The Project Gutenberg eBook of The Descent of Man and Selection in Relation to Sex, by Charles Darwin

In many large groups of birds it may be observed that the plumage is differently coloured in each species, yet that certain spots, marks, or stripes, though likewise differently coloured, are retained by all the species. 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 almost an exactly reversed manner to 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 small part black which is white in the parent-species.191

Formation and variability of the Ocelli or eye-like Spots on the Plumage of Birds.—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 at least with insects has probably been a simple one; 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 belonging to the same family, the ocelli are even still more variable. In some specimens (A, fig. 52) 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 (A¹) ocellus, 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).192 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, from the comparison of allied species, to follow, 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;193 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 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 simplest 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 bearing in mind the zoned feathers of the mongrel offspring from differently-coloured fowls, and the extraordinary variability of the ocelli in many Lepidoptera, the formation of these beautiful ornaments can hardly be a highly complex process, and probably depends on some slight and graduated change in the nature of the tissues.

Gradation of Secondary Sexual Characters.—Cases of gradation are important for us, as they shew that it is at least possible that highly complex ornaments may have been 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 ancient and extinct progenitors; but this is obviously impossible. We may, however, generally gain a clue by comparing all the species of a group, if it be a large one; for some of them will probably retain, at least in a partial manner, 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 some one or two strongly-characterised cases, for instance that of the peacock, in order to discover if any light can thus 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 to form the oval disc or ocellus, which is certainly one of the most beautiful objects in the world. This consists of an iridescent, intensely blue, indented centre, surrounded by a rich green zone, and 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 perhaps deserves notice; the barbs, for a space along one of the concentric zones are destitute, to a greater or less degree, of their barbules, so that a part of the disc is surrounded by an almost transparent zone, which gives to it a highly-finished aspect. But I have elsewhere described 194 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. 53), of indentations, or rather breaks. These indentations are common to the Indian and Javan peacocks (Pavo cristatus and P. muticus); and they seemed to me 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 of other birds; and so with all the other characters of the peacock. Let us look to the allied Gallinaceæ 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, p. 90) 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, somewhat like that of this same kind of peacock. The ocelli on the wings and tail of the several species of Polyplectron 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 which is edged with cream-colour, so that the ocellus is here surrounded with differently, though not brightly, shaded concentric zones. The unusual length of the tail-coverts is another highly remarkable character in Polyplectron; for in some of the species they are half as long, and in others two-thirds of the length of the true tail-feathers. The tail-coverts are ocellated, as in the peacock. Thus the several species of Polyplectron manifestly make a graduated approach in the length of their tail-coverts, in the zoning of the ocelli, and in some other characters, to the peacock.

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 at the lower or basal end alone. 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 be quite obliterated.

The tail-feathers in both species of 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 of Polyplectron in order to discover whether the ocelli in any of them shewed any tendency to disappear, and, to my great satisfaction, I was successful. 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 rudimentary vestige 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 resemble the tail-coverts of the peacock. Now in this species the two central tail-feathers alone are ornamented, each with two brightly-coloured ocelli, the ocelli having completely disappeared from the inner sides of 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 the principle of gradation throws light on the steps by which the magnificent train of the peacock has been acquired, hardly anything more is needed. We may 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; and we shall then see in our mind’s eye, a bird possessing 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; we shall see in short, a Polyplectron. The indentation of the central disc and surrounding zones of the ocellus in both species of peacock, seems to me to speak plainly in favour of this view; and this structure is otherwise inexplicable. The males of Polyplectron are no doubt very beautiful birds, but their beauty, when viewed from a little distance, cannot be compared, as I formerly saw in the Zoological Gardens, 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 of the most beautiful males, rendered the peacock the most splendid of living birds.

Argus pheasant.—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 within sockets, and which 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 various 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.

The wing-feathers, bearing the ocelli, are covered with dark stripes or rows of dark spots, each stripe or row running obliquely down the outer side of the shaft to an ocellus. The spots are generally elongated in a transverse line 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.

