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.[129]
The fundamental or ground colours of animals ar 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. Taylor 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 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.[130]
Mr. Tylor was of opinion 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 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.[131] 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.[132]
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 development of tegumentary appendages. Among birds the most brilliant colours are possessed by those which have developed frills, crests, and elongated tails like the humming-birds; immense tail-coverts like the peacock; enormously expanded wing-feathers, as in the argus-pheasant; or magnificent plumes from the region of the coracoids in many of the birds of paradise. It is to be noted, also, that all these accessory plumes spring from parts of the body which, in other species, are distinguished by patches of colour; so that we may probably impute the development of colour and of accessory plumage to the same fundamental cause.
Among insects, the most brilliant and varied coloration occurs in the butterflies and moths, groups in which the wing-membranes have received their greatest expansion, and whose specialisation has been carried furthest in the marvellous scaly covering which is the seat of the colour. It is suggestive, that the only other group in which functional wings are much coloured is that of the dragonflies, where the membrane is exceedingly expanded. In like manner, the colours of beetles, though greatly inferior to those of the lepidoptera, occur in a group in which the anterior pair of wings has been thickened and modified in order to protect the vital parts, and in which these wing-covers (elytra), in the course of development in the different groups, must have undergone great changes, and have been the seat of very active growth.
The Origin of Accessory Plumes.
Mr. Darwin supposes, that these have in almost every case been developed by the preference of female birds for such males as possessed them in a higher degree than others; but this theory does not account for the fact that these plumes usually appear in a few definite parts of the body. We require some cause to initiate the development in one part rather than in another. Now, the view that colour has arisen over surfaces where muscular and nervous development is considerable, and the fact that it appears especially upon the accessory or highly developed plumes, leads us to inquire whether the same cause has not primarily determined the development of these plumes. The immense tuft of golden plumage in the best known birds of paradise (Paradisea apoda and P. minor) springs from a very small area on the side of the breast. Mr. Frank E. Beddard, who has kindly examined a specimen for me, says that "this area lies upon the pectoral muscles, and near to the point where the fibres of the muscle converge towards their attachment to the humerus. The plumes arise, therefore, close to the most powerful muscle of the body, and near to where the activities of that muscle would be at a maximum. Furthermore, the area of attachment of the plumes is just above the point where the arteries and nerves for the supply of the pectoral muscles, and neighbouring regions, leave the interior of the body. The area of attachment of the plume is, also, as you say in your letter, just above the junction of the coracoid and sternum." Ornamental plumes of considerable size rise from the same part in many other species of paradise birds, sometimes extending laterally in front, so as to form breast shields. They also occur in many humming-birds, and in some sun-birds and honey-suckers; and in all these cases there is a wonderful amount of activity and rapid movement, indicating a surplus of vitality, which is able to manifest itself in the development of these accessory plumes.[133]