The Descent of Man, and Selection in Relation to Sex

on the same feather. Thus in Fig. 57 stripe A runs to ocellus a; B runs to ocellus b; stripe C is broken in the upper part, and runs down to the next succeeding ocellus, not represented in the woodcut; D to the next lower one, and so with the stripes E and F. Lastly, the several ocelli are separated from each other by a pale surface bearing irregular black marks. [Fig. 58. Basal part of the secondary wing feather, nearest to the body.] I will next describe the other extreme of the series, namely, the first trace of an ocellus. The short secondary wing-feather (Fig. 58), nearest to the body, is marked like the other feathers, with oblique, longitudinal, rather irregular, rows of very dark spots. The basal spot, or that nearest the shaft, in the five lower rows (excluding the lowest one) is a little larger than the other spots of the same row, and a little more elongated in a transverse direction. It differs also from the other spots by being bordered on its upper side with some dull fulvous shading. But this spot is not in any way more remarkable than those on the plumage of many birds, and might easily be overlooked. The next higher spot does not differ at all from the upper ones in the same row. The larger basal spots occupy exactly the same relative position on these feathers as do the perfect ocelli on the longer wing-feathers. By looking to the next two or three succeeding wing-feathers, an absolutely insensible gradation can be traced from one of the last-described basal spots, together with the next higher one in the same row, to a curious ornament, which cannot be called an ocellus, and which I will name, from the want of a better term, an “elliptic ornament.” These are shewn in the accompanying figure (Fig. 59). We here see several oblique rows, A, B, C, D, etc. (see the lettered diagram on the right hand), of dark spots of the usual character. Each row of spots runs down to and is connected with one of the elliptic ornaments, in exactly the same manner as each stripe in Fig. 57 runs down to and is connected with one of the ball-and-socket ocelli. Looking to any one row, for instance, B, in Fig. 59, the lowest mark (b) is thicker and considerably longer than the upper spots, and has its left extremity pointed and curved upwards. This black mark is abruptly bordered on its upper side by a rather broad space of richly shaded tints, beginning with a narrow brown zone, which passes into orange, and this into a pale leaden tint, with the end towards the shaft much paler. These shaded tints together fill up the whole inner space of the elliptic ornament. The mark (b) corresponds in every respect with the basal shaded spot of the simple feather described in the last paragraph (Fig. 58), but is more highly developed and more brightly coloured. Above and to the right of this spot (b, Fig. 59), with its bright shading, there is a long narrow, black mark (c), belonging to the same row, and which is arched a little downwards so as to face (b). This mark is sometimes broken into two portions. It is also narrowly edged on the lower side with a fulvous tint. To the left of and above c, in the same oblique direction, but always more or less distinct from it, there is another black mark (d). This mark is generally sub-triangular and irregular in shape, but in the one lettered in the diagram it is unusually narrow, elongated, and regular. It apparently consists of a lateral and broken prolongation of the mark (c), together with its confluence with a broken and prolonged part of the next spot above; but I do not feel sure of this. These three marks, b, c, and d, with the intervening bright shades, form together the so-called elliptic ornament. These ornaments placed parallel to the shaft, manifestly correspond in position with the ball-and-socket ocelli. Their extremely elegant appearance cannot be appreciated in the drawing, as the orange and leaden tints, contrasting so well with the black marks, cannot be shewn. [Fig. 60. An ocellus in an intermediate condition between the elliptic ornament and the perfect ball-and-socket ocellus.] Between one of the elliptic ornaments and a perfect ball-and-socket ocellus, the gradation is so perfect that it is scarcely possible to decide when the latter term ought to be used. The passage from the one into the other is effected by the elongation and greater curvature in opposite directions of the lower black mark (b, Fig. 59), and more especially of the upper one (c), together with the contraction of the elongated sub-triangular or narrow mark (d), so that at last these three marks become confluent, forming an irregular elliptic ring. This ring is gradually rendered more and more circular and regular, increasing at the same time in diameter. I have here given a drawing (Fig. 60) of the natural size of an ocellus not as yet quite perfect. The lower part of the black ring is much more curved than is the lower mark in the elliptic ornament (b, Fig. 59). The upper part of the ring consists of two or three separate portions; and there is only a trace of the thickening of the portion which forms the black mark above the white shade. This white shade itself is not as yet much concentrated; and beneath it the surface is brighter coloured than in a perfect ball-and-socket ocellus. Even in the most perfect ocelli traces of the junction of three or four elongated black marks, by which the ring has been formed, may often be detected. The irregular sub-triangular or narrow mark (d, Fig. 59), manifestly forms, by its contraction and equalisation, the thickened portion of the ring above the white shade on a perfect ball-and-socket ocellus. The lower part of the ring is invariably a little thicker than the other parts (Fig. 57), and this follows from the lower black mark of the elliptic ornament (b, Fig. 59) having