In cases such as we have discussed, it is obvious that there is no possible explanation except through selection. This brings us to the last kind of secondary sexual characters, and the one in regard to which doubt has been most frequently expressed,--decorative colours and decorative forms, the brilliant plumage of the male pheasant, the humming-birds, and the bird of Paradise, as well as the bright colours of many species of butterfly, from the beautiful blue of our little Lycaenidae to the magnificent azure of the large Morphinae of Brazil. In a great many cases, though not by any means in all, the male butterflies are "more beautiful" than the females, and in the Tropics in particular they shine and glow in the most superb colours.
I really see no reason why we should doubt the power of sexual selection, and I myself stand wholly on Darwin's side. Even though we certainly cannot assume that the females exercise a conscious choice of the "handsomest" mate, and deliberate like the judges in a court of justice over the perfections of their wooers, we have no reason to doubt that distinctive forms (decorative feathers) and colours have a particularly exciting effect upon the female, just as certain odours have among animals of so many different groups, including the butterflies. The doubts which existed for a considerable time, as a result of fallacious experiments, as to whether the colours of flowers really had any influence in attracting butterflies have now been set at rest through a series of more careful investigations; we now know that the colours of flowers are there on account of the butterflies, as Sprengel first showed, and that the blossoms of Phanerogams are selected in relation to them, as Darwin pointed out.
Certainly it is not possible to bring forward any convincing proof of the origin of decorative colours through sexual selection, but there are many weighty arguments in favour of it, and these form a body of presumptive evidence so strong that it almost amounts to certainty.
In the first place, there is the analogy with other secondary sexual characters. If the song of birds and the chirping of the cricket have been evolved through sexual selection, if the penetrating odours of male animals,--the crocodile, the musk-deer, the beaver, the carnivores, and, finally, the flower-like fragrances of the butterflies have been evolved to their present pitch in this way, why should decorative colours have arisen in some other way? Why should the eye be less sensitive to SPECIFICALLYMALE colours and other VISIBLE signs ENTICING TO THE FEMALE, than the olfactory sense to specifically male odours, or the sense of hearing to specifically male sounds? Moreover, the decorative feathers of birds are almost always spread out and displayed before the female during courtship.
I have elsewhere ("The Evolution Theory", London, 1904, I. page 219.)pointed out that decorative colouring and sweet-scentedness may replace one another in Lepidoptera as well as in flowers, for just as some modestly coloured flowers (mignonette and violet) have often a strong perfume, while strikingly coloured ones are sometimes quite devoid of fragrance, so we find that the most beautiful and gaily-coloured of our native Lepidoptera, the species of Vanessa, have no scent-scales, while these are often markedly developed in grey nocturnal Lepidoptera. Both attractions may, however, be combined in butterflies, just as in flowers. Of course, we cannot explain why both means of attraction should exist in one genus, and only one of them in another, since we do not know the minutest details of the conditions of life of the genera concerned. But from the sporadic distribution of scent-scales in Lepidoptera, and from their occurrence or absence in nearly related species, we may conclude that fragrance is a relatively MODERN acquirement, more recent than brilliant colouring.
One thing in particular that stamps decorative colouring as a product of selection is ITS GRADUAL INTENSIFICATION by the addition of new spots, which we can quite well observe, because in many cases the colours have been first acquired by the males, and later transmitted to the females by inheritance. The scent-scales are never thus transmitted, probably for the same reason that the decorative colours of many birds are often not transmitted to the females: because with these they would be exposed to too great elimination by enemies. Wallace was the first to point out that in species with concealed nests the beautiful feathers of the male occurred in the female also, as in the parrots, for instance, but this is not the case in species which brood on an exposed nest. In the parrots one can often observe that the general brilliant colouring of the male is found in the female, but that certain spots of colour are absent, and these have probably been acquired comparatively recently by the male and have not yet been transmitted to the female.
Isolation of the group of individuals which is in process of varying is undoubtedly of great value in sexual selection, for even a solitary conspicuous variation will become dominant much sooner in a small isolated colony, than among a large number of members of a species.