With animals of all kinds, whenever colour has been modified for some special purpose, this has been, as far 392as we can judge, either for protection or as an attraction between the sexes. With many species of butterflies the upper surfaces of the wings are obscurely coloured, and this in all probability leads to their escaping observation and danger. But butterflies when at rest would be particularly liable to be attacked by their enemies; and almost all the kinds when resting raise their wings vertically over their backs, so that the lower sides alone are exposed to view. Hence it is this side which in many cases is obviously coloured so as to imitate the surfaces on which these insects commonly rest. Dr. Rössler, I believe, first noticed the similarity of the closed wings of certain Vanessæ and other butterflies to the bark of trees. Many analogous and striking facts could be given. The most interesting one is that recorded by Mr. Wallace506 of a common Indian and Sumatran butterfly (Kallima), which disappears like magic when it settles in a bush; for it hides its head and antennæ between its closed wings, and these in form, colour, and veining cannot be distinguished from a withered leaf together with the footstalk. In some other cases the lower surfaces of the wings are brilliantly coloured, and yet are protective; thus in Thecla rubi the wings when closed are of an emerald green and resemble the young leaves of the bramble, on which this butterfly in the spring may often be seen seated.
Although the obscure tints of the upper or under surface of many butterflies no doubt serve to conceal them, yet we cannot possibly extend this view to the brilliant and conspicuous colours of many kinds, such as our admiral and peacock Vanessæ, our white 393cabbage-butterflies (Pieris), or the great swallow-tail Papilio which haunts the open fens—for these butterflies are thus rendered visible to every living creature. With these species both sexes are alike; but in the common brimstone butterfly (Gonepteryx rhamni), the male is of an intense yellow, whilst the female is much paler; and in the orange-tip (Anthocharis cardamines) the males alone have the bright orange tips to their wings. In these cases the males and females are equally conspicuous, and it is not credible that their difference in colour stands in any relation to ordinary protection. Nevertheless it is possible that the conspicuous colours of many species may be in an indirect manner beneficial, as will hereafter be explained, by leading their enemies at once to recognise them as unpalatable. Even in this case it does not certainly follow that their bright colours and beautiful patterns were acquired for this special purpose. In some other remarkable cases, beauty has been gained for the sake of protection, through the imitation of other beautiful species, which inhabit the same district and enjoy an immunity from attack by being in some way offensive to their enemies.
The female of our orange-tip butterfly, above referred to, and of an American species (Anth. genutia) probably shew us, as Mr. Walsh has remarked to me, the primordial colours of the parent-species of the genus; for both sexes of four or five widely-distributed species are coloured in nearly the same manner. We may infer here, as in several previous cases, that it is the males of Anth. cardamines and genutia which have departed from the usual type of colouring of their genus. In the Anth. sara from California, the orange-tips have become partially developed in the female; for her wings are tipped with reddish-orange, but paler than in the394 male, and slightly different in some other respects. In an allied Indian form, the Iphias glaucippe, the orange-tips are fully developed in both sexes. In this Iphias the under surface of the wings marvellously resembles, as pointed out to me by Mr. A. Butler, a pale-coloured leaf; and in our English orange-tip, the under surface resembles the flower-head of the wild parsley, on which it may be seen going to rest at night.507 The same reasoning power which compels us to believe that the lower surfaces have here been coloured for the sake of protection, leads us to deny that the wings have been tipped, especially when this character is confined to the males, with bright orange for the same purpose.
Turning now to Moths: most of these rest motionless with their wings depressed during the whole or greater part of the day; and the upper surfaces of their wings are often shaded and coloured in an admirable manner, as Mr. Wallace has remarked, for escaping detection. With most of the Bombycidæ and Noctuidæ,508 when at rest, the front-wings overlap and conceal the hind-wings; so that the latter might be brightly coloured without much risk; and they are thus coloured in many species of both families. During the act of flight, moths would often be able to escape from their enemies; nevertheless, as the hind-wings are then fully exposed to view, their bright colours must generally have been acquired at the cost of some little risk. But the following fact shews us how cautious we ought to be in drawing conclusions on this head. The common yellow under-wings 395(Triphaena) often fly about during the day or early evening, and are then conspicuous from the colour of their hind-wings. It would naturally be thought that this would be a source of danger; but Mr. J. Jenner Weir believes that it actually serves them as a means of escape, for birds strike at these brightly coloured and fragile surfaces, instead of at the body. For instance, Mr. Weir turned into his aviary a vigorous specimen of Triphaena pronuba, which was instantly pursued by a robin; but the bird’s attention being caught by the coloured wings, the moth was not captured until after about fifty attempts, and small portions of the wings were repeatedly broken off. He tried the same experiment, in the open air, with a T. fimbria and swallow; but the large size of this moth probably interfered with its capture.509 We are thus reminded of a statement made by Mr. Wallace,510 namely, that in the Brazilian forests and Malayan islands, many common and highly-decorated butterflies are weak flyers, though furnished with a broad expanse of wings; and they “are often captured with pierced and broken wings, as if they had been seized by birds, from which they had escaped: if the wings had been much smaller in proportion to the body, it seems probable that the insect would more frequently have been struck or pierced in a vital part, and thus the increased expanse of the wings may have been indirectly beneficial.”