I will next describe the other extreme of the series, namely the first trace of an ocellus. The short secondary wing-feather (fig. 57), nearest to the body, is marked like the other feathers, with oblique, longitudinal, rather irregular, rows of spots. The lowest spot, or that nearest the shaft, in the five lower rows (excluding the basal row) is a little larger than the other spots in 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 quite overlooked. The next higher spot in each row does not differ at all from the upper ones in the same row, although in the following series it becomes, as we shall see, greatly modified. The larger spots occupy exactly the same relative position on this feather as those occupied by the perfect ocelli on the longer wing-feathers.

By looking to the next two or three succeeding secondary wing-feathers, an absolutely insensible gradation can be traced from one of the above-described lower 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. 58). We here see several oblique rows, A, B, C, D (see the lettered diagram), &c., 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. 56 runs down to, and is connected with, one of the ball-and-socket ocelli. Looking to any one row, for instance, B, the lowest spot or 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. This mark corresponds in every respect with the larger, shaded spot, described in the last paragraph (fig. 57), but is more highly developed and more brightly coloured. To the right and above this spot (b), 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). 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 is unusually narrow, elongated, and regular. It apparently consists of a lateral and broken prolongation of the mark (c), as I infer from traces of similar prolongations from the succeeding upper spots; 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 stand in a line parallel to the shaft, and 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.

The successive steps in the shading of the ball-and-socket ocelli can be followed out with equal clearness. The brown, orange, and pale-leaden 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. 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 was before especially noticed), the line of separation being oblique, in the same direction with 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. 57) having some dull fulvous shading on the upper side.

We have now seen that a perfect series can be followed, from two almost simple spots, at first quite distinct from each other, to one of 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 which have been 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, will deny that a simple dark spot with some fulvous shading might be converted, through the approximation and modification of the 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 extremely pretty, some thinking them even more beautiful 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 improvements in the pattern and shading; and this process has been carried on until the wonderful ball-and-socket ocelli have been 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 reflected 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 more clearly see) 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 darkness. Mr. Gould several years ago pointed out to me a humming-bird, the Urosticte benjamini, remarkable from the curious differences presented by the two 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 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,196 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 E. 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. 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.”197 If this be admitted, we can perceive that the males which 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 consequently have left a larger number of offspring to inherit their newly-acquired beauty.

CHAPTER XV.

Birds—continued.

Discussion why the males alone of some species, and both sexes of other species, are brightly coloured—On sexually-limited inheritance, as applied to various structures and to brightly-coloured plumage—Nidification in relation to colour—Loss of nuptial plumage during the winter.

We have in this chapter to consider, why with many kinds of birds the female has not received the same ornaments as the male; and why with many others, both sexes are equally, or almost equally, ornamented? In the following chapter we shall consider why in some few rare cases the female is more conspicuously coloured than the male.

In my ‘Origin of Species’198 I briefly suggested that the long tail of the peacock would be inconvenient, and the conspicuous black colour of the male capercailzie dangerous, to the female during the period of incubation; and consequently that the transmission of these characters from the male to the female offspring had been checked through natural selection. I still think that this may have occurred in some few instances: but after mature reflection on all the facts which I have been able to collect, I am now inclined to believe that when the sexes differ, the successive variations have generally been from the first limited in their transmission to the same sex in which they first appeared. Since my remarks appeared, the subject of sexual coloration has been discussed in some very interesting papers by Mr. Wallace,199 who believes that in almost all cases the successive variations tended at first to be transmitted equally to both sexes; but that the female was saved, through natural selection, from acquiring the conspicuous colours of the male, owing to the danger which she would thus have incurred during incubation.

