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Descent of Man and Selection in Relation to Sex, Part 2

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. <h3>GRADATION OF SECONDARY SEXUAL CHARACTERS.</h3> [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 a highly finished aspect. But I have elsewhere described (50. ‘Variation of Animals and Plants under Domestication,’ vol. i. p. 254.) an exactly analogous variation in the hackles of a sub-variety of the game-cock, in which the tips, having a metallic lustre, “are separated from the lower part of the feather by a symmetrically shaped transparent zone, composed of the naked portions of the barbs.” The lower margin or base of the dark-blue centre of the ocellus is deeply indented on the line of the shaft. The surrounding zones likewise shew traces, as may be seen in the drawing (Fig. 54), of indentations, or rather breaks. These indentations are common to the Indian and Javan peacocks (Pavo cristatus and P. muticus); and they seem to deserve particular attention, as probably connected with the development of the ocellus; but for a long time I could not conjecture their meaning. If we admit the principle of gradual evolution, there must formerly have existed many species which presented every successive step between the wonderfully elongated tail-coverts of the peacock and the short tail-coverts of all ordinary birds; and again between the magnificent ocelli of the former, and the simpler ocelli or mere coloured spots on other birds; and so with all the other characters of the peacock. Let us look to the allied Gallinaceae for any still-existing gradations. The species and sub-species of Polyplectron inhabit countries adjacent to the native land of the peacock; and they so far resemble this bird that they are sometimes called peacock-pheasants. I am also informed by Mr. Bartlett that they resemble the peacock in their voice and in some of their habits. During the spring the males, as previously described, strut about before the comparatively plain-coloured females, expanding and erecting their tail and wing-feathers, which are ornamented with numerous ocelli. I request the reader to turn back to the drawing (Fig. 51) of a Polyplectron; In P. napoleonis the ocelli are confined to the tail, and the back is of a rich metallic blue; in which respects this species approaches the Java peacock. P. hardwickii possesses a peculiar top-knot, which is also somewhat like that of the Java peacock. In all the species the ocelli on the wings and tail are either circular or oval, and consist of a beautiful, iridescent, greenish-blue or greenish-purple disc, with a black border. This border in P. chinquis shades into brown, edged with cream colour, so that the ocellus is here surrounded with variously shaded, though not bright, concentric zones. The unusual length of the tail-coverts is another remarkable character in Polyplectron; for in some of the species they are half, and in others two-thirds as long as the true tail-feathers. The tail-coverts are ocellated as in the peacock. Thus the several species of Polyplectron manifestly make a graduated approach to the peacock in the length of their tail-coverts, in the zoning of the ocelli, and in some other characters. [Fig. 55. Part of a tail-covert of Polyplectron chinquis, with the two ocelli of natural size. Fig. 56. Part of a tail-covert of Polyplectron malaccense, with the two ocelli, partially confluent, of natural size.] Notwithstanding this approach, the first species of Polyplectron which I examined almost made me give up the search; for I found not only that the true tail-feathers, which in the peacock are quite plain, were ornamented with ocelli, but that the ocelli on all the feathers differed fundamentally from those of the peacock, in there being two on the same feather (Fig. 55), one on each side of the shaft. Hence I concluded that the early progenitors of the peacock could not have resembled a Polyplectron. But on continuing my search, I observed that in some of the species the two ocelli stood very near each other; that in the tail-feathers of P. hardwickii they touched each other; and, finally, that on the tail-coverts of this same species as well as of P. malaccense (Fig. 56) they were actually confluent. As the central part alone is confluent, an indentation is left at both the upper and lower ends; and the surrounding coloured zones are likewise indented. A single ocellus is thus formed on each tail-covert, though still plainly betraying its double origin. These confluent ocelli differ from the single ocelli of the peacock in having an indentation at both ends, instead of only at the lower or basal end. The explanation, however, of this difference is not difficult; in some species of Polyplectron the two oval ocelli on the same feather stand parallel to each other; in other species (as in P. chinquis) they converge towards one end; now the partial confluence of two convergent ocelli would manifestly leave a much deeper indentation at the divergent than at the convergent end. It is also manifest that if the convergence were strongly pronounced and the confluence complete, the indentation at the convergent end would tend to disappear. The tail-feathers in both species of the peacock are entirely destitute of ocelli, and this apparently is related to their being covered up and concealed by the long tail-coverts. In this respect they differ remarkably from the tail-feathers of Polyplectron, which in most of the species are ornamented with larger ocelli than those on the tail-coverts. Hence I was led carefully to examine the tail-feathers of the several species, in order to discover whether their ocelli shewed any tendency to disappear; and to my great satisfaction, this appeared to be so. The central tail-feathers of P. napoleonis have the two ocelli on each side of the shaft perfectly developed; but the inner ocellus becomes less and less conspicuous on the more exterior tail-feathers, until a mere shadow or rudiment is left on the inner side of the outermost feather. Again, in P. malaccense, the ocelli on the tail-coverts are, as we have seen, confluent; and these feathers are of unusual length, being two-thirds of the length of the tail-feathers, so