The males and females of the same species of butterfly are known521 in several cases to inhabit different stations, the former commonly basking in the sunshine, the latter haunting gloomy forests. It is therefore possible that different conditions of life may have acted directly on the two sexes; but this is not probable,522 as in the adult state they are exposed during a very short period to different conditions; and the larvæ of both are exposed to the same conditions. Mr. Wallace believes 404that the less brilliant colours of the female have been specially gained in all or almost all cases for the sake of protection. On the contrary it seems to me more probable that the males alone, in the large majority of cases, have acquired their bright colours through sexual selection, the females having been but little modified. Consequently the females of distinct but allied species ought to resemble each other much more closely than do the males of the same species; and this is the general rule. The females thus approximately show us the primordial colouring of the parent-species of the group to which they belong. They have, however, almost always been modified to a certain extent by some of the successive steps of variation, through the accumulation of which the males were rendered beautiful, having been transferred to them. The males and females of allied though distinct species will also generally have been exposed during their prolonged larval state to different conditions, and may have been thus indirectly affected; though with the males any slight change of colour thus caused will often have been completely masked by the brilliant tints gained through sexual selection. When we treat of Birds, I shall have to discuss the whole question whether the differences in colour between the males and females have been in part specially gained by the latter as a protection; so that I will here only give unavoidable details.
In all cases when the more common form of equal inheritance by both sexes has prevailed, the selection of bright-coloured males would tend to make the females bright-coloured; and the selection of dull-coloured females would tend to make the males dull. If both processes were carried on simultaneously, they would tend to neutralise each other. As far as I can see, it would be extremely difficult to change through selection the405 one form of inheritance into the other. But by the selection of successive variations, which were from the first sexually limited in their transmission, there would not be the slightest difficulty in giving bright colours to the males alone, and at the same time or subsequently, dull colours to the females alone. In this latter manner female butterflies and moths may, as I fully admit, have been rendered inconspicuous for the sake of protection, and widely different from their males.
Mr. Wallace523 has argued with much force in favour of his view that when the sexes differ, the female has been specially modified for the sake of protection; and that this has been effected by one form of inheritance, namely, the transmission of characters to both sexes, having been changed through the agency of natural selection into the other form, namely, transmission to one sex. I was at first strongly inclined to accept this view; but the more I have studied the various classes throughout the animal kingdom, the less probable it has appeared. Mr. Wallace urges that both sexes of the Heliconidæ, Danaidæ, Acroeidæ are equally brilliant because both are protected from the attacks of birds and other enemies, by their offensive odour; but that in other groups, which do not possess this immunity, the females have been rendered inconspicuous, from having more need of protection than the males. This supposed difference in the “need of protection by the two sexes” is rather deceptive, and requires some discussion. It is obvious that brightly-coloured individuals, whether males or females, would equally attract, and obscurely-coloured individuals equally escape, the 406attention of their enemies. But we are concerned with the effects of the destruction or preservation of certain individuals of either sex, on the character of the race. With insects, after the male has fertilised the female, and after the latter has laid her eggs, the greater or less immunity from danger of either sex could not possibly have any effect on the offspring. Before the sexes have performed their proper functions, if they existed in equal numbers and if they strictly paired (all other circumstances being the same), the preservation of the males and females would be equally important for the existence of the species and for the character of the offspring. But with most animals, as is known to be the case with the domestic silk-moth, the male can fertilise two or three females; so that the destruction of the males would not be so injurious to the species as that of the females. On the other hand, Dr. Wallace believes that with moths the progeny from a second or third fertilisation is apt to be weakly, and therefore would not have so good chance of surviving. When the males exist in much greater numbers than the females, no doubt many males might be destroyed with impunity to the species; but I cannot see that the results of ordinary selection for the sake of protection would be influenced by the sexes existing in unequal numbers; for the same proportion of the more conspicuous individuals, whether males or females, would probably be destroyed. If indeed the males presented a greater range of variation in colour, the result would be different; but we need not here follow out such complex details. On the whole I cannot perceive that an inequality in the numbers of the two sexes would influence in any marked manner the effects of ordinary selection on the character of the offspring.
Female Lepidoptera require, as Mr. Wallace insists,407 some days to deposit their fertilised ova and to search for a proper place; during this period (whilst the life of the male was of no importance) the brighter-coloured females would be exposed to danger and would be liable to be destroyed. The duller-coloured females on the other hand would survive, and thus would influence, it might be thought, in a marked manner the character of the species,—either of both sexes or of one sex, according to which form of inheritance prevailed. But it must not be forgotten that the males emerge from the cocoon-state some days before the females, and during this period, whilst the unborn females were safe, the brighter-coloured males would be exposed to danger; so that ultimately both sexes would probably be exposed during a nearly equal length of time to danger, and the elimination of conspicuous colours would not be much more effective in the one than the other sex.
