Henry Eliot Howard

“ There cannot be territories without boundaries of some description; there cannot well be boundaries without disputes arising as to those boundaries; nor, one would imagine, can there be disputes without consciousness as a factor entering into the situation; and so on, until by a simple mental process we conceive of a state in bird life analogous to that which we know to be customary amongst ourselves. ”

“ Nevertheless, even though she may have discovered a male ready to breed, success is not necessarily assured to her; for with multitudes of individuals striving to procreate their kind, it would be surprising if there were no clashing of interests, if no two females were ever to meet in the same occupied territory. Competition of this kind is not uncommon, and the final appeal is to the law of battle, just as an appeal to physical strength sometimes decides the question of the initial ownership of a territory. ”

“ Competition doubtless varies at different periods and in different districts according to the numerical standing of the species in a given locality and according to the numerical standing of others that require similar conditions of existence; at times it may even be absent, just as at any moment it may become acute. These examples show how profoundly the evolution of the breeding territory may have been influenced by relationships in the inorganic world, and they give some idea of the intricate nature of the problem with which we have to deal. ”

“ Nevertheless, if the battles between males of the same species are directly related to the occupation of a position suitable for breeding purposes, if those which occur between males of closely related forms can be traced to a similar source, if the females take their share in the defence of the ground that is occupied, if, in short, the competition is as severe as I believe it to be, and is wholly responsible for the strife which is prevalent at the commencement of the breeding season—then such competition must have introduced profound modifications in the distribution of species ”

“ Nevertheless if every male were to make use of its powers whether it were in occupation of a territory or not, if the wandering individual had an equal chance of attracting a mate, then it would be idle to attempt to establish any relation between "song" on the one hand, and "territory" on the other, and impossible to regard the voice as the medium through which an effectual union of the sexes is procured. But there is reason to believe that the males utilise their powers of producing sound only under certain well-defined conditions. ”

“ There are direct relationships, such as we have been speaking of, which are essential to the every-day working of the process, and others which are indirect, though none the less important for they must have exercised an influence throughout the ages. These latter are furnished by the physical—the inorganic world, by climate, by the supply of the particular kind of breeding stations, by the scarcity or abundance of the necessary food and by the relative position of the food supply to the places suitable for breeding. ”

“ The answer is the same in both cases—to facilitate reproduction. But why should a small bird require so many square yards, whilst a very much larger one is satisfied with so small an area? The explanation must be sought in the conditions of existence. The Reed-Bunting has no difficulty in finding a position suitable for the construction of its nest; there are acres of waste land and reedy swamps capable of supplying food for large numbers of individuals, and the necessary situations for countless nests. But its young, like those of many another species, are born in a very helpless state. ”

“ All such adjustments have, however, been brought about by relationships which have gradually become interwoven in the tissue of the race.
The intolerance that the male displays towards other individuals, usually of the same sex, leads to a vast amount of strife. Nowhere in the animal world are conflicts more frequent, more prolonged, and more determined than in the sexual life of birds; and though they are acknowledged to be an important factor in the life of the individual, yet there is much difference of opinion as to the exact position they occupy in the drama of bird life. ”

“ But since we know so little of the organic changes which determine sexual behaviour, and have no means of ascertaining the nature of the impulse which is first aroused, we can only deal with the situation from the point at which the internal organic changes reflect themselves in the behaviour to a degree which is visible to an external observer. That point is reached when large numbers of species, forsaking the normal routine of existence to which they have been accustomed for some months, suddenly adopt a radical change in their mode of behaviour. ”

“ The bird takes a short flight, searches a bush here and some rushes there, returns, and after a while repeats the performance; we on our part mark the extreme limits reached in each direction, and by continued observation discover that these limits are seldom exceeded, that definition grows more and more pronounced, and that by degrees the movements of the bird are confined within a restricted area. ”

“ We find, however, in the migrants, that the males are earlier than the females in reaching the breeding grounds, and, in resident species, that they desert the females and retire alone to their prospective territories, so that there is a difference in the behaviour of the sexes at the very commencement of the sexual process. What is the immediate consequence? Since the male isolates itself, it follows, if the union of the sexes is to be effected, that the discovery of a mate must rest largely with the female. ”