that in both these respects they approach the tail-coverts of the peacock. Now in P. malaccense, the two central tail-feathers alone are ornamented, each with two brightly-coloured ocelli, the inner ocellus having completely disappeared from all the other tail-feathers. Consequently the tail-coverts and tail-feathers of this species of Polyplectron make a near approach in structure and ornamentation to the corresponding feathers of the peacock. As far, then, as gradation throws light on the steps by which the magnificent train of the peacock has been acquired, hardly anything more is needed. If we picture to ourselves a progenitor of the peacock in an almost exactly intermediate condition between the existing peacock, with his enormously elongated tail-coverts, ornamented with single ocelli, and an ordinary gallinaceous bird with short tail-coverts, merely spotted with some colour, we shall see a bird allied to Polyplectron—that is, with tail-coverts, capable of erection and expansion, ornamented with two partially confluent ocelli, and long enough almost to conceal the tail-feathers, the latter having already partially lost their ocelli. The indentation of the central disc and of the surrounding zones of the ocellus, in both species of peacock, speaks plainly in favour of this view, and is otherwise inexplicable. The males of Polyplectron are no doubt beautiful birds, but their beauty, when viewed from a little distance, cannot be compared with that of the peacock. Many female progenitors of the peacock must, during a long line of descent, have appreciated this superiority; for they have unconsciously, by the continued preference for the most beautiful males, rendered the peacock the most splendid of living birds. <h3>ARGUS PHEASANT.</h3> Another excellent case for investigation is offered by the ocelli on the wing-feathers of the Argus pheasant, which are shaded in so wonderful a manner as to resemble balls lying loose within sockets, and consequently differ from ordinary ocelli. No one, I presume, will attribute the shading, which has excited the admiration of many experienced artists, to chance—to the fortuitous concourse of atoms of colouring matter. That these ornaments should have been formed through the selection of many successive variations, not one of which was originally intended to produce the ball-and-socket effect, seems as incredible as that one of Raphael’s Madonnas should have been formed by the selection of chance daubs of paint made by a long succession of young artists, not one of whom intended at first to draw the human figure. In order to discover how the ocelli have been developed, we cannot look to a long line of progenitors, nor to many closely-allied forms, for such do not now exist. But fortunately the several feathers on the wing suffice to give us a clue to the problem, and they prove to demonstration that a gradation is at least possible from a mere spot to a finished ball-and-socket ocellus. [Fig. 57. Part of secondary wing-feather of Argus pheasant, shewing two perfect ocelli, a and b. A, B, C, D, etc., are dark stripes running obliquely down, each to an ocellus. [Much of the web on both sides, especially to the left of the shaft, has been cut off.] Fig.59. Portion of one of the secondary wing-feathers near to the body, shewing the so-called elliptic ornaments. The right-hand figure is given merely as a diagram for the sake of the letters of reference. A, B, C, D, etc. Rows of spots running down to and forming the elliptic ornaments. b. Lowest spot or mark in row B. c. The next succeeding spot or mark in the same row. d. Apparently a broken prolongation of the spot c. in the same row B.] The wing-feathers, bearing the ocelli, are covered with dark stripes (Fig. 57) or with rows of dark spots (Fig. 59), each stripe or row of spots running obliquely down the outer side of the shaft to one of the ocelli. The spots are generally elongated in a line transverse to the row in which they stand. They often become confluent either in the line of the row—and then they form a longitudinal stripe—or transversely, that is, with the spots in the adjoining rows, and then they form transverse stripes. A spot sometimes breaks up into smaller spots, which still stand in their proper places. It will be convenient first to describe a perfect ball-and-socket ocellus. This consists of an intensely black circular ring, surrounding a space shaded so as exactly to resemble a ball. The figure here given has been admirably drawn by Mr. Ford and well engraved, but a woodcut cannot exhibit the exquisite shading of the original. The ring is almost always slightly broken or interrupted (Fig. 57) at a point in the upper half, a little to the right of and above the white shade on the enclosed ball; it is also sometimes broken towards the base on the right hand. These little breaks have an important meaning. The ring is always much thickened, with the edges ill-defined towards the left-hand upper corner, the feather being held erect, in the position in which it is here drawn. Beneath this thickened part there is on the surface of the ball an oblique almost pure-white mark, which shades off downwards into a pale-leaden hue, and this into yellowish and brown tints, which insensibly become darker and darker towards the lower part of the ball. It is this shading which gives so admirably the effect of light shining on a convex surface. If one of the balls be examined, it will be seen that the lower part is of a brown tint and is indistinctly separated by a curved oblique line from the upper part, which is yellower and more leaden; this curved oblique line runs at right angles to the longer axis of the white patch of light, and indeed of all the shading; but this difference in colour, which cannot of course be shewn in the woodcut, does not in the least interfere with the perfect shading of the ball. It should be particularly observed that each ocellus stands in obvious connection either with a dark stripe, or with a longitudinal row of dark spots, for both occur indifferently 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 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.  <div style="break-after:column;"></div>

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