The Descent of Man, Charles Darwin [top 100 books of all time checklist txt] 📗

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 progenitors of the genus were 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 to this action the modification of the plumage in the males alone, seeing that the females and the young, though similarly exposed, have not been affected. Hardly any fact 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 will 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 either on both sexes, or from their constitutional differences chiefly on one sex. We see only that this is subordinate in importance to the accumulated results of selection. Judging, however, from a wide-spread analogy, when 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 a certain amount of fluctuating variability. In this case sexual selection, which depends on an element liable to change—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 (from what we know of the results on domestic animals of man’s unintentional selection), be surprising 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 or representative species, together with their young, differ hardly at all from one another, so that the males alone can be distinguished, yet the females of most 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 Gallinaceae: 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 one another very closely in colour, whilst the males differ to an extraordinary degree. So it is with the females of most of the Cotingidae, Fringillidae, and many other families. There can indeed be no doubt that, as a general rule, the females have been less modified 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 (7. Wallace, ‘The Malay Archipelago,’ vol. ii. 1869, p. 394.); 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 (8. These species are described with coloured figures, by M. F. Pollen, in ‘Ibis,’ 1866, p. 275.), 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 inexplicable ones, which occur independently of man’s selection in certain sub-breeds of the game-fowl, in which the females are very different, whilst the males can hardly be distinguished. (9. ‘Variation of Animals,’ etc., vol. i. p. 251.)

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 greater or less transference 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, etc., we shall see that they differ from one another 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 indistinguishable. There is no improbability in the plain, though peculiarly 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 distinctly 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 (10. Macgillivray, ‘History of British Birds,’ vol. i. pp. 172-174.), we may conclude that his plumage has been influenced by sexual selection, at least to a certain extent. If so, we may further infer that 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 by the males through sexual selection, both during former and recent times. 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 blue- breast (Cyanecula suecica) has a rich blue breast, including a sub- triangular red mark; now marks of nearly 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 Gallinaceae offer many analogous cases; for none of the species, such as partridges, quails, guinea-fowls, etc., 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 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, it 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 incubation, 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. (11. See, on this subject, chap. xxiii. in the ‘Variation of Animals and Plants under Domestication.’) 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.

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