“ Indirectly its purpose is that of the whole process, the rearing of offspring. But inasmuch as a certain measure of success could be attained, and that perhaps often, without all the complications introduced by the territory, there are manifestly advantages to be gained by its inclusion in the scheme. The difficulties which beset the path of reproduction are by no means always the same—all manner of adjustments have to be made to suit the needs of different species. ”

“ I suppose, because the competition thus created would be a means of maintaining efficiency, the females, by common consent, are supposed to supply the condition under which the pugnacious nature of the male is rendered susceptible to appropriate stimulation. And so long as the evidence seemed to show that battles were confined to the male sex, so long were there grounds for hoping that their origin might be traced to such competition. But female fights with female, pair with pair, and, which is still more remarkable, a pair will attack a single male or a single female ”

“ But manifestly all will not be in a like satisfactory position to accomplish this, for whereas the isolated pairs will have free access to all the food in the immediate vicinity of the nest, those which have built in proximity to one another, meeting competition in every direction, will be compelled to roam farther afield and waste much valuable time by doing so; and under conditions which can well be imagined, even this slight loss of time will be sufficient to impede the growth of the delicate offspring, or to lead perhaps to still greater disaster. ”

“ And since such a radical departure from the normal routine of behaviour could scarcely appear generation after generation in so many widely divergent forms, and still be so uniform in occurrence each returning season, if it were not founded upon some congenital basis, it is probable that the journey, whether it be the extensive one of the Warbler or the short one of the Reed-Bunting, is undertaken in response to some inherited disposition, and probable also that the disposition bears some relation to the few acres in which the bird ultimately finds a resting place. ”

“ The amount of space occupied by each individual is therefore a matter of urgent importance. A few square feet of rock sufficient for the immediate purpose of incubation is all that can be allowed if the species is to maintain its position in the struggle for existence. ”

“ 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. ”

“ For instance, when they are on their way to the breeding grounds, or moving from locality to locality in search of isolation, or when they desert their territories temporarily, as certain of the residents often do, they are generally silent; but when they are in occupation of their territories they become vociferous—and this is notoriously the case during the early hours of the day, which is the period of maximum activity so far as sexual behaviour is concerned. ”

“ An acquired mode of activity becomes by repetition ingrained in the life of the individual, so that an action performed to-day is liable to be repeated to-morrow so long as it does not prejudice the existence or annul the fertility of the individual. ”

“ If, then, different individuals were to jostle one another from adjoining positions, and each one were to attempt to occupy a ledge in solitary state, not only would the successful ones gain no advantage from the additional space over which they exercised dominion, but inasmuch as many members that were fitted to breed would be precluded from doing so, the status of the species as a whole would be seriously affected. ”

“ Its young at birth are by no means helpless in the sense that the young Reed-Bunting is, and food is readily procured. But breeding stations are scarce, for although there are many miles of cliff-bound coast, yet not every type of rock formation produces the fissures and ledges upon which the bird rests. ”

“ For all practical purposes they are without covering of any description and consequently require protection from the elements, warmth from the body of the brooding bird, and repeated supplies of nourishment. A threefold burden is thus imposed upon the parents: they must find food for themselves, they must afford protection to the young by brooding, and they must supply them with the necessary food at regular intervals. ”

“ With the majority of species it is difficult to do this. So many square miles of suitable breeding ground are inhabited by so few ReedBuntings that, even supposing certain members were to establish an ascendency over too wide an area, it would be impossible to discover by actual observation whether the race as a whole were being adversely affected. ”

“ Hence vast stretches of coast-line remain uninhabited, and the birds are forced to concentrate at certain points, where year after year they assemble in countless numbers from distant parts of the ocean. ”

“ The same result can be obtained without our having recourse to so complex a principle of explanation, and that by the law of habit formation. In common with other animals, birds are subject to this law in a marked degree. ”

“ In due course eggs will be laid and incubation performed, and thus far all alike will probably be successful. Here, however, a critical point is reached. If the young are to be freed from the risk of exposure, the parents must find the necessary supply of food rapidly. ”

“ By vast numbers of individuals hurrying from one part of the globe to another, from one country to another, and even from mid-ocean to the coasts; by detachments travelling from one district to another ”