In some analogous cases, namely with birds having a distinct summer and winter plumage, but with the two sexes nearly alike, certain closely-allied species can easily be distinguished in their summer or nuptial plumage, yet are undistinguishable in their winter as well as in their immature plumage. This is the case with some of the closely-allied Indian wagtails or Motacillæ. Mr. Swinhoe[234] informs me that three species of Ardeola, a genus of herons, which represent each other on separate continents, are “most strikingly different” when ornamented with their summer plumes, but are hardly, if at all, distinguishable during the winter. The young also of these three species in their immature plumage closely resemble the adults in their winter dress. This case is all the more interesting because with two other species of Ardeola both sexes retain, during the winter and summer, nearly the same plumage as that possessed by the three first species during the winter and in their immature state; and this plumage, which is common to several distinct species at different ages and seasons, probably shews us how the progenitor of the genus was coloured. In all these cases, the nuptial plumage which we may assume was originally acquired by the adult males during the breeding-season, and transmitted to the adults of both sexes at the corresponding season, has been modified, whilst the winter and immature plumages have been left unchanged.
The question naturally arises, how is it that in these latter cases the winter plumage of both sexes, and in the former cases the plumage of the adult females, as well as the immature plumage of the young, have not been at all affected? The species which represent each other in distinct countries will almost always have been exposed to somewhat different conditions, but we can hardly attribute the modification of the plumage in the males alone to this action, seeing that the females and the young, though similarly exposed, have not been affected. Hardly any fact in nature shews us more clearly how subordinate in importance is the direct action of the conditions of life, in comparison with the accumulation through selection of indefinite variations, than the surprising difference between the sexes of many birds; for both sexes must have consumed the same food and have been exposed to the same climate. Nevertheless we are not precluded from believing that in the course of time new conditions may produce some direct effect; we see only that this is subordinate in importance to the accumulated results of selection. When, however, a species migrates into a new country, and this must precede the formation of representative species, the changed conditions to which they will almost always have been exposed will cause them to undergo, judging from a widely-spread analogy, a certain amount of fluctuating variability. In this case sexual selection, which depends on an element eminently liable to change—namely the taste or admiration of the female—will have had new shades of colour or other differences to act on and accumulate; and as sexual selection is always at work, it would (judging from what we know of the results on domestic animals of man’s unintentional selection), be a surprising fact if animals inhabiting separate districts, which can never cross and thus blend their newly-acquired characters, were not, after a sufficient lapse of time, differently modified. These remarks likewise apply to the nuptial or summer plumage, whether confined to the males or common to both sexes.
Although the females of the above closely-allied species, together with their young, differ hardly at all from each other, so that the males alone can be distinguished, yet in most cases the females of the species within the same genus obviously differ from each other. The differences, however, are rarely as great as between the males. We see this clearly in the whole family of the Gallinaceæ: the females, for instance, of the common and Japan pheasant, and especially of the gold and Amherst pheasant—of the silver pheasant and the wild fowl—resemble each other very closely in colour, whilst the males differ to an extraordinary degree. So it is with the females of most of the Cotingidæ, Fringillidæ, and many other families. There can indeed be no doubt that, as a general rule, the females have been modified to a less extent than the males. Some few birds, however, offer a singular and inexplicable exception; thus the females of Paradisea apoda and P. papuana differ from each other more than do their respective males;[235] the female of the latter species having the under surface pure white, whilst the female P. apoda is deep brown beneath. So, again, as I hear from Professor Newton, the males of two species of Oxynotus (shrikes), which represent each other in the islands of Mauritius and Bourbon,[236] differ but little in colour, whilst the females differ much. In the Bourbon species the female appears to have partially retained an immature condition of plumage, for at first sight she “might be taken for the young of the Mauritian species.” These differences may be compared with those which occur, independently of selection by man, and which we cannot explain, in certain sub-breeds of the game-fowl, in which the females are very different, whilst the males can hardly be distinguished.[237]
As I account so largely by sexual selection for the differences between the males of allied species, how can the differences between the females be accounted for in all ordinary cases? We need not here consider the species which belong to distinct genera; for with these, adaptation to different habits of life, and other agencies, will have come into play. In regard to the differences between the females within the same genus, it appears to me almost certain, after looking through various large groups, that the chief agent has been the transference, in a greater or less degree, to the female of the characters acquired by the males through sexual selection. In the several British finches, the two sexes differ either very slightly or considerably; and if we compare the females of the greenfinch, chaffinch, goldfinch, bullfinch, crossbill, sparrow, &c., we shall see that they differ from each other chiefly in the points in which they partially resemble their respective males; and the colours of the males may safely be attributed to sexual selection. With many gallinaceous species the sexes differ to an extreme degree, as with the peacock, pheasant, and fowl, whilst with other species there has been a partial or even complete transference of character from the male to the female. The females of the several species of Polyplectron exhibit in a dim condition, and chiefly on the tail, the splendid ocelli of their males. The female partridge differs from the male only in the red mark on her breast being smaller; and the female wild turkey only in her colours being much duller. In the guinea-fowl the two sexes are undistinguishable. There is no improbability in the plain, though peculiar spotted plumage of this latter bird having been acquired through sexual selection by the males, and then transmitted to both sexes; for it is not essentially different from the much more beautifully-spotted plumage, characteristic of the males alone of the Tragopan pheasants.
