The Descent of Man – Day 106 of 151

Formation and Variability of the Ocelli or Eye-like Spots on the Plumage of Birds

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.

Figure 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.

Gradation of Secondary Sexual Characters

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.

Figure 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 bas 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.

Part of a tail-covert of Polyplectron chinquis, with the two ocelli of natural size.

Figure 55: Part of a tail-covert of Polyplectron chinquis, with the two ocelli of natural size.

Part of a tail-covert of Polyplectron malaccense, with the two ocelli, partially confluent, of natural size.

Figure 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.

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