This view necessitates a tedious discussion on a difficult point, namely whether the transmission of a character, which is at first inherited by both sexes, can be subsequently limited in its transmission, by means of selection, to one sex alone. We must bear in mind, as shewn in the preliminary chapter on sexual selection, that characters which are limited in their development to one sex are always latent in the other. An imaginary illustration will best aid us in seeing the difficulty of the case: we may suppose that a fancier wished to make a breed of pigeons, in which the males alone should be coloured of a pale blue, whilst the females retained their former slaty tint. As with pigeons characters of all kinds are usually transmitted to both sexes equally, the fancier would have to try to convert this latter form of inheritance into sexually-limited transmission. All that he could do would be to persevere in selecting every male pigeon which was in the least degree of a paler blue; and the natural result of this process, if steadily carried on for a long time, and if the pale variations were strongly inherited or often recurred, would be to make his whole stock of a lighter blue. But our fancier would be compelled to match, generation after generation, his pale blue males with slaty females, for he wishes to keep the latter of this colour. The result would generally be the production either of a mongrel piebald lot, or more probably the speedy and complete loss of the pale-blue colour, for the primordial slaty tint would be transmitted with prepotent force. Supposing, however, that some pale-blue males and slaty females were produced during each successive generation, and were always crossed together; then the slaty females would have, if I may use the expression, much blue blood in their veins, for their fathers, grandfathers, etc., will all have been blue birds. Under these circumstances it is conceivable (though I know of no distinct facts rendering it probable) that the slaty females might acquire so strong a latent tendency to pale-blueness, that they would not destroy this colour in their male offspring, their female offspring still inheriting the slaty tint. If so, the desired end of making a breed with the two sexes permanently different in colour might be gained.

The extreme importance, or rather necessity, of the desired character in the above case, namely, pale-blueness, being present though in a latent state in the female, so that the male offspring should not be deteriorated, will be best appreciated as follows: the male of Sœmmerring’s pheasant has a tail thirty-seven inches in length, whilst that of the female is only eight inches; the tail of the male common pheasant is about twenty inches, and that of the female twelve inches long. Now if the female Sœmmerring pheasant with her short tail were crossed with the male common pheasant, there can be no doubt that the male hybrid offspring would have a much longer tail than that of the pure offspring of the common pheasant. On the other hand, if the female common pheasant, with her tail nearly twice as long as that of the female Sœmmerring pheasant, were crossed with the male of the latter, the male hybrid offspring would have a much shorter tail than that of the pure offspring of Sœmmerring’s pheasant.200

Our fancier, in order to make his new breed with the males of a decided pale-blue tint, and the females unchanged, would have to continue selecting the males during many generations; and each stage of paleness would have to be fixed in the males, and rendered latent in the females. The task would be an extremely difficult one, and has never been tried, but might possibly succeed. The chief obstacle would be the early and complete loss of the pale-blue tint, from the necessity of reiterated crosses with the slaty female, the latter not having at first any latent tendency to produce pale-blue offspring.

On the other hand, if one or two males were to vary ever so slightly in paleness, and the variations were from the first limited in their transmission to the male sex, the task of making a new breed of the desired kind would be easy, for such males would simply have to be selected and matched with ordinary females. An analogous case has actually occurred, for there are breeds of the pigeon in Belgium 201 in which the males alone are marked with black striæ. In the case of the fowl, variations of colour limited in their transmission to the male sex habitually occur. Even when this form of inheritance prevails, it might well happen that some of the successive steps in the process of variation might be transferred to the female, who would then come to resemble in a slight degree the male, as occurs in some breeds of the fowl. Or again, the greater number, but not all, of the successive steps might be transferred to both sexes, and the female would then closely resemble the male. There can hardly be a doubt that this is the cause of the male pouter pigeon having a somewhat larger crop, and of the male carrier pigeon having somewhat larger wattles, than their respective females; for fanciers have not selected one sex more than the other, and have had no wish that these characters should be more strongly displayed in the male than in the female, yet this is the case with both breeds.

The same process would have to be followed, and the same difficulties would be encountered, if it were desired to make a breed with the females alone of some new colour.

