the females prefer certain males on account of the beauty of their plumage." Mr. Hewitt was convinced "that the female almost invariably prefers the most vigorous, defiant, and mettlesome male;" and Mr. Tegetmeier, "that a gamecock, though disfigured by being dubbed, and with his hackles trimmed, would be accepted as readily as a male retaining all his natural ornaments." "1 Evidence is adduced that a female pigeon will sometimes take an antipathy to a particular male without any assignable cause; or, in other cases, will take a strong fancy to some one bird, and will desert her own mate for him; but it is not stated that superiority or inferiority of plumage has anything to do with these fancies. Two instances are indeed given, of male birds being rejected, which had lost their ornamental plumage; but in both cases (a widow-finch and a silver pheasant) the long tail-plumes are the indication of sexual maturity. Such cases do not support the idea that males with the tail-feathers a trifle longer, or the colours a trifle brighter, are generally preferred, and that those which are only a little inferior are as generally rejected, and this is what is absolutely needed to establish the theory of the development of these plumes by means of the choice of the female.

It will be seen, that female birds have unaccountable likes and dislikes in the matter of their partners, just as we have ourselves, and this may afford us an illustration. A young man, when courting, brushes or curls his hair, and has his moustache, beard, or whiskers in perfect order, and no doubt his sweetheart admires them; but this does not prove that she marries him on account of these ornaments, still less that hair, beard, whiskers, and moustache were developed by the continued preferences of the female sex. So, a girl likes to see her lover well and fashionably dressed, and he always dresses as well as he can when he visits her; but we cannot conclude from this that the whole series of male costumes, from the brilliantly coloured, puffed, and slashed doublet and hose of the Elizabethan period, through the gorgeous coats, long waistcoats, and pigtails of the early Georgian era, down to the funereal dress-suit of the present day, are the direct result of female preference. In like manner, female birds may be 1 Descent of Man, pp. 417, 418, 420.

charmed or excited by the fine display of plumage by the males; but there is no proof whatever that slight differences in that display have any effect in determining their choice of a partner.

Display of Decorative Plumage.

The extraordinary manner in which most birds display their plumage at the time of courtship, apparently with the full knowledge that it is beautiful, constitutes one of Mr. Darwin's strongest arguments. It is, no doubt, a very curious and interesting phenomenon, and indicates a connection between the exertion of particular muscles and the development of colour and ornament; but, for the reasons just given, it does not prove that the ornament has been developed by female choice. During excitement, and when the organism develops superabundant energy, many animals find it pleasurable to exercise their various muscles, often in fantastic ways, as seen in the gambols of kittens, lambs, and other young animals. But at the time of pairing, male birds are in a state of the most perfect development, and possess an enormous store of vitality; and under the excitement of the sexual passion they perform strange antics or rapid flights, as much probably from an internal impulse to motion and exertion as with any desire to please their mates. Such are the rapid descent of the snipe, the soaring and singing of the lark, and the dances of the cock-of-the-rock and of many other birds.

It is very suggestive that similar strange movements are performed by many birds which have no special developments of plumage. Goatsuckers, geese, carrion-vultures, and many other birds of plain plumage have been observed to dance, spread their wings or tails, and perform strange love-antics. The courtship of the great albatross, a most unwieldy and plain coloured bird, has been thus described by Professor Moseley: "The male, standing by the female on the nest, raises his wings, spreads his tail and elevates it, throws up his head with the bill in the air, or stretches it straight out, or forwards, as far as he can, and then utters a curious cry."1 Mr. Jenner Weir informs me that "the male blackbird is full of action, spreads out his glossy wing and tail, turns his rich golden

1 Notes of a Naturalist on the Challenger.

beak towards the female, and chuckles with delight," while he has never seen the more plain coloured thrush demonstrative to the female. The linnet distends his rosy breast, and slightly expands his brown wings and tail; while the various gay coloured Australian finches adopt such attitudes and postures as, in every case, to show off their variously coloured plumage to the best advantage.1

A Theory of Animal Coloration.

Having rejected Mr. Darwin's theory of female choice as incompetent to account for the brilliant colours and markings of the higher animals, the preponderance of these colours and markings in the male sex, and their display during periods of activity or excitement, I may be asked what explanation I have to offer as a preferable substitute. In my Tropical Nature I have already indicated such a theory, which I will now briefly explain, supporting it by some additional facts and arguments, which appear to me to have great weight, and for which I am mainly indebted to a most interesting and suggestive posthumous work by Mr. Alfred Tylor.2

The fundamental or ground colours of animals are, as has been shown in preceding chapters, very largely protective, and it is not improbable that the primitive colours of all animals were so. During the long course of animal development other modes of protection than concealment by harmony of colour arose, and thenceforth the normal development of colour due to the complex chemical and structural changes ever going on in the organism, had full play; and the colours thus produced were again and again modified by natural selection for purposes of warning, recognition, mimicry, or special protection, as has been already fully explained in the preceding chapters.