It is a more important consideration that female Lepidoptera, as Mr. Wallace remarks, and as is known to every collector, are generally slower flyers than the males. Consequently the latter, if exposed to greater danger from being conspicuously coloured, might be able to escape from their enemies, whilst the similarly-coloured females would be destroyed; and thus the females would have the most influence in modifying the colour of their progeny.
There is one other consideration: bright colours, as far as sexual selection is concerned, are commonly of no service to the females; so that if the latter varied in brightness, and the variations were sexually limited in their transmission, it would depend on mere chance whether the females had their bright colours increased; and this would tend throughout the Order to diminish the number of species with brightly-coloured females408 in comparison with the species having brightly-coloured males. On the other hand, as bright colours are supposed to be highly serviceable to the males in their love-struggles, the brighter males (as we shall see in the chapter on Birds) although exposed to rather greater danger, would on an average procreate a greater number of offspring than the duller males. In this case, if the variations were limited in their transmission to the male sex, the males alone would be rendered more brilliantly coloured; but if the variations were not thus limited, the preservation and augmentation of such variations would depend on whether more evil was caused to the species by the females being rendered conspicuous, than good to the males by certain individuals being successful over their rivals.
As there can hardly be a doubt that both sexes of many butterflies and moths have been rendered dull-coloured for the sake of protection, so it may have been with the females alone of some species in which successive variations towards dullness first appeared in the female sex and were from the first limited in their transmission to
the same sex. If not thus limited, both sexes would become dull-coloured. We shall immediately see, when we treat of mimickry, that the females alone of certain butterflies have been rendered extremely beautiful for the sake of protection, without any of the successive protective variations having been transferred to the male, to whom they could not possibly have been in the least degree injurious, and therefore could not have been eliminated through natural selection. Whether in each particular species, in which the sexes differ in colour, it is the female which has been specially modified for the sake of protection; or whether it is the male which has been specially modified for the sake of sexual attraction, the409 female having retained her primordial colouring only slightly changed through the agencies before alluded to; or whether again both sexes have been modified, the female for protection and the male for sexual attraction, can only be definitely decided when we know the life-history of each species.
Without distinct evidence, I am unwilling to admit that a double process of selection has long been going on with a multitude of species,—the males having been rendered more brilliant by beating their rivals; and the females more dull-coloured by having escaped from their enemies. We may take as an instance the common brimstone butterfly (Gonepteryx), which appears early in the spring before any other kind. The male of this species is of a far more intense yellow than the female, though she is almost equally conspicuous; and in this case it does not seem probable that she specially acquired her pale tints as a protection, though it is probable that the male acquired his bright colours as a sexual attraction. The female of Anthocharis cardamines does not possess the beautiful orange tips to her wings with which the male is ornamented; consequently she closely resembles the white butterflies (Pieris) so common in our gardens; but we have no evidence that this resemblance is beneficial. On the contrary, as she resembles both sexes of several species of the same genus inhabiting various quarters of the world, it is more probable that she has simply retained to a large extent her primordial colours.
Various facts support the conclusion that with the greater number of brilliantly-coloured Lepidoptera, it is the male which has been modified; the two sexes having come to differ from each other, or to resemble each other, according to which form of inheritance has prevailed. Inheritance is governed by so many un410known laws or conditions, that they seem to us to be most capricious in their action;524 and we can so far understand how it is that with closely-allied species the sexes of some differ to an astonishing degree, whilst the sexes of others are identical in colour. As the successive steps in the process of variation are necessarily all transmitted through the female, a greater or less number of such steps might readily become developed in her; and thus we can understand the frequent gradations from an extreme difference to no difference at all between the sexes of the species within the same group. These cases of gradation are much too common to favour the supposition that we here see females actually undergoing the process of transition and losing their brightness for the sake of protection; for we have every reason to conclude that at any one time the greater number of species are in a fixed condition. With respect to the differences between the females of the species in the same genus or family, we can perceive that they depend, at least in part, on the females partaking of the colours of their respective males. This is well illustrated in those groups in which the males are ornamented to an extraordinary degree; for the females in these groups generally partake to a certain extent of the splendour of their male partners. Lastly, we continually find, as already remarked, that the females of almost all the species in the same genus, or even family, resemble each other much more closely in colour than do the males; and this indicates that the males have undergone a greater amount of modification than the females.
411Mimickry.—This principle was first made clear in an admirable paper by Mr. Bates,525 who thus threw a flood of light on many obscure problems. It had previously been observed that certain butterflies in S. America belonging to quite distinct families, resembled the Heliconidæ so closely in every stripe and shade of colour that they could not be distinguished except by an experienced entomologist. As the Heliconidæ are coloured in their usual manner, whilst the others depart from the usual colouring of the groups to which they belong, it is clear that the latter are the imitators, and the Heliconidæ the imitated. Mr. Bates further observed that the imitating species are comparatively rare, whilst the imitated swarm in large numbers; the two sets living mingled together. From the fact of the Heliconidæ being conspicuous and beautiful insects, yet so numerous in individuals and species, he concluded that they must be protected from the attacks of birds by some secretion or odour; and this hypothesis has now been confirmed by a considerable body of curious evidence.526 From these considerations Mr. Bates inferred that the butterflies which imitate the protected species had acquired their present marvellously deceptive appearance, through variation and natural selection, in order to be mistaken for the protected kinds and thus to escape being devoured. No explanation is here attempted of the brilliant colours of the imitated, but only of the imitating butterflies. We must account for the colours of the former in the same general manner, as in the cases previously discussed in this chapter. Since the publication of Mr. Bates’ paper, similar and equally striking facts have been observed 412by Mr. Wallace527 in the Malayan region, and by Mr. Trimen in South Africa.