Display.—The bright colours of butterflies and of some moths are specially arranged for display, whether or not they serve in addition as a protection. Bright 396colours would not be visible during the night; and there can be no doubt that moths, taken as a body, are much less gaily decorated than butterflies, all of which are diurnal in their habits. But the moths in certain families, such as the Zygænidæ, various Sphingidæ, Uraniidæ, some Arctiidæ and Saturniidæ, fly about during the day or early evening, and many of these are extremely beautiful, being far more brightly coloured than the strictly nocturnal kinds. A few exceptional cases, however, of brightly-coloured nocturnal species have been recorded.511
There is evidence of another kind in regard to display. Butterflies, as before remarked, elevate their wings when at rest, and whilst basking in the sunshine often alternately raise and depress them, thus exposing to full view both surfaces; and although the lower surface is often coloured in an obscure manner as a protection, yet in many species it is as highly coloured as the upper surface, and sometimes in a very different manner. In some tropical species the lower surface is even more brilliantly coloured than the upper.512 In one English fritillary, the Argynnis aglaia, the lower surface alone is ornamented with shining silver discs. Nevertheless, as a general rule, the upper surface, which is probably the most fully exposed, is coloured more brightly and in a more diversified manner than the lower. Hence the lower surface generally affords 397to entomologists the most useful character for detecting the affinities of the various species.
Now if we turn to the enormous group of moths, which do not habitually expose to full view the under surface of their wings, this side is very rarely, as I hear from Mr. Stainton, coloured more brightly than the upper side, or even with equal brightness. Some exceptions to the rule, either real or apparent, must be noticed, as that of Hypopira, specified by Mr. Wormald.513 Mr. R. Trimen informs me that in Guenée’s great work, three moths are figured, in which the under surface is much the most brilliant. For instance, in the Aust
ralian Gastrophora the upper surface of the fore-wing is pale greyish-ochreous, while the lower surface is magnificently ornamented by an ocellus of cobalt-blue, placed in the midst of a black mark, surrounded by orange-yellow, and this by bluish-white. But the habits of these three moths are unknown; so that no explanation can be given of their unusual style of colouring. Mr. Trimen also informs me that the lower surface of the wings in certain other Geometræ514 and quadrifid Noctuæ are either more variegated or more brightly-coloured than the upper surface; but some of these species have the habit of “holding their wings quite erect over their backs, retaining them in this position for a considerable time,” and thus exposing to view the under surface. Other species when settled on the ground or herbage have the habit of now and then suddenly and slightly lifting up their wings. Hence the lower surface of the wings being more brightly-coloured than the upper sur398face in certain moths is not so anomalous a circumstance as it at first appears. The Saturniidæ include some of the most beautiful of all moths, their wings being decorated, as in our British Emperor moth, with fine ocelli; and Mr. T. W. Wood515 observes that they resemble butterflies in some of their movements; “for instance, in the gentle waving up and down of the wings, as if for display, which is more characteristic of diurnal than of nocturnal Lepidoptera.”
It is a singular fact that no British moths, nor as far as I can discover hardly any foreign species, which are brilliantly coloured, differ much in colour according to sex; though this is the case with many brilliant butterflies. The male, however, of one American moth, the Saturnia Io, is described as having its fore-wings deep yellow, curiously marked with purplish-red spots; whilst the wings of the female are purple-brown, marked with grey lines.516 The British moths which differ sexually in colour are all brown, or various tints of dull yellow, or nearly white. In several species the males are much darker than the females,517 and these belong to groups which generally fly about during the afternoon. On the other hand, in many genera, as Mr. Stainton informs me, 399the males have the hind-wings whiter than those of the female—of which, fact Agrotis exclamationis offers a good instance. The males are thus rendered more conspicuous than the females, whilst flying about in the dusk. In the Ghost Moth (Hepialus humuli) the difference is more strongly marked; the males being white and the females yellow with darker markings. It is difficult to conjecture what the meaning can be of these differences between the sexes in the shades of darkness or lightness; but we can hardly suppose that they are the result of mere variability with sexually-limited inheritance, independently of any benefit thus derived.