Lastly, our fancier might wish to make a breed with the two sexes differing from each other, and both from the parent-species. Here the difficulty would be extreme, unless the successive variations were from the first sexually limited on both sides, and then there would be no difficulty. We see this with the fowl; thus the two sexes of the pencilled Hamburghs differ greatly from each other, and from the two sexes of the aboriginal Gallus bankiva; and both are now kept constant to their standard of excellence by continued selection, which would be impossible unless the distinctive characters of both were limited in their transmission. The Spanish fowl offers a more curious case; the male has an immense comb, but some of the successive variations, by the accumulation of which it was acquired, appear to have been transferred to the female; for she has a comb many times larger than that of the females of the parent-species. But the comb of the female differs in one respect from that of the male, for it is apt to lop over; and within a recent period it has been ordered by the fancy that this should always be the case, and success has quickly followed the order. Now the lopping of the comb must be sexually limited in its transmission, otherwise it would prevent the comb of the male from being perfectly upright, which would be abhorrent to every fancier. On the other hand the uprightness of the comb in the male must likewise be a sexually-limited character, otherwise it would prevent the comb of the female from lopping over.

From the foregoing illustrations, we see that even with almost unlimited time at command, it would be an extremely difficult and complex process, though perhaps not impossible, to change through selection one form of transmission into the other. Therefore, without distinct evidence in each case, I am unwilling to admit that this has often been effected with natural species. On the other hand by means of successive variations, which were from the first sexually limited in their transmission, there would not be the least difficulty in rendering a male bird widely different in colour or in any other character from the female; the latter being left unaltered, or slightly altered, or specially modified for the sake of protection.

As bright colours are of service to the males in their rivalry with other males, such colours would be selected, whether or not they were transmitted exclusively to the same sex. Consequently the females might be expected often to partake of the brightness of the males to a greater or less degree; and this occurs with a host of species. If all the successive variations were transmitted equally to both sexes, the females would be undistinguishable from the males; and this likewise occurs with many birds. If, however, dull colours were of high importance for the safety of the female during incubation, as with many ground birds, the females which varied in brightness, or which received through inheritance from the males any marked accession of brightness, would sooner or later be destroyed. But the tendency in the males to continue for an indefinite period transmitting to their female offspring their own brightness, would have to be eliminated by a change in the form of inheritance; and this, as shewn by our previous illustration, would be extremely difficult. The more probable result of the long-continued destruction of the more brightly-coloured females, supposing the equal form of transmission to prevail, would be the lessening or annihilation of the bright colours of the males, owing to their continually crossing with the duller females. It would be tedious to follow out all the other possible results; but I may remind the reader, as shewn in the eighth chapter, that if sexually-limited variations in brightness occurred in the females, even if they were not in the least injurious to them and consequently were not eliminated, yet they would not be favoured or selected, for the male usually accepts any female, and does not select the more attractive individuals; consequently these variations would be liable to be lost, and would have little influence on the character of the race; and this will aid in accounting for the females being commonly less brightly-coloured than the males.

In the chapter just referred to, instances were given, and any number might have been added, of variations occurring at different ages, and inherited at the same age. It was also shewn that variations which occur late in life are commonly transmitted to the same sex in which they first appeared; whilst variations occurring early in life are apt to be transmitted to both sexes; not that all the cases of sexually-limited transmission can thus be accounted for. It was further shewn that if a male bird varied by becoming brighter whilst young, such variations would be of no service until the age for reproduction had arrived, and there was competition between rival males. If we suppose that three-fourths of the young males of any species are on an average destroyed by various enemies; then the chances would be as three to one against any one individual more brightly-coloured than usual surviving to propagate its kind. But in the case of birds which live on the ground and which commonly need the protection of dull colours, bright tints would be far more dangerous to the young and inexperienced than to the adult males. Consequently the males which varied in brightness whilst young would suffer much destruction and be eliminated through natural selection; on the other hand the males which varied in this manner when nearly mature, notwithstanding that they were exposed to some additional danger, might survive, and from being favoured through sexual selection, would procreate their kind. The brightly-coloured young males being destroyed and the mature ones being successful in their courtship, may account, on the principle of a relation existing between the period of variation and the form of transmission, for the males alone of many birds, having acquired and transmitted brilliant colours to their male offspring alone. But I by no means wish to maintain that the influence of age on the form of transmission is indirectly the sole cause of the great difference in brilliancy between the sexes of many birds.

