Henry Eliot Howard

“ The question now arises as to how it comes about that the area occupied by each individual conforms in broad outline to that which has proved beneficial for the welfare of the species as a whole. We shall find that up to a point the answer is a simple one. No one could study the behaviour of animals without observing the important part that habit plays in the life of the individual; an action performed to-day is liable to be repeated to-morrow and the following day until it becomes ingrained in the life of the individual. ”

“ What is the biological value to the Guillemot of an inherited nature which, for the Bunting, has utility in relation to the supply of food for the young? Up to a point, the act of securing a territory has like value for each respective species, whether the area occupied be large or small—that is to say, it enables the one sex to discover the other with reasonable promptitude. ”

“ I suggest that its evolution has afforded the condition under which this beneficial procedure has developed—free to mate when they will, free to seek food where they will, free to pursue their normal routine of existence, and to meet all exigencies as they arise in their ordinary daily life—whilst free to do this, their future, as a pair, is nevertheless secure. ”

“ Hence, while preserving freedom of movement for each individual, the territory will render their future, as a pair, secure.
No doubt the course of behaviour, as we observe it to-day in the lives of many species, is the outcome of, rather than the condition which has led to, the evolution of the territory. Thus, in many cases, we find that early mating is the rule rather than the exception; we find that the sexes frequently separate to seek their food, and fly away temporarily in different directions ”

“ For a bird, success in the attainment of reproduction does not merely imply the successful discharge of the sexual function; much more is demanded; it must find somewhere to build its nest and to lay its eggs, it must shield its young from extremes of temperature and protect them from enemies, and it must be in a position to supply them with food at regular intervals. ”

“ There is a gorse-covered common which I have in mind, a favourite breeding resort of this species. Between this common and the surrounding country, the birds constantly pass to and fro. If you watch a particular male you will observe that it sings for a while in its territory, that it then rises in the air and disappears from view, and finally that it returns to the tree, bush, or mound which constitutes its headquarters, where it again sings. ”

“ There are many species whose success in rearing offspring is largely dependent upon the rapidity with which they can obtain food; and it makes but little difference which species we choose out of many—Finch, Bunting, Warbler, or Chat. I shall choose the Buntings, as their life-history in broad outline conforms to the general type, and, moreover, their behaviour is fresh in my mind. The young are born in a very helpless state ”

“ We have, then, the congenital basis which leads to the occupation of a position, and to the enmity shown by the owner of the position towards other individuals; and this congenital basis is found alike in many widely divergent forms, living under equally widely divergent conditions; we have acquired accommodation; and we have relationships in the organic and inorganic world—and the outcome of it all is a system of behaviour which we, who can perceive the end to which such behaviour is tending, are justified in speaking of as "a disposition to secure a territory." ”

“ A community, however, in the true sense of the word, is a collection of individuals brought together, not primarily as a result of shortage of breeding ground, but in consequence of advantages of communal ownership over individual ownership. ”

“ But every locality cannot supply sufficient trees of the right kind, appropriately situated and in suitable relation to the food supply, in which numbers of nests can be built in close proximity; so that if more than one community were to attempt to establish itself in a limited area, the supply of food or the supply of trees might become a pressing problem. Each community must therefore be prepared to defend its own interests, and each must be regarded as one unit and the area occupied as one territory within which are included a number of lesser territories. ”

“ Now when we bear in mind how large an expanse of coast is formed of blown sand or of rocks of low altitude, and how many miles of cliff fail to supply the three essential conditions that we have been considering, we can see that suitable breeding stations must be limited both in number and extent. From a wide expanse of ocean hosts of individuals are therefore obliged to converge at certain definite points; and hence, each recurring season, there must arise a competition for positions at the station, just as there is competition between individual Buntings for positions in the marsh. ”

“ Let us see how far this law of habit formation may have been effective in defining the extent of the area occupied. When a male Warbler arrives at its destination in the spring it seeks out a suitable environment, and, having found a place unoccupied by any other male, settles in it and remains there—its behaviour up to this point being determined by racial preparation. After the fatigue of the journey its movements are at first sluggish; hunger, however, asserts itself and a search is made for food ”

“ The life of the Guillemot during the winter is a strenuous one; we know that large numbers succumb in stormy weather, and we can infer that slight constitutional defects might make all the difference between failure and success; and, therefore, the less severely the constitution of the parent is taxed during reproduction, and the more securely the constitution of the offspring is built up, the greater prospect will both have of resisting the hardships of the winter successfully. Much, then, will depend upon the distance the parents have to travel in order to obtain food. ”

“ Nevertheless the isolation of the individual is, in a sense, just as complete as that of the individual Bunting, for each one is just as vigilant in resisting - intrusion upon its few square feet as the Bunting is in guarding its many square yards, so that the evidence seems to show that that part of the inherited nature which is the basis of the territory is much the same in both species. ”

