What meaning does the song convey to a male that is unestablished? Does the bird recognise that it is forestalled; does it foresee and fear the possibility of a conflict, and conclude that the attempt to settle is not worth while? I do not imagine that it thinks about it at all. How then does the warning warn? We will endeavour to answer this question, but, in order to do so, we must review the stages by which a territory is secured.
We take as our starting point the internal organic changes which are known to occur. These changes are correlated with other changes, manifested by a conspicuous alteration in behaviour--to wit, the disappearance of sociability and its replacement by isolation. Having found a station which meets the requirements of its racial characteristics, the male establishes itself for a season, becomes vociferous, displays hostility towards others of its kind, and in due course is discovered by a female. The whole is thus an inter-related whole, a chain of activities which follow one another in ordered sequence. Now we have seen that it is neither pugnacious nor vociferous until the territory is actually occupied; we have seen that the fact of occupation is the condition under which the instincts of pugnacity and of song are rendered susceptible to appropriate stimulation; we have discussed the nature of the stimulus in each case, and we wish to know the sort of meaning that the song conveys to an individual which is still in the preliminary stage of seeking a station. In sequential order we have the following: (1) internal organic changes which lead to isolation, (2) the appropriate environment which gives rise to an impulse to remain in it, (3) the occupation of a territory which is the condition under which the instincts are rendered susceptible to stimulation, (4) the various stimuli. Each is dependent upon that which precedes it, and no part can be subtracted without failure of the biological end in view, neither can the different stages be combined in different order. So that, in considering the significance of song to an unestablished male, we are dealing with the situation at a point at which all the latent activities have not been fully felt, for all that so far has occurred is the change from sociability to isolation determined by internal organic changes. The bird has not established a territory because it has not come into contact with the appropriate environment, and it is not pugnacious because the condition which renders its instinct susceptible is absent; and so, as it wanders from place to place and hears the voices of males here or males there, it merely behaves in accordance with that part of its nature which predominates just at that particular moment--the impulse to avoid them.