As with all birds in which the sexes differ in colour, it is an interesting question whether the males alone have been modified through sexual selection, the females being left, as far as this agency is concerned, unchanged or only partially changed; or whether the females have been specially modified through natural selection for the sake of protection, I will discuss this question at considerable length, even at greater length than its intrinsic importance deserves; for various curious collateral points may thus be conveniently considered.

Before we enter on the subject of colour, more especially in reference to Mr. Wallace’s conclusions, it may be useful to discuss under a similar point of view some other differences between the sexes. A breed of fowls formerly existed in Germany 202 in which the hens were furnished with spurs; they were good layers, but they so greatly disturbed their nests with their spurs that they could not be allowed to sit on their own eggs. Hence at one time it appeared to me probable that with the females of the wild Gallinaceæ the development of spurs had been checked through natural selection, from the injury thus caused to their nests. This seemed all the more probable as the wing-spurs, which could not be injurious during nidification, are often as well developed in the female as in the male; though in not a few cases they are rather larger in the male. When the male is furnished with leg-spurs the female almost always exhibits rudiments of them,—the rudiment sometimes consisting of a mere scale, as with the species of Gallus. Hence it might be argued that the females had aboriginally been furnished with well-developed spurs, but that these had subsequently been lost either through disuse or natural selection. But if this view be admitted, it would have to be extended to innumerable other cases; and it implies that the female progenitors of the existing spur-bearing species were once encumbered with an injurious appendage.

In some few genera and species, as in Galloperdix, Acomus, and the Javan peacock (Pavo muticus), the females, as well as the males, possess well-developed spurs. Are we to infer from this fact that they construct a different sort of nest, not liable to be injured by their spurs, from that made by their nearest allies, so that there has been no need for the removal of their spurs? Or are we to suppose that these females especially require spurs for their defence? It is a more probable conclusion that both the presence and absence of spurs in the females result from different laws of inheritance having prevailed, independently of natural selection. With the many females in which spurs appear as rudiments, we may conclude that some few of the successive variations, through which they were developed in the males, occurred very early in life, and were as a consequence transferred to the females. In the other and much rarer cases, in which the females possess fully developed spurs, we may conclude that all the successive variations were transferred to them; and that they gradually acquired the inherited habit of not disturbing their nests.

The vocal organs and the variously-modified feathers for producing sound, as well as the proper instincts for using them, often differ in the two sexes, but are sometimes the same in both. Can such differences be accounted for by the males having acquired these organs and instincts, whilst the females have been saved from inheriting them, on account of the danger to which they would have been exposed by attracting the attention of birds or beasts of prey? This does not seem to me probable, when we think of the multitude of birds which with impunity gladden the country with their voices during the spring.203 It is a safer conclusion that as vocal and instrumental organs are of special service only to the males during their courtship, these organs were developed through sexual selection and continued use in this sex alone—the successive variations and the effects of use having been from the first limited in their transmission in a greater or less degree to the male offspring.

Many analogous cases could be advanced; for instance the plumes on the head, which are generally longer in the male than in the female, sometimes of equal length in both sexes, and occasionally absent in the female,—these several cases sometimes occurring in the same group of birds. It would be difficult to account for a difference of this kind between the sexes on the principle of the female having been benefited by possessing a slightly shorter crest than the male, and its consequent diminution or complete suppression through natural selection. But I will take a more favourable case, namely, the length of the tail. The long train of the peacock would have been not only inconvenient but dangerous to the peahen during the period of incubation and whilst accompanying her young. Hence there is not the least à priori improbability in the development of her tail having been checked through natural selection. But the females of various pheasants, which apparently are exposed on their open nests to as much danger as the peahen, have tails of considerable length. The females as well as the males of the Menura superba have long tails, and they build a domed nest, which is a great anomaly in so large a bird. Naturalists have wondered how the female Menura could manage her tail during incubation; but it is now known 204 that she “enters the nest head first, and then turns round with her tail sometimes over her back, but more often bent round by her side. Thus in time the tail becomes quite askew, and is a tolerable guide to the length of time the bird has been sitting.” Both sexes of an Australian kingfisher (Tanysiptera sylvia) have the middle tail-feathers greatly lengthened; and as the female makes her nest in a hole, these feathers become, as I am informed by Mr. R. B. Sharpe, much crumpled during nidification.

