The Courtship of Animals

by W. P. Pycraft

In the males of all healthy animals the periodic stimulus to reproduction finds expression in more or less striking eccentricity of conduct. Sometimes, as the foregoing instances have shown, this has been exaggerated by the development of long, resplendent plumes: sometimes by brilliant coloration, displayed either by the plumage or by bare areas of skin, or by both, while in not a few cases attitudes, to our eyes grotesque and made still more so by the aid of inflatable pouches, are the outward and visible sign of the raging fires within. For the completion of this chapter yet other instances of this kind must be cited, instances which reveal a further elaboration of some of the more striking of these tricks of posturing; or which concern the growth of the aggressive instincts, which are proclaimed by the development of armature often of a very formidable character. As the sequel shows, however, there are no hard and fast dividing lines between these several modes of expression.

That remarkable bird, the Ruff (Machetes pugnax), now, alas! no longer to be met with in our fens, exhibits a curiously composite character in the phases of its love display.

Preparations for this are begun in the early spring by the assumption of what is called a “nuptial dress,” which is worn only by the male, and which contrasts in a very conspicuous manner with the plumage worn during the rest of the year. The most striking features of this dress are the great, erectile, Elizabethan ruff which encircles the neck immediately behind the head, and the long, tongue-shaped “ears” which surmount the head itself. These exhibit a most remarkable diversity in their coloration, and it is no exaggeration to say that no two are ever alike. Red, cream-colour, buff, black, white; spotted, streaked, freckled and barred are the only descriptions that can be applied to them, for the combinations of their hues and patterns seem infinite. Having grasped this fact, the eye next turns to the colouring of the rest of the body, and it will be found that here too is the same diversity, though less conspicuously so; and finally it will be noticed that at this time the feathers around the base of the beak have been replaced by yellow or orange-coloured papillæ. The females also now wear a dress differing from that of the so-called “winter plumage,” but it does not present any very striking features nor any form of ornamentation comparable to that of the males.

The Ruff is a polygamous bird, which, in its display, presents some curious and puzzling features, one of which consists of a sort of tournament between rival males. At the break of day the performers, selecting such eminences as the fen-lands afford, assemble apparently to display their finery, for a couple of males will often stand facing one another with frills erected and beaks touching the ground, silent and immovable, for perhaps half a minute. Sooner or later, however, they will commence to spar, and this presently leads to blows, during which one of the combatants will attempt to seize the other by the wings. However, no damage seems to be inflicted during such encounters, which are by no means aimless or profitless, for during such bouts the weaker, less vigorous birds are driven from the field, and the victor in consequence wins for himself a larger harem.

When the actual pairing time arrives the parade of the frills begins again. The amorous instincts, it is important to notice, are awakened earlier in the males, so that by the time the females have attained to a like condition the least mettlesome males have been driven off. What follows is not the selection by the females of the finest performers so much as a process of sorting out, whereby the females discover and cleave to those males which are readiest for mating. This display succeeds in revealing both the most mettlesome males and the most amorous females, who, however, would seem to require great persistence and much demonstration on the part of the males before they can be finally aroused to the pitch necessary for pairing. Again and again a male may be seen to approach an apparently very unconcerned female, and then to crouch down before her with his beak pressed to the ground and his frill and “ears” set off to their fullest. For some seconds he will remain lost in apparent contemplation, then with a dazed, far-off, expression he will look up, to find, as often as not, that she is still apparently feeding, quite unmoved by his protestations; or that she has even flown off and left him. Pursuit speedily follows, and the performance is repeated until at last she too catches the flame of passion and permits, or rather invites, the final act of sexual congress.

Though these birds on occasion will fight, and savagely, they cannot inflict serious damage on one another by reason of the relative feebleness of their beaks and legs, which are but ill-adapted for violent measures. Inasmuch as the Ruff is a polygamous species, these bloodless battles have a peculiar interest. They show that the preponderance of females, which polygamy implies, is not, as is commonly supposed to be the case, due to a high death-rate among the males by fighting. The same is true of the Wydah-birds, and their kin, the only polygamous species among the Passeres.

In this connection it is to be remarked that fighting, of a more or less sanguinary character, is apparently universal among birds, the conflicts being waged not so much in the way of squabbles for the possession of females as for the acquisition and retention of territory and all that this entails during the breeding season and, to a much less extent, in the defence of the eggs and young. But to this point we must return. For the moment it will be more profitable to focus attention on the character of this fighting. In the first place, it is by no means necessary that the combatants should be armed. The “dove of peace” at this time of the year appears in a new and not always pleasing light, for not only will he fight his neighbours, but he does not always show that gentleness towards his wife with which tradition has credited him. The little Humming-bird would seem to be as little capable of fighting as a bird could be, yet few are more pugnacious. The naturalist Gosse tells of a pair which had torn one another’s tongues out in their blind fury; and everybody knows that Robins and Tits fight savagely to preserve their chosen haunts from invasion by their neighbours. In some birds this pugnacity has become an overmastering passion. Some of the Quails, and a species of Rail (Gallicrex cristatus), a near relation of the Moorhen, are commonly kept by the natives of the East, as our forefathers kept Fighting-cocks, for the sake of seeing them fight one another. Yet, save in the case of the Fighting-cock, neither of these birds possesses any aggressive weapons.

Among the game-birds, however, powerful armature, in the shape of long, pointed, spurs on the legs are met with. In the Jungle-fowls and Pheasants only a single pair are found on each leg, but in other species, as in the Francolins, there are several pairs, and these birds, it is instructive to notice, are notorious for the ferocity of their encounters. It is said that in the Indian Swamp-Francolin (Francolinus gularis) nearly every individual is marked by scars and wounds received in duels with rivals.

Certain members of the Plover-tribe, and certain Anserine birds, have developed spurs of a very formidable character on the wings. Among the Plover-tribe the best example of such armoured species is the Egyptian Spur-winged Plover (Hoplopterus). This bird, after the fashion of its unarmoured relatives, such as the Common Lapwing, fights by turning suddenly in the air and striking with the wings. In the case of the formidably armed Egyptian bird the result is often fatal; but with our Lapwing a fatal result is rare, since but slightly swollen knobs take the place of spurs. In Hoplopterus and in the Jacana this spur arises from the base of the thumb, but in the Spur-winged Goose (Plectropterus) it is borne by one of the wrist bones (the radial) while in the aberrant Geese-like birds (Palamedea and Chauna) there are two spurs on each wring, one at each end of the metacarpus. That these weapons have come into being in response to need seems a very natural conclusion, but it is one which presents many difficulties when more closely examined. The wing spurs, differing widely in their nature as they do, in one case borne on a carpal bone, in others on the metacarpus, seem rather to owe their origin to fortuitous variations which have become, so to speak, adopted by selection, than to a response to the oft-repeated stimuli incidental to fighting. The latter explanation is Lamarckian and to-day finds favour with but few. The stimulus theory seems to be effectually discounted by the existence of the spurs on the legs of gallinaceous birds. That these owe their origin to impacts, or blows, seems more than doubtful: and one can hardly see how they could have served any useful purpose until they had attained a sufficient length to serve as weapons. Even if we suppose that the spurs of, say, the Jungle-fowl or the Francolin have been derived from tuberosities such as are found on the legs of the French Partridge (Caccabis rufa), we should still lack evidence that the use of the legs in fighting caused the origin of the tuberosities.