and flow of a given species in a given district. Some favourite haunt is deserted for a year, or for a term of years, and is then revisited; or, if it is always occupied, the number of inhabitants fluctuates--plenty of pairs in this season, only a few in that. Many intricate relationships, both external and internal, contribute towards this state of affairs.

Fluctuation in a downward direction, or temporary extinction, is brought about by changes in the physical world, by changes in the available supply of food, by the increase of enemies, or by adverse climatic conditions; whilst fluctuation in an upward direction, though due indirectly to a combination of circ.u.mstances in the external world favourable to the survival of large numbers of individuals, is directly determined by the impulse to seek isolation. As individuals of different species establish themselves, and form kingdoms and lesser kingdoms, we can watch the gradual quickening into life of moorland and forest and we can observe the manner in which it all comes to pa.s.s.

Males that for weeks or months have lived in society, drifting from locality to locality according to the abundance of food or its scarcity, now set forth alone and settle first here and then there in search of isolation. Lapwings settle in the water meadows, and, finding themselves forestalled, pa.s.s on in search of other ground; Blackbirds arrive in a coppice or in a hedgerow and, meeting with opposition, disappear; and the Curlew, wandering with no fixed abode but apparently with a fixity of purpose, searches out the moorland where it can find the particular environmental conditions to which its inherited nature will respond. In fact, wherever we choose to look, we can observe in a general way the gradual appropriation of breeding ground; and if we fix our attention upon particular males, we can watch the method by which success or failure is achieved.

On more than one occasion I have watched the efforts of Reed-Buntings to appropriate territories in a marsh that was already inhabited. Sometimes their efforts met with success, at other times with failure. In the former case, the males, whose ground was intruded upon, were severally forced to yield part of their holding and were thus left in possession of a smaller area. The success of the intruder seemed to depend upon persistent determination, rather than upon superior skill in battle.