originally been thicker than the upper mark (c). Every step can be followed in the process of confluence and modification; and the black ring which surrounds the ball of the ocellus is unquestionably formed by the union and modification of the three black marks, b, c, d, of the elliptic ornament. The irregular zigzag black marks between the successive ocelli (Fig. 57) are plainly due to the breaking up of the somewhat more regular but similar marks between the elliptic ornaments. The successive steps in the shading of the ball-and-socket ocelli can be followed out with equal clearness. The brown, orange, and pale-leadened narrow zones, which border the lower black mark of the elliptic ornament, can be seen gradually to become more and more softened and shaded into each other, with the upper lighter part towards the left-hand corner rendered still lighter, so as to become almost white, and at the same time more contracted. But even in the most perfect ball-and-socket ocelli a slight difference in the tints, though not in the shading, between the upper and lower parts of the ball can be perceived, as before noticed; and the line of separation is oblique, in the same direction as the bright coloured shades of the elliptic ornaments. Thus almost every minute detail in the shape and colouring of the ball-and-socket ocelli can be shewn to follow from gradual changes in the elliptic ornaments; and the development of the latter can be traced by equally small steps from the union of two almost simple spots, the lower one (Fig. 58) having some dull fulvous shading on its upper side. [Fig. 61. Portion near summit of one of the secondary wing-feathers, bearing perfect ball-and-socket ocelli. a. Ornamented upper part. b. Uppermost, imperfect ball-and-socket ocellus. (The shading above the white mark on the summit of the ocellus is here a little too dark.) c. Perfect ocellus.] The extremities of the longer secondary feathers which bear the perfect ball-and-socket ocelli, are peculiarly ornamented (Fig. 61). The oblique longitudinal stripes suddenly cease upwards and become confused; and above this limit the whole upper end of the feather (a) is covered with white dots, surrounded by little black rings, standing on a dark ground. The oblique stripe belonging to the uppermost ocellus (b) is barely represented by a very short irregular black mark with the usual, curved, transverse base. As this stripe is thus abruptly cut off, we can perhaps understand from what has gone before, how it is that the upper thickened part of the ring is here absent; for, as before stated, this thickened part apparently stands in some relation with a broken prolongation from the next higher spot. From the absence of the upper and thickened part of the ring, the uppermost ocellus, though perfect in all other respects, appears as if its top had been obliquely sliced off. It would, I think, perplex any one, who believes that the plumage of the Argus pheasant was created as we now see it, to account for the imperfect condition of the uppermost ocellus. I should add that on the secondary wing-feather farthest from the body all the ocelli are smaller and less perfect than on the other feathers, and have the upper part of the ring deficient, as in the case just mentioned. The imperfection here seems to be connected with the fact that the spots on this feather shew less tendency than usual to become confluent into stripes; they are, on the contrary, often broken up into smaller spots, so that two or three rows run down to the same ocellus. There still remains another very curious point, first observed by Mr. T.W. Wood (51. The ‘Field,’ May 28, 1870.), which deserves attention. In a photograph, given me by Mr. Ward, of a specimen mounted as in the act of display, it may be seen that on the feathers which are held perpendicularly, the white marks on the ocelli, representing light reflected from a convex surface, are at the upper or further end, that is, are directed upwards; and the bird whilst displaying himself on the ground would naturally be illuminated from above. But here comes the curious point; the outer feathers are held almost horizontally, and their ocelli ought likewise to appear as if illuminated from above, and consequently the white marks ought to be placed on the upper sides of the ocelli; and, wonderful as is the fact, they are thus placed! Hence the ocelli on the several feathers, though occupying very different positions with respect to the light, all appear as if illuminated from above, just as an artist would have shaded them. Nevertheless they are not illuminated from strictly the same point as they ought to be; for the white marks on the ocelli of the feathers which are held almost horizontally, are placed rather too much towards the further end; that is, they are not sufficiently lateral. We have, however, no right to expect absolute perfection in a part rendered ornamental through sexual selection, any more than we have in a part modified through natural selection for real use; for instance, in that wondrous organ the human eye. And we know what Helmholtz, the highest authority in Europe on the subject, has said about the human eye; that if an optician had sold him an instrument so carelessly made, he would have thought himself fully justified in returning it. (52. ‘Popular Lectures on Scientific Subjects,’ Eng. trans. 1873, pp. 219, 227, 269, 390.) We have now seen that a perfect series can be followed, from simple spots to the wonderful ball-and-socket ornaments. Mr. Gould, who kindly gave me some of these feathers, fully agrees with me in the completeness of the gradation. It is obvious that the stages in development exhibited by the feathers on the same bird do not at all necessarily shew us the steps passed through by the extinct progenitors of the species; but they probably give us the clue to the