It should be observed that, in some instances, the transference of characters from the male to the female has been effected apparently at a remote period, the male having subsequently undergone great changes, without transferring to the female any of his later-gained characters. For instance, the female and the young of the black-grouse (Tetrao tetrix) resemble pretty closely both sexes and the young of the red-grouse T. Scoticus; and we may consequently infer that the black-grouse is descended from some ancient species, of which both sexes were coloured in nearly the same manner as the red-grouse. As both sexes of this latter species are more plainly barred during the breeding-season than at any other time, and as the male differs slightly from the female in his more strongly-pronounced red and brown tints,[238] we may conclude that his plumage has been, at least to a certain extent, influenced by sexual selection. If so, we may further infer that the nearly similar plumage of the female black-grouse was similarly produced at some former period. But since this period the male black-grouse has acquired his fine black plumage, with his forked and outwardly-curled tail-feathers; but of these characters there has hardly been any transference to the female, excepting that she shews in her tail a trace of the curved fork.
We may therefore conclude that the females of distinct though allied species have often had their plumage rendered more or less different by the transference in various degrees, of characters acquired, both during former and recent times, by the males through sexual selection. But it deserves especial attention that brilliant colours have been transferred much more rarely than other tints. For instance, the male of the red-throated bluebreast (Cyanecula suecica) has a rich blue breast, including a sub-triangular red mark; now marks of approximately the same shape have been transferred to the female, but the central space is fulvous instead of red, and is surrounded by mottled instead of blue feathers. The Gallinaceæ offer many analogous cases; for none of the species, such as partridges, quails, guinea-fowls, &c., in which the colours of the plumage have been largely transferred from the male to the female, are brilliantly coloured. This is well exemplified with the pheasants, in which the male is generally so much more brilliant than the female; but with the Eared and Cheer pheasants (Crossoptilon auritum and Phasianus Wallichii) the two sexes closely resemble each other and their colours are dull. We may go so far as to believe that if any part of the plumage in the males of these two pheasants had been brilliantly coloured, this would not have been transferred to the females. These facts strongly support Mr. Wallace’s view that with birds which are exposed to much danger during nidification, the transference of bright colours from the male to the female has been checked through natural selection. We must not, however, forget that another explanation, before given, is possible; namely, that the males which varied and became bright, whilst they were young and inexperienced, would have been exposed to much danger, and would generally have been destroyed; the older and more cautious males, on the other hand, if they varied in a like manner, would not only have been able to survive, but would have been favoured in their rivalry with other males. Now variations occurring late in life tend to be transmitted exclusively to the same sex, so that in this case extremely bright tints would not have been transmitted to the females. On the other hand, ornaments of a less conspicuous kind, such as those possessed by the Eared and Cheer pheasants, would not have been dangerous, and if they appeared during early youth, would generally have been transmitted to both sexes.
In addition to the effects of the partial transference of characters from the males to the females, some of the differences between the females of closely-allied species may be attributed to the direct or definite action of the conditions of life.[239] With the males any such action would generally have been masked by the brilliant colours gained through sexual selection; but not so with the females. Each of the endless diversities in plumage, which we see in our domesticated birds is, of course, the result of some definite cause; and under natural and more uniform conditions, some one tint, assuming that it was in no way injurious, would almost certainly sooner or later prevail. The free intercrossing of the many individuals belonging to the same species would ultimately tend to make any change of colour, thus induced, uniform in character.