“ But all the young cannot be fed, neither are they ready to be fed, at the same moment; and the parents have besides to find food for themselves, and the nest has to be cleaned—all of which necessitates the young being exposed to the elements at frequent intervals. Now it is impossible to observe the instinctive routine, of the parents, when the young need attention, without being impressed with the conative aspect of their behaviour. Why, we ask, are the movements of the female so brisk; why does she seek food and clean the nest so hurriedly ”

“ The individual may fail to establish itself within a community, but, even if it succeeds, the community may fail to establish the rights of communal ownership; hence it has to face a twofold possibility of failure, and if it lacked the inherited nature which leads the Guillemot to secure a position upon the ledge, or the Bunting to obtain a position in the marsh, the chances are that it would fail in the attainment of reproduction. ”

“ But though the nest is in full view and within a few inches of her perch, and though the young stretch out their necks, yet so strong is the former habit that she is compelled to return to the opposite side and approach the nest by the usual circuitous route before she distributes the food amongst her offspring. ”

“ But it must not be supposed that each pair finds, or even attempts to find, the whole of the food within its territory, or that it is necessary for the theory that it should do so; all that is required is that such overcrowding as might lead to prolonged absence on the part of the parents and inordinate exposure of the young shall be avoided. So that the problem has to be considered not merely from the point of view of the individual, but from the larger point of view of all the pairs inhabiting a given area. ”

“ Hence cliffs which are situated away from the water, or from which the water recedes at low tide, or which are surrounded by an area of shallow water, and are thus not in proximity to the feeding ground, even though they may fulfil the second and third condition, will not answer the requirements of a breeding station.
Of no less importance is the type of rock-formation. Not every formation affords the necessary ledges upon which the egg can be deposited with safety—the face of the cliff may be too smooth, or too jagged, or the shelves may run at too acute an angle. ”

“ Sometimes male and female accompany one another—that is, they leave simultaneously and likewise return; at other times, though they depart together, the male returns alone; or the male may disappear in one direction whilst the female does so in another—and, on the whole, there is a sameness in the direction of flight taken by the same pairs on different occasions. ”

“ And so, when we come to consider the question of reproduction in the higher forms of life, we are apt to focus attention too much upon the sexual function and too little upon the contributory factors, the failure of any one of which would mean failure of the whole. ”

“ But the use of this word is nevertheless open to criticism, for it denotes a human end upon which the highest faculties have been brought to bear, and consequently we have to be on our guard lest our conception of the "territory" should tend to soar upwards into regions which require a level of mental development not attained by the bird. It is necessary to bear this in mind now we have come to consider the meaning of the territory, or rather the position that it occupies in the whole scheme of reproduction. ”

“ Each Warbler seeks its food within the precincts of its own particular domain, and, except in occasional instances, neither resorts to neutral ground nor makes excursions into the locality immediately surrounding the territory, as does the Bunting. Probably it would be disastrous if it attempted to do so, for since its young at birth are so delicate and so susceptible to changes of temperature, it cannot afford to be absent from them for long. Of the two experiments made with young Whitethroats, one was made under favourable and the other under unfavourable conditions. ”

“ Next, after mating has taken place, the position occupied by the male acts as a headquarters to which the birds can always repair, and becomes a bond of union which is serviceable in that it prevents any possibility of their drifting apart. And in the third place, the males become pugnacious and in this way secure for themselves areas which vary in size according to the conditions of existence of the species, so that there is no possibility of too many congregating in this locality, and all the less likelihood of too few finding their way to that ”

“ How, then, can the theory apply to a species that breeds under such conditions? What part can the territory play in furthering the life of the individual when large numbers of nests are built closely together in the same tree? ”

“ A rookery is an example of a true community. Neither shortage of nesting accommodation nor scarcity of food can account for Rooks assembling together to breed; for if the different pairs which go to make up the rookery were to scatter throughout the surrounding neighbourhood, they would, as a rule, find plenty of trees in which to build their nests, and plenty of food. ”

“ Now the power of locomotion is so highly developed in birds that it may seem unreasonable to suppose that males and females would have any difficulty in meeting when their inherited nature required that they should do so, still less reasonable to suggest that this power might even act as a hindrance to successful mating. Nevertheless, if we try to picture to ourselves the conditions which would obtain if the movements of both sexes were in no wise controlled, and mating were solely dependent upon fortuitous gatherings, we shall come. ”

“ Either the face of the cliff is made up of a series of broken precipices, or the face of the precipices is too smooth, or the otherwise suitable ledges are situated too near the water, or the water recedes from the base of the cliff at low tide. Many miles of rock-bound coast are thus useless for the purpose of reproduction. ”

“ Next, let us consider their position were a disposition to establish a territory to form part of the inherited nature of the male. Each one will then be free to seek food when and where it wills and to associate with other individuals without the risk of permanent separation from its mate; and, no matter how long an interval may elapse between mating and nest-building, each one will be in a position to find the other when the appropriate moment for doing so arrives. ”