From the foregoing statements 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 exhibited as if for display. Hence I am led to suppose that the females generally prefer, or are most excited by the more brilliant males; for on any other supposition the males would be ornamented, as far as we can see, for no purpose. We know that ants and certain lamellicorn beetles are capable of feeling an attachment for each other, and that ants recognise their fellows after an interval of several months. Hence there is no abstract improbability in the Lepidoptera, which probably stand nearly or quite as high in the scale as these insects, having sufficient mental capacity to admire bright colours. They certainly discover flowers by colour, and, as I have elsewhere shewn, the plants which are fertilised exclusively by the wind never have a conspicuously-coloured corolla. The Humming-bird Sphinx may often be seen to swoop down from a distance on a bunch of flowers in the midst of green foliage;400 and I have been assured by a friend, that these moths repeatedly visited flowers painted on the walls of a room in the South of France. The common white butterfly, as I hear from Mr. Doubleday, often flies down to a bit of paper on the ground, no doubt mistaking it for one of its own species. Mr. Collingwood518 in speaking of the difficulty of collecting certain butterflies in the Malay Archipelago, states that “a dead specimen pinned upon a conspicuous twig will often arrest an insect of the same species in its headlong flight, and bring it down within easy reach of the net, especially if it be of the opposite sex.”
The courtship of butterflies is a prolonged affair. The males sometimes fight together in rivalry; and many may be seen pursuing or crowding round the same female. If, then, the females do not prefer one male to another, the pairing must be left to mere chance, and this does not appear to me a probable event. If, on the other hand, the females habitually, or even occasionally, prefer the more beautiful males, the colours of the latter will have been rendered brighter by degrees, and will have been transmitted to both sexes or to one sex, according to which law of inheritance prevailed. The process of sexual selection will have been much facilitated, if the conclusions arrived at from various kinds of evidence in the supplement to the ninth chapter can be trusted; namely that the males of many Lepidoptera, at least in the imago state, greatly exceed in number the females.
Some facts, however, are opposed to the belief that female butterflies prefer the more beautiful males; thus, as I have been assured by several observers, fresh females may frequently be seen paired with battered, faded or 401dingy males; but this is a circumstance which could hardly fail often to follow from the males emerging from their cocoons earlier than the females. With moths of the family of the Bombycidæ, the sexes pair immediately after assuming the imago state; for they cannot feed, owing to the rudimentary condition of their mouths. The females, as several entomologists have remarked to me, lie in an almost torpid state, and appear not to evince the least choice in regard to their partners, This is the case with the common silk-moth (B. mori), as I have been told by some continental and English breeders. Dr. Wallace, who has had such immense experience in breeding Bombyx cynthia, is convinced that the females evince no choice or preference. He has kept above 300 of these moths living together, and has often found the most vigorous females mated with stunted males. The reverse apparently seldom occurs; for, as he believes, the more vigorous males pass over the weakly females, being attracted by those endowed with most vitality. Although we have been indirectly induced to believe that the females of many species prefer the more beautiful males, I have no reason to suspect, either with moths or butterflies, that the males are attracted by the beauty of the females. If the more beautiful females had been continually preferred, it is almost certain, from the colours of butterflies being so frequently transmitted to one sex alone, that the females would often have been rendered more beautiful than their male partners. But this does not occur except in a few instances; and these can be explained, as we shall presently see, on the principle of mimickry and protection.
As sexual selection primarily depends on variability, a few words must be added on this subject. In respect402 to colour there is no difficulty, as any number of highly variable Lepidoptera could be named. One good instance will suffice. Mr. Bates shewed me a whole series of specimens of Papilio sesostris and childrenæ; in the latter the males varied much in the extent of the beautifully enamelled green patch on the fore-wings, and in the size of the white mark, as well as of the splendid crimson stripe on the hind-wings; so that there was a great contrast between the most and least gaudy males. The male of Papilio sesostris, though a beautiful insect, is much less so than P. childrenæ. It likewise varies a little in the size of the green patch on the fore-wings, and in the occasional appearance of a small crimson stripe on the hind-wings, borrowed, as it would seem, from its own female; for the females of this and of many other species in the Æneas group possess this crimson stripe. Hence between the brightest specimens of P. sesostris and the least bright of P. childrenæ, there was but a small interval; and it was evident that as far as mere variability is concerned, there would be no difficulty in permanently increasing by means of selection the beauty of either species. The variability is here almost confined to the male sex; but Mr. Wallace and Mr. Bates have shewn519 that the females of some other species
are extremely variable, the males being nearly constant. As I have before mentioned the Ghost Moth (Hepialus humuli) as one of the best instances in Britain of a difference in colour between the sexes of moths, it may be worth adding520 that in the Shetland 403Islands, males are frequently found which closely resemble the females. In a future chapter I shall have occasion to shew that the beautiful eye-like spots or ocelli, so common on the wings of many Lepidoptera, are eminently variable.
On the whole, although many serious objections may be urged, it seems probable that most of the species of Lepidoptera which are brilliantly coloured, owe their colours to sexual selection, excepting in certain cases, presently to be mentioned, in which conspicuous colours are beneficial as a protection. From the ardour of the male throughout the animal kingdom, he is generally willing to accept any female; and it is the female which usually exerts a choice. Hence if sexual selection has here acted, the male, when the sexes differ, ought to be the most brilliantly coloured; and this undoubtedly is the ordinary rule. When the sexes are brilliantly coloured and resemble each other, the characters acquired by the males appear to have been transmitted to both sexes. But will this explanation of the similarity and dissimilarity in colour between the sexes suffice?