In these two cases the great length of the tail-feathers must be in some degree inconvenient to the female; and as in both species the tail-feathers of the female are somewhat shorter than those of the male, it might be argued that their full development had been prevented through natural selection. Judging from these cases, if with the peahen, the development of the tail had been checked only when it became inconveniently or dangerously long, she would have acquired a much longer tail than she actually possesses; for her tail is not nearly so long, relatively to the size of her body, as that of many female pheasants, nor longer than that of the female turkey. It must also be borne in mind, that in accordance with this view as soon as the tail of the peahen became dangerously long, and its development was consequently checked, she would have continually reacted on her male progeny, and thus have prevented the peacock from acquiring his present magnificent train. We may therefore infer that the length of the tail in the peacock and its shortness in the peahen are the result of the requisite variations in the male having been from the first transmitted to the male offspring alone.

We are led to a nearly similar conclusion with respect to the length of the tail in the various species of pheasants. In the Eared pheasant (Crossoptilon auritum) the tail is of equal length in both sexes, namely, sixteen or seventeen inches; in the common pheasant it is about twenty inches long in the male, and twelve in the female; in Sœmmerring’s pheasant, thirty-seven inches in the male, and only eight in the female; and lastly in Reeve’s pheasant it is sometimes actually seventy-two inches long in the male and sixteen in the female. Thus in the several species, the tail of the female differs much in length, irrespectively of that of the male; and this can be accounted for as it seems to me, with much more probability, by the laws of inheritance,—that is by the successive variations having been from the first more or less closely limited in their transmission to the male sex,—than by the agency of natural selection, owing to the length of tail having been injurious in a greater or less degree to the females of the several species.

We may now consider Mr. Wallace’s arguments, in regard to the sexual coloration of birds. He believes that the bright tints originally acquired through sexual selection by the males, would in all or almost all cases have been transmitted to the females, unless the transference had been checked through natural selection. I may here remind the reader that various facts bearing on this view have already been given under reptiles, amphibians, fishes, and lepidoptera. Mr. Wallace rests his belief chiefly, but not exclusively, as we shall see in the next chapter, on the following statement,205 that when both sexes are coloured in a strikingly-conspicuous manner the nest is of such a nature as to conceal the sitting bird; but when there is a marked contrast of colour between the sexes, the male being gay and the female dull-coloured, the nest is open and exposes the sitting bird to view. This coincidence, as far as it goes, certainly supports the belief that the females which sit on open nests have been specially modified for the sake of protection. Mr. Wallace admits that there are, as might have been expected, some exceptions to his two rules, but it is a question whether the exceptions are not so numerous as seriously to invalidate them.

There is in the first place much truth in the Duke of Argyll’s remark 206 that a large domed nest is more conspicuous to an enemy, especially to all tree-haunting carnivorous animals, than a smaller open nest. Nor must we forget that with many birds which build open nests the males sit on the eggs and aid in feeding the young as well as the females: this is the case, for instance, with Pyranga æstiva,207 one of the most splendid birds in the United States, the male being vermilion, and the female light brownish-green. Now if brilliant colours had been extremely dangerous to birds whilst sitting on their open nests, the males in these cases would have suffered greatly. It might, however, be of such paramount importance to the male to be brilliantly coloured, in order to beat his rivals, that this would more than compensate for some additional danger.