Mr. Tylor has, however, called attention to an important principle which underlies the various patterns or ornamental markings of animals—namely, that diversified coloration follows the chief lines of structure, and changes at points, such as the joints, where function changes. He says, "If we take highly decorated species-that is, animals marked by 1 Descent of Man, pp. 401, 402.

2 Coloration in Animals and Plants, London, 1886.

alternate dark or light bands or spots, such as the zebra, some deer, or the carnivora, we find, first, that the region of the spinal column is marked by a dark stripe; secondly, that the regions of the appendages, or limbs, are differently marked ; thirdly, that the flanks are striped or spotted, along or between the regions of the lines of the ribs; fourthly, that the shoulder and hip regions are marked by curved lines; fifthly, that the pattern changes, and the direction of the lines, or spots, at the head, neck, and every joint of the limbs; and lastly, that the tips of the ears, nose, tail, and feet, and the eye are emphasised in colour. In spotted animals the greatest length of the spot is generally in the direction of the largest development of the skeleton."

This structural decoration is well seen in many insects. In caterpillars, similar spots and markings are repeated in each segment, except where modified for some form of protection. In butterflies, the spots and bands usually have reference to the form of the wing and the arrangement of the nervures; and there is much evidence to show that the primitive markings are always spots in the cells, or between the nervures, or at the junctions of nervures, the extension and coalescence of these spots forming borders, bands, or blotches, which have become modified in infinitely varied ways for protection, warning, or recognition. Even in birds, the distribution of colours and markings follows generally the same law. The crown of the head, the throat, the ear-coverts, and the eyes have usually distinct tints in all highly coloured birds; the region of the furcula has often a distinct patch of colour, as have the pectoral muscles, the uropygium or root of the tail, and the under tail-coverts.1

Mr. Tylor was of opinion that the primitive form of ornamentation consisted of spots, the confluence of these in certain directions forming lines or bands; and, these again, sometimes coalescing into blotches, or into more or less uniform tints covering a large portion of the surface of the body. The young lion and tiger are both spotted; and in the Java hog (Sus vittatus) very young animals are banded, but have spots over the shoulders and thighs. These spots run into stripes 1 Coloration of Animals, Pl. X, p. 90; and Pls. II, III, and IV, pp. 30, 40, 42.

as the animal grows older; then the stripes expand, and at last, meeting together, the adult animal becomes of a uniform dark brown colour. So many of the species of deer are spotted when young, that Darwin concludes the ancestral form, from which all deer are derived, must have been spotted. Pigs and tapirs are banded or spotted when young; an imported young specimen of Tapirus Bairdi was covered with white spots in longitudinal rows, here and there forming short stripes. Even the horse, which Darwin supposes to be descended from a striped animal, is often spotted, as in dappled horses; and great numbers show a tendency to spottiness, especially on the haunches.

Ocelli may also be developed from spots, or from bars, as pointed out by Mr. Darwin. Spots are an ordinary form of marking in disease, and these spots sometimes run together, forming blotches. There is evidence that colour markings are in some way dependent on nerve distribution. In the disease known as frontal herpes, an eruption occurs which corresponds exactly to the distribution of the ophthalmic division of the fifth cranial nerve, mapping out all its little branches even to the one which goes to the tip of the nose. In a Hindoo suffering from herpes the pigment was destroyed in the arm along the course of the ulnar nerve, with its branches along both sides of one finger and the half of another. In the leg the sciatic and scaphenous nerves were partly mapped out, giving to the patient the appearance of an anatomical diagram.2

These facts are very interesting, because they help to explain the general dependence of marking on structure which has been already pointed out. For, as the nerves everywhere follow the muscles, and these are attached to the various bones, we see how it happens, that the tracts in which distinct developments of colour appear, should so often be marked out by the chief divisions of the bony structure in vertebrates, and by the segments in the annulosa. There is, however, another correspondence of even greater interest and importance. Brilliant colours usually appear just in proportion to the

1 See coloured Fig. in Proc. Zool. Soc., 1871, p. 626.

2 A. Tylor's Coloration, p. 40; and Photograph in Hutchinson's Illustrations of Clinical Surgery, quoted by Tylor. His interpretation of the facts may, however, be erroneous.