The Descent of Man, and Selection in Relation to Sex

colour of the eyes have been preserved or augmented through sexual selection in the males? This is very doubtful; for Mr. Bartlett shewed me in the Zoological Gardens that the inside of the mouth of this Buceros is black in the male and flesh-coloured in the female; and their external appearance or beauty would not be thus affected. I observed in Chile (46. ‘Zoology of the Voyage of H.M.S. “Beagle,”’ 1841, p. 6.) that the iris in the condor, when about a year old, is dark-brown, but changes at maturity into yellowish-brown in the male, and into bright red in the female. The male has also a small, longitudinal, leaden-coloured, fleshy crest or comb. The comb of many gallinaceous birds is highly ornamental, and assumes vivid colours during the act of courtship; but what are we to think of the dull-coloured comb of the condor, which does not appear to us in the least ornamental? The same question may be asked in regard to various other characters, such as the knob on the base of the beak of the Chinese goose (Anser cygnoides), which is much larger in the male than in the female. No certain answer can be given to these questions; but we ought to be cautious in assuming that knobs and various fleshy appendages cannot be attractive to the female, when we remember that with savage races of man various hideous deformities—deep scars on the face with the flesh raised into protuberances, the septum of the nose pierced by sticks or bones, holes in the ears and lips stretched widely open—are all admired as ornamental. Whether or not unimportant differences between the sexes, such as those just specified, have been preserved through sexual selection, these differences, as well as all others, must primarily depend on the laws of variation. On the principle of correlated development, the plumage often varies on different parts of the body, or over the whole body, in the same manner. We see this well illustrated in certain breeds of the fowl. In all the breeds the feathers on the neck and loins of the males are elongated, and are called hackles; now when both sexes acquire a top-knot, which is a new character in the genus, the feathers on the head of the male become hackle-shaped, evidently on the principle of correlation; whilst those on the head of the female are of the ordinary shape. The colour also of the hackles forming the top-knot of the male, is often correlated with that of the hackles on the neck and loins, as may be seen by comparing these feathers in the golden and silver-spangled Polish, the Houdans, and Creve-coeur breeds. In some natural species we may observe exactly the same correlation in the colours of these same feathers, as in the males of the splendid Gold and Amherst pheasants. The structure of each individual feather generally causes any change in its colouring to be symmetrical; we see this in the various laced, spangled, and pencilled breeds of the fowl; and on the principle of correlation the feathers over the whole body are often coloured in the same manner. We are thus enabled without much trouble to rear breeds with their plumage marked almost as symmetrically as in natural species. In laced and spangled fowls the coloured margins of the feathers are abruptly defined; but in a mongrel raised by me from a black Spanish cock glossed with green, and a white game-hen, all the feathers were greenish-black, excepting towards their extremities, which were yellowish-white; but between the white extremities and the black bases, there was on each feather a symmetrical, curved zone of dark-brown. In some instances the shaft of the feather determines the distribution of the tints; thus with the body-feathers of a mongrel from the same black Spanish cock and a silver-spangled Polish hen, the shaft, together with a narrow space on each side, was greenish-black, and this was surrounded by a regular zone of dark-brown, edged with brownish-white. In these cases we have feathers symmetrically shaded, like those which give so much elegance to the plumage of many natural species. I have also noticed a variety of the common pigeon with the wing-bars symmetrically zoned with three bright shades, instead of being simply black on a slaty-blue ground, as in the parent-species. In many groups of birds the plumage is differently coloured in the several species, yet certain spots, marks, or stripes are retained by all. Analogous cases occur with the breeds of the pigeon, which usually retain the two wing-bars, though they may be coloured red, yellow, white, black, or blue, the rest of the plumage being of some wholly different tint. Here is a more curious case, in which certain marks are retained, though coloured in a manner almost exactly the opposite of what is natural; the aboriginal pigeon has a blue tail, with the terminal halves of the outer webs of the two outer tail feathers white; now there is a sub-variety having a white instead of a blue tail, with precisely that part black which is white in the parent-species. (47. Bechstein, ‘Naturgeschichte Deutschlands,’ B. iv. 1795, s. 31, on a sub-variety of the Monck pigeon.) FORMATION AND VARIABILITY OF THE OCELLI OR EYE-LIKE SPOTS ON THE PLUMAGE OF BIRDS. [Fig. 53. Cyllo leda, Linn., from a drawing by Mr. Trimen, shewing the extreme range of variation in the ocelli. A. Specimen, from Mauritius, upper surface of fore-wing. A1. Specimen, from Natal, ditto. B. Specimen, from Java, upper surface of hind-wing. B1. Specimen, from Mauritius, ditto.] As no ornaments are more beautiful than the ocelli on the feathers of various birds, on the hairy coats of some mammals, on the scales of reptiles and fishes, on the skin of amphibians, on the wings of many Lepidoptera and other insects, they deserve to be especially noticed. An ocellus consists of a spot within a ring of another colour, like the pupil within the iris, but the central spot is often surrounded by additional concentric