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Display

The bright colours of many butterflies and of some moths are specially arranged for display, so that they may be readily seen. During the night colours are not visible, and there can be no doubt that the nocturnal moths, taken as a body, are much less gaily decorated than butterflies, all of which are diurnal in their habits. But the moths of certain families, such as the Zygaenidae, several Sphingidae, Uraniidae, some Arctiidae and Saturniidae, fly about during the day or early evening, and many of these are extremely beautiful, being far brighter coloured than the strictly nocturnal kinds. A few exceptional cases, however, of bright-coloured nocturnal species have been recorded. (14. For instance, Lithosia; but Prof. Westwood (‘Modern Class. of Insects,’ vol. ii. p. 390) seems surprised at this case. On the relative colours of diurnal and nocturnal Lepidoptera, see ibid. pp. 333 