The Descent of Man – Day 108 of 151

An ocellus in an intermediate condition between the elliptic ornament and the perfect ball-and-socket ocellus.

Figure 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.

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.

Figure 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 actual steps, and they at least prove to demonstration that a gradation is possible. Bearing in mind how carefully the male Argus pheasant displays his plumes before the female, as well as the many facts rendering it probable that female birds prefer the more attractive males, no one who admits the agency of sexual selection in any case will deny that a simple dark spot with some fulvous shading might be converted, through the approximation and modification of two adjoining spots, together with some slight increase of colour, into one of the so-called elliptic ornaments. These latter ornaments have been shewn to many persons, and all have admitted that they are beautiful, some thinking them even more so than the ball-and-socket ocelli. As the secondary plumes became lengthened through sexual selection, and as the elliptic ornaments increased in diameter, their colours apparently became less bright; and then the ornamentation of the plumes had to be gained by an improvement in the pattern and shading; and this process was carried on until the wonderful ball-and-socket ocelli were finally developed. Thus we can understand–and in no other way as it seems to me–the present condition and origin of the ornaments on the wing-feathers of the Argus pheasant.

From the light afforded by the principle of gradation–from what we know of the laws of variation–from the changes which have taken place in many of our domesticated birds–and, lastly, from the character (as we shall hereafter see more clearly) of the immature plumage of young birds–we can sometimes indicate, with a certain amount of confidence, the probable steps by which the males have acquired their brilliant plumage and various ornaments; yet in many cases we are involved in complete darkness. Mr. Gould several years ago pointed out to me a humming-bird, the Urosticte benjamini, remarkable for the curious differences between the sexes. The male, besides a splendid gorget, has greenish-black tail-feathers, with the four central ones tipped with white; in the female, as with most of the allied species, the three outer tail-feathers on each side are tipped with white, so that the male has the four central, whilst the female has the six exterior feathers ornamented with white tips. What makes the case more curious is that, although the colouring of the tail differs remarkably in both sexes of many kinds of humming-birds, Mr. Gould does not know a single species, besides the Urosticte, in which the male has the four central feathers tipped with white.

The Duke of Argyll, in commenting on this case (53. ‘The Reign of Law,’ 1867, p. 247.), passes over sexual selection, and asks, “What explanation does the law of natural selection give of such specific varieties as these?” He answers “none whatever”; and I quite agree with him. But can this be so confidently said of sexual selection? Seeing in how many ways the tail-feathers of humming-birds differ, why should not the four central feathers have varied in this one species alone, so as to have acquired white tips? The variations may have been gradual, or somewhat abrupt as in the case recently given of the humming-birds near Bogota, in which certain individuals alone have the “central tail-feathers tipped with beautiful green.” In the female of the Urosticte I noticed extremely minute or rudimental white tips to the two outer of the four central black tail-feathers; so that here we have an indication of change of some kind in the plumage of this species. If we grant the possibility of the central tail-feathers of the male varying in whiteness, there is nothing strange in such variations having been sexually selected. The white tips, together with the small white ear-tufts, certainly add, as the Duke of Argyll admits, to the beauty of the male; and whiteness is apparently appreciated by other birds, as may be inferred from such cases as the snow-white male of the Bell-bird. The statement made by Sir R. Heron should not be forgotten, namely, that his peahens, when debarred from access to the pied peacock, would not unite with any other male, and during that season produced no offspring. Nor is it strange that variations in the tail-feathers of the Urosticte should have been specially selected for the sake of ornament, for the next succeeding genus in the family takes its name of Metallura from the splendour of these feathers. We have, moreover, good evidence that humming-birds take especial pains in displaying their tail-feathers; Mr. Belt (54. ‘The Naturalist in Nicaragua,’ 1874, p. 112.), after describing the beauty of the Florisuga mellivora, says, “I have seen the female sitting on a branch, and two males displaying their charms in front of her. One would shoot up like a rocket, then suddenly expanding the snow-white tail, like an inverted parachute, slowly descend in front of her, turning round gradually to shew off back and front…The expanded white tail covered more space than all the rest of the bird, and was evidently the grand feature in the performance. Whilst one male was descending, the other would shoot up and come slowly down expanded. The entertainment would end in a fight between the two performers; but whether the most beautiful or the most pugnacious was the accepted suitor, I know not.” Mr. Gould, after describing the peculiar plumage of the Urosticte, adds, “that ornament and variety is the sole object, I have myself but little doubt.” (55. ‘Introduction to the Trochilidae,’ 1861, p. 110.) If this be admitted, we can perceive that the males which during former times were decked in the most elegant and novel manner would have gained an advantage, not in the ordinary struggle for life, but in rivalry with other males, and would have left a larger number of offspring to inherit their newly-acquired beauty.

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