zones. The ocelli on the tail-coverts of the peacock offer a familiar example, as well as those on the wings of the peacock-butterfly (Vanessa). Mr. Trimen has given me a description of a S. African moth (Gynanisa isis), allied to our Emperor moth, in which a magnificent ocellus occupies nearly the whole surface of each hinder wing; it consists of a black centre, including a semi-transparent crescent-shaped mark, surrounded by successive, ochre-yellow, black, ochre-yellow, pink, white, pink, brown, and whitish zones. Although we do not know the steps by which these wonderfully beautiful and complex ornaments have been developed, the process has probably been a simple one, at least with insects; for, as Mr. Trimen writes to me, “no characters of mere marking or coloration are so unstable in the Lepidoptera as the ocelli, both in number and size.” Mr. Wallace, who first called my attention to this subject, shewed me a series of specimens of our common meadow-brown butterfly (Hipparchia janira) exhibiting numerous gradations from a simple minute black spot to an elegantly-shaded ocellus. In a S. African butterfly (Cyllo leda, Linn.), belonging to the same family, the ocelli are even still more variable. In some specimens (A, Fig. 53) large spaces on the upper surface of the wings are coloured black, and include irregular white marks; and from this state a complete gradation can be traced into a tolerably perfect ocellus (A1), and this results from the contraction of the irregular blotches of colour. In another series of specimens a gradation can be followed from excessively minute white dots, surrounded by a scarcely visible black line (B), into perfectly symmetrical and large ocelli (B1). (48. This woodcut has been engraved from a beautiful drawing, most kindly made for me by Mr. Trimen; see also his description of the wonderful amount of variation in the coloration and shape of the wings of this butterfly, in his ‘Rhopalocera Africae Australis,’ p. 186.) In cases like these, the development of a perfect ocellus does not require a long course of variation and selection. With birds and many other animals, it seems to follow from the comparison of allied species that circular spots are often generated by the breaking up and contraction of stripes. In the Tragopan pheasant faint white lines in the female represent the beautiful white spots in the male (49. Jerdon, ‘Birds of India,’ vol. iii. p. 517.); and something of the same kind may be observed in the two sexes of the Argus pheasant. However this may be, appearances strongly favour the belief that on the one hand, a dark spot is often formed by the colouring matter being drawn towards a central point from a surrounding zone, which latter is thus rendered lighter; and, on the other hand, that a white spot is often formed by the colour being driven away from a central point, so that it accumulates in a surrounding darker zone. In either case an ocellus is the result. The colouring matter seems to be a nearly constant quantity, but is redistributed, either centripetally or centrifugally. The feathers of the common guinea-fowl offer a good instance of white spots surrounded by darker zones; and wherever the white spots are large and stand near each other, the surrounding dark zones become confluent. In the same wing-feather of the Argus pheasant dark spots may be seen surrounded by a pale zone, and white spots by a dark zone. Thus the formation of an ocellus in its most elementary state appears to be a simple affair. By what further steps the more complex ocelli, which are surrounded by many successive zones of colour, have been generated, I will not pretend to say. But the zoned feathers of the mongrels from differently coloured fowls, and the extraordinary variability of the ocelli on many Lepidoptera, lead us to conclude that their formation is not a complex process, but depends on some slight and graduated change in the nature of the adjoining tissues. [Fig. 54. Feather of Peacock, about two-thirds of natural size, drawn by Mr. Ford. The transparent zone is represented by the outermost white zone, confined to the upper end of the disc.] Cases of gradation are important, as shewing us that highly complex ornaments may be acquired by small successive steps. In order to discover the actual steps by which the male of any existing bird has acquired his magnificent colours or other ornaments, we ought to behold the long line of his extinct progenitors; but this is obviously impossible. We may, however, generally gain a clue by comparing all the species of the same group, if it be a large one; for some of them will probably retain, at least partially, traces of their former characters. Instead of entering on tedious details respecting various groups, in which striking instances of gradation could be given, it seems the best plan to take one or two strongly marked cases, for instance that of the peacock, in order to see if light can be thrown on the steps by which this bird has become so splendidly decorated. The peacock is chiefly remarkable from the extraordinary length of his tail-coverts; the tail itself not being much elongated. The barbs along nearly the whole length of these feathers stand separate or are decomposed; but this is the case with the feathers of many species, and with some varieties of the domestic fowl and pigeon. The barbs coalesce towards the extremity of the shaft forming the oval disc or ocellus, which is certainly one of the most beautiful objects in the world. It consists of an iridescent, intensely blue, indented centre, surrounded by a rich green zone, this by a broad coppery-brown zone, and this by five other narrow zones of slightly different iridescent shades. A trifling character in the disc deserves notice; the barbs, for a space along one of the concentric zones are more or less destitute of their barbules, so that a part of the disc is surrounded by an almost transparent zone, which gives it