Mr. Wallace admits that with the King-crows (Dicrurus), Orioles, and Pittidæ, the females are conspicuously coloured, yet they build open nests; but he urges that the birds of the first group are highly pugnacious and could defend themselves; that those of the second group take extreme care in concealing their open nests, but this does not invariably hold good;208 and that with the birds of the third group the females are brightly coloured chiefly on the under surface. Besides these cases the whole great family of pigeons, which are sometimes brightly, and almost always conspicuously coloured, and which are notoriously liable to the attacks of birds of prey, offers a serious exception to the rule, for pigeons almost always build open and exposed nests. In another large family, that of the Humming-birds, all the species build open nests, yet with some of the most gorgeous species the sexes are alike; and in the majority, the females, though less brilliant than the males, are very brightly coloured. Nor can it be maintained that all female humming-birds, which are brightly coloured, escape detection by their tints being green, for some display on their upper surfaces red, blue, and other colours.209

In regard to birds which build in holes or construct domed nests, other advantages, as Mr. Wallace remarks, besides concealment are gained, such as shelter from the rain, greater warmth, and in hot countries protection from the rays of the sun;210 so that it is no valid objection to his view that many birds having both sexes obscurely coloured build concealed nests.211 The female Hornbills (Buceros), for instance, of India and Africa are protected, during nidification, with extraordinary care, for the male plaisters up the hole in which the female sits on her eggs, and leaves only a small orifice through which he feeds her; she is thus kept a close prisoner during the whole period of incubation;212 yet female hornbills are not more conspicuously coloured than many other birds of equal size which build open nests. It is a more serious objection to Mr. Wallace’s view, as is admitted by him, that in some few groups the males are brilliantly coloured and the females obscure, and yet the latter hatch their eggs in domed nests. This is the case with the Grallinæ of Australia, the Superb Warblers (Maluridæ) of the same country, the Sun-birds (Nectariniæ), and with several of the Australian Honey-suckers or Meliphagidæ.213

If we look to the birds of England we shall see that there is no close and general relation between the colours of the female and the nature of the nest constructed by her. About forty of our British birds (excluding those of large size which could defend themselves) build in holes in banks, rocks, or trees, or construct domed nests. If we take the colours of the female goldfinch, bullfinch, or blackbird, as a standard of the degree of conspicuousness, which is not highly dangerous to the sitting female, then out of the above forty birds, the females of only twelve can be considered as conspicuous to a dangerous degree, the remaining twenty-eight being inconspicuous.214 Nor is there any close relation between a well-pronounced difference in colour between the two sexes, and the nature of the nest constructed. Thus the male house-sparrow (Passer domesticus) differs much from the female, the male tree-sparrow (P. montanus) differs hardly at all, and yet both build well-concealed nests. The two sexes of the common fly-catcher (Muscicapa grisola) can hardly be distinguished, whilst the sexes of the pied fly-catcher (M. luctuosa) differ considerably, and both build in holes. The female blackbird (Turdus merula) differs much, the female ring-ouzel (T. torquatus) differs less, and the female common thrush (T. musicus) hardly at all from their respective males; yet all build open nests. On the other hand, the not very distantly-allied water-ouzel (Cinclus aquaticus) builds a domed nest, and the sexes differ about as much as in the case of the ring-ouzel. The black and red grouse (Tetrao tetrix and T. Scoticus) build open nests, in equally well-concealed spots, but in the one species the sexes differ greatly, and in the other very little.

Notwithstanding the foregoing objections, I cannot doubt, after reading Mr. Wallace’s excellent essay, that looking to the birds of the world, a large majority of the species in which the females are conspicuously coloured (and in this case the males with rare exceptions are equally conspicuous), build concealed nests for the sake of protection. Mr. Wallace enumerates 215 a long series of groups in which this rule holds good; but it will suffice here to give, as instances, the more familiar groups of kingfishers, toucans, trogons, puff-birds (Capitonidæ), plaintain-eaters (Musophagæ), woodpeckers, and parrots. Mr. Wallace believes that in these groups, as the males gradually acquired through sexual selection their brilliant colours, these were transferred to the females and were not eliminated by natural selection, owing to the protection which they already enjoyed from their manner of nidification. According to this view, their present manner of nesting was acquired before their present colours. But it seems to me much more probable that in most cases as the females were gradually rendered more and more brilliant from partaking of the colours of the male, they were gradually led to change their instincts (supposing that they originally built open nests), and to seek protection by building domed or concealed nests. No one who studies, for instance, Audubon’s account of the differences in the nests of the same species in the Northern and Southern United States,216 will feel any great difficulty in admitting that birds, either by a change (in the strict sense of the word) of their habits, or through the natural selection of so-called spontaneous variations of instinct, might readily be led to modify their manner of nesting.

This way of viewing the relation, as far as it holds good, between the bright colours of female birds and their manner of nesting, receives some support from certain analogous cases occurring in the Sahara Desert. Here, as in most other deserts, various birds, and many other animals, have had their colours adapted in a wonderful manner to the tints of the surrounding surface. Nevertheless there are, as I am informed by the Rev. Mr. Tristram, some curious exceptions to the rule; thus the male of the Monticola cyanea is conspicuous from his bright blue colour, and the female almost equally conspicuous from her mottled brown and white plumage; both sexes of two species of Dromolæa are of a lustrous black; so that these three birds are far from receiving protection from their colours, yet they are able to survive, for they have acquired the habit, when in danger, of taking refuge in holes or crevices in the rocks.

With respect to the above-specified groups of birds, in which the females are conspicuously coloured and build concealed nests, it is not necessary to suppose that each separate species had its nidifying instinct specially modified; but only that the early progenitors of each group were gradually led to build domed or concealed nests; and afterwards transmitted this instinct, together with their bright colours, to their modified descendants. This conclusion, as far as it can be trusted, is interesting, namely, that sexual selection, together with equal or nearly equal inheritance by both sexes, have indirectly determined the manner of nidification of whole groups of birds.

Even in the groups in which, according to Mr. Wallace, the females from being protected during nidification, have not had their bright colours eliminated through natural selection, the males often differ in a slight, and occasionally in a considerable degree, from the females. This is a significant fact, for such differences in colour must be accounted for on the principle of some of the variations in the males having been from the first limited in their transmission to the same sex; as it can hardly be maintained that these differences, especially when very slight, serve as a protection to the female. Thus all the species in the splendid group of the Trogons build in holes; and Mr. Gould gives figures 217 of both sexes of twenty-five species, in all of which, with one partial exception, the sexes differ sometimes slightly, sometimes conspicuously, in colour,—the males being always more beautiful than the females, though the latter are likewise beautiful. All the species of kingfisher build in holes, and with most of the species the sexes are equally brilliant, and thus far Mr. Wallace’s rule holds good; but in some of the Australian species the colours of the females are rather less vivid than those of the male; and in one splendidly-coloured species, the sexes differ so much that they were at first thought to be specifically distinct.218 Mr. R. B. Sharpe, who has especially studied this group, has shewn me some American species (Ceryle) in which the breast of the male is belted with black. Again, in Carcineutes, the difference between the sexes is conspicuous: in the male the upper surface is dull-blue banded with black, the lower surface being partly fawn-coloured, and there is much red about the head; in the female the upper surface is reddish-brown banded with black, and the lower surface white with black markings. It is an interesting fact, as shewing how the same peculiar style of sexual colouring often characterises allied forms, that in three species of Dacelo the male differs from the female only in the tail being dull-blue banded with black, whilst that of the female is brown with blackish bars; so that here the tail differs in colour in the two sexes in exactly the same manner as the whole upper surface in the sexes of Carcineutes.

A. Specimen, from Mauritius, upper surface of fore-wing	B. Specimen, from Java, upper surface of hind-wing.
A¹. Specimen, from Natal, ditto.	B¹. Specimen, from Mauritius, ditto.

A, B, C, &c. Rows of spots running down to and forming the elliptic ornaments.	c. The next succeeding spot or mark in the same row.
b. Lowest spot or mark in row B.	d. Apparently a broken prolongation of the spot c in the same row B.

CHAPTER XV.

Birds—continued.