No one doubts that both sexes of many birds have had their colours adapted for the sake of protection; and it is possible that the females alone of some species may have been thus modified. Although it would be a difficult, perhaps an impossible process, as shewn in the last chapter, to convert through selection one form of transmission into another, there would not be the least difficulty in adapting the colours of the female, independently of those of the male, to surrounding objects, through the accumulation of variations which were from the first limited in their transmission to the female sex. If the variations were not thus limited, the bright tints of the male would be deteriorated or destroyed. Whether the females alone of many species have been thus specially modified, is at present very doubtful. I wish I could follow Mr. Wallace to the full extent; for the admission would remove some difficulties. Any variations which were of no service to the female as a protection would be at once obliterated, instead of being lost simply by not being selected, or from free intercrossing, or from being eliminated when transferred to the male and in any way injurious to him. Thus the plumage of the female would be kept constant in character. It would also be a relief if we could admit that the obscure tints of both sexes of many birds had been acquired and preserved for the sake of protection,—for example, of the hedge-warbler or kitty-wren (Accentor modularis and Troglodytes vulgaris), with respect to which we have no sufficient evidence of the action of sexual selection. We ought, however, to be cautious in concluding that colours which appear to us dull, are not attractive to the females of certain species; we should bear in mind such cases as that of the common house-sparrow, in which the male differs much from the female, but does not exhibit any bright tints. No one probably will dispute that many gallinaceous birds which live on the open ground have acquired their present colours, at least in part, for the sake of protection. We know how well they are thus concealed; we know that ptarmigans, whilst changing from their winter to their summer plumage, both of which are protective, suffer greatly from birds of prey. But can we believe that the very slight differences in tints and markings between, for instance, the female black and red-grouse serve as a protection? Are partridges, as they are now coloured, better protected than if they had resembled quails? Do the slight differences between the females of the common pheasant, the Japan and golden pheasants, serve as a protection, or might not their plumages have been interchanged with impunity? From what Mr. Wallace has observed of the habits of certain gallinaceous birds in the East he thinks that such slight differences are beneficial. For myself, I will only say that I am not convinced.
Formerly when I was inclined to lay much stress on the principle of protection, as accounting for the less bright colours of female birds, it occurred to me that possibly both sexes and the young might aboriginally have been brightly coloured in an equal degree; but that subsequently, the females from the danger incurred during incubation, and the young from being inexperienced, had been rendered dull as a protection. But this view is not supported by any evidence, and is not probable; for we thus in imagination expose during past times the females and the young to danger, from which it has subsequently been necessary to shield their modified descendants. We have, also, to reduce, through a gradual process of selection, the females and the young to almost exactly the same tints and markings, and to transmit them to the corresponding sex and period of life. It is also a somewhat strange fact, on the supposition that the females and the young have partaken during each stage of the process of modification of a tendency to be as brightly coloured as the males, that the females have never been rendered dull-coloured without the young participating in the same change; for there are no instances, as far as I can discover, of species with the females dull-coloured and the young bright-coloured. A partial exception, however, is offered by the young of certain woodpeckers, for they have “the whole upper part of the head tinged with red,” which afterwards either decreases into a mere circular red line in the adults of both sexes, or quite disappears in the adult females.[240]
Finally, with respect to our present class of cases, the most probable view appears to be that successive variations in brightness or in other ornamental characters, occurring in the males at a rather late period of life have alone been preserved; and that most or all of these variations owing to the late period of life at which they appeared, have been from the first transmitted only to the adult male offspring. Any variations in brightness which occurred in the females or in the young would have been of no service to them, and would not have been selected; moreover, if dangerous, would have been eliminated. Thus the females and the young will either have been left unmodified, or, and this has much more commonly occurred, will have been partially modified by receiving through transference from the males some of the successive variations. Both sexes have perhaps been directly acted on by the conditions of life to which they have long been exposed; but the females from not being otherwise much modified will best exhibit any such effects. These changes and all others will have been kept uniform by the free intercrossing of many individuals. In some cases, especially with ground birds, the females and the young may possibly have been modified, independently of the males, for the sake of protection, so as to have acquired the same dull-coloured plumage.