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conscious sexual selection, that is, the actual choice by the females of the more brilliantly-coloured males, I believe very little if any effect is directly due. It is undoubtedly proved that in birds the females do sometimes exert a choice; but the evidence of this fact collected by Mr. Darwin (Descent of Man, chap. xiv.) does not prove that colour determines that choice, while much of the strongest evidence is directly opposed to this view. All the facts appear to be consistent with the choice depending on a variety of male characteristics, with some of which colour is often correlated. Thus it is the opinion of some of the best observers that vigour and liveliness are most attractive, and these are no doubt usually associated with intensity of colour. Again, the display of the various ornamental appendages of the male during courtship may be attractive; but these appendages, with their bright colours or shaded patterns, are due probably to general laws of growth, and to that superabundant vitality which we have seen to be a cause of colour. But there are many considerations which seem to show that the possession of these ornamental appendages and bright colours in the male is not an important character functionally, and that it has not been produced by the action of conscious sexual selection. Amid the copious mass of facts and opinions collected by Mr. Darwin as to the display of colour and ornaments by the male birds, there is a total absence of any evi. dence that the females admire or even notice this display. The hen, the turkey, and the pea-fowl go on feeding while the male is displaying his finery; and there is reason to believe that it is his persistency and energy rather than his beauty which wins the day. Again, evidence collected by Mr. Darwin himself proves that cach bird finds a mate under any circumstances. He gives a number of cases of one of a pair of birds being shot, and the survivor being alway's found paired again almost immediately. This is sufliciently explained on the assumption that the destruction of birls by various causes is continually leaving widlows and widlowers in nearly equal proportions, and thus each one finds a fresh mate ; and it learls to the conclusion that permanently uprareal birls are very scarce; so that, speaking broadly, every bird finds a mate and breeds. But this would almost or quite neutralize any effect of sexual selection of colour or ornament, since the less liighly-coloured birds woull be at no disadvantage :18 regards leaving healthy offspring. If, however, heightened colour is correlated with health and viyour; and if these healthy and vigorous birds provide best for their young, and leave offspring which, being equally healthy and vigorous, can best provide for themselves—which cannot be denied ; then natural selection becomes a preserver and intensifier of colour.
Another most important consideration is, that male butterflies rival or even excel the most gorgeous male birls in bright colours and elegant patterns; and among these there is literally not one particle of evidence that the female is influenced by colour, or even that she has any power of choice; while there is much direct evidence to the contrary (Descent of Jlan, p. 318). The weakness of the evidence for conscious sexual selection among these insects is so palpable, that Mr. Darwin is obliged to supplement it by the singularly inconclusive argument that, “ Unless the female prefer one male to another, the
pairing must be left to mere chance, and this does not appear probable” (1. c. p. 317). But he has just said — "The males sometimes fight together in rivalry, and many may be seen pursuing or crowding round the same female;" while in the case of the silk-moths,—“the females appear not to evince the least choice in regard to their partners.” Surely the plain inference from all this is, that males fight and struggle for the almost passive female; and that the most vigorous and energetic, the strongest-winged or the most persevering, wins her. How can there be chance in this ? Natural selection woulil here act, as in birds, in perpetuating the strongest and most vigorous males; and as these would usually be the more highly coloured of their race, the same results would be produced as regards the intensification and variation of colour in the one case as in the other.
Let us now see how these principles will apply to some of the cases adduced by Mr. Darwin in support of his theory of conscious sexual selection.
In Descent of Jan, 2nd ed., pp. 307-316, we find an elaborate account of the various modes of colouring of butterflies and moths, proving that the coloured parts are always more or less displayed, and that they have some evident relation to an observer. Mr. Darwin then says : “From the several foregoing facts it is impossible to admit that the brilliant colours of butterflies, and of some few moths, have commonly been acquired for the sake of protection. We have seen that their colours and elegant patterns are arranged and exbibited as if for display. llence I am led to believe that the females prefer or are most excited by the more brilliant males; for on any other supposition the males would, as
far as we can see, be ornamented to no purpose" (l.c., p. 316). I am not aware that any one has ever maintained that the brilliant colours of butterflies have “commonly been acquired for the sake of protection," yet Mr. Darwin has himself referred to cases in which the brilliant colour is so placed as to serve for protection; as for example, the eye-spots on the hind wings of moths, which are pierced by birds and so save the vital parts of the insect; while the bright patch on the orange-tip butterflies which Jr. Darwin denies are protective, may serve the same purpose. It is in fact somewhat remarkable how very generally the black spots, ocelli, or bright patches of colour are on the tips, margins, or discs of the wings; and as the insects are necessarily visible while flying, and this is the time when they are most suliject to attacks by insectivorous birds, the position of the more conspicuous parts at some distance from the body may be a real protection to them. Again, Mr. Darwin admits that the white colour of the male ghost-moth may render it more easily seen by the female while flying about in the dusk; and if to this we add that it will be also more readily distinguished from allied species, we have a reason for diverse ornamentation in these insects quite sufficient to account for most of the facts, without believing in the selection of brilliant males by the females, for which there is not a particle of evidence.
Probulle use of the IIorns of Bectles.-A somewhat analogous case is furnished by the immense horns of some beetles of the families Copridæ and Dynastidæ, which Mr. Darwin admits are not used for fighting, and
See M. Fabre's testimony on this point, Descent of Man, p. 291.
therefore concludes are ornaments, developed through selection of the larger-horned males by the females. But it has been overlooked that these horns may be protective. The males probably fly about most, as is usually the case with male insects; and as they generally fly at dusk they are subject to the attacks of large-mouthed goatsuckers and podargi, as well as of insect-eating owls. Now the long, pointed or forked horns, often divergent, or movable with the head, would render it very difficult for these birds to swallow such insects, and would therefore be an efficient protection; just as are the hooked spines of some stingless ants and the excessively hard integuments of many beetles, against the smaller insectivorous birds.
Cause of the greater Brilliancy of some Female Insects.--The facts given by Mr. Darwin to show that butterflies and other insects can distinguish colours and are attracted by colours similar to their own, are quite consistent with the view that colour, which continually tends to appear, is utilised for purposes of identification and distinction, when not required to be modified or suppressed for the purpose of protection. The cases of the females of some species of Thecla, Callidryas, Colias, and Ilipparchia, which have more conspicuous markings than the male, may be due to several causes: to obtain greater distinction from other species; for protection from birds, as in the case of the yellow-underwing moths; while sometimes--as in Ilipparchia—the lower intensity of colouring in the female may lead to more contrasted markings. Mr. Darwin thinks that here the males have selected the more beautiful females; although one chief fact in support