As some writers528 have felt much difficulty in understanding how the first steps in the process of mimickry could have been effected through natural selection, it may be well to remark that the process probably has never commenced with forms widely dissimilar in colour. But with two species moderately like each other, the closest resemblance if beneficial to either form could readily be thus gained; and if the imitated form was subsequently and gradually modified through sexual selection or any other means, the imitating form would be led along the same track, and thus be modified to almost any extent, so that it might ultimately assume an appearance or colouring wholly unlike that of the other members of the group to which it belonged. As extremely slight variations in colour would not in many cases suffice to render a species so like another protected species as to lead to its preservation, it should be remembered that many species of Lepidoptera are liable to considerable and abrupt variations in colour. A few instances have been given in this chapter; but under this point of view Mr. Bates’ original paper on mimickry, as well as Mr. Wallace’s papers, should be consulted.
In the foregoing cases both sexes of the imitating species resemble the imitated; but occasionally the 413female alone mocks a brilliantly-coloured and protected species inhabiting the same district. Consequently the female differs in colour from her own male, and, which is a rare and anomalous circumstance, is the more brightly-coloured of the two. In all the few species of Pieridæ, in which the female is more conspicuously coloured than the male, she imitates, as I am informed by Mr. Wallace, some protected species inhabiting the same region. The female of Diadema anomala is rich purple-brown with almost the whole surface glossed with satiny blue, and she closely imitates the Euplœa midamus, “one of the commonest butterflies of the East;” whilst the male is bronzy or olive-brown, with only a slight blue gloss on the outer parts of the wings.529 Both sexes of this Diadema and of D. bolina follow the same habits of life, so that the differences in colour between the sexes cannot be accounted for by exposure to different conditions;530 even if this explanation were admissible in other instances.531
The above cases of female butterflies which are more brightly-coloured than the males, shew us, firstly, that variations have arisen in a state of nature in the female sex, and have been transmitted exclusively, or almost exclusively, to the same sex; and, secondly, that this form of inheritance has not been determined through natural selection. For if we assume that the females, before they became brightly coloured in imitation of some protected kind, were exposed during each season for a longer period to danger than the males; or if we assume that 414they coul
d not escape so swiftly from their enemies, we can understand how they alone might originally have acquired through natural selection and sexually-limited inheritance their present protective colours. But except on the principle of these variations having been transmitted exclusively to the female offspring, we cannot understand why the males should have remained dull-coloured; for it would surely not have been in any way injurious to each individual male to have partaken by inheritance of the protective colours of the female, and thus to have had a better chance of escaping destruction. In a group in which brilliant colours are so common as with butterflies, it cannot be supposed that the males have been kept dull-coloured through sexual selection by the females rejecting the individuals which were rendered as beautiful as themselves. We may, therefore, conclude that in these cases inheritance by one sex is not due to the modification through natural selection of a tendency to equal inheritance by both sexes.
It may be well here to give an analogous case in another Order, of characters acquired only by the female, though not in the least injurious, as far as we can judge, to the male. Amongst the Phasmidæ, or spectre-insects, Mr. Wallace states that “it is often the females alone that so strikingly resemble leaves, while the males show only a rude approximation.” Now, whatever may be the habits of these insects, it is highly improbable that it could be disadvantageous to the males to escape detection by resembling leaves.532 Hence we may conclude 415that the females alone in this latter as in the previous cases originally varied in certain characters; these characters having been preserved and augmented through ordinary selection for the sake of protection and from the first transmitted to the female offspring alone.
Bright Colours of Caterpillars.—Whilst reflecting on the beauty of many butterflies, it occurred to me that some caterpillars were splendidly coloured, and as sexual selection could not possibly have here acted, it appeared rash to attribute the beauty of the mature insect to this agency, unless the bright colours of their larvæ could be in some manner explained. In the first place it may be observed that the colours of caterpillars do not stand in any close correlation with those of the mature insect. Secondly, their bright colours do not 416serve in any ordinary manner as a protection. As an instance of this, Mr. Bates informs me that the most conspicuous caterpillar which he ever beheld (that of a Sphinx) lived on the large green leaves of a tree on the open llanos of South America; it was about four inches in length, transversely banded with black and yellow, and with its head, legs, and tail of a bright red. Hence it caught the eye of any man who passed by at the distance of many yards, and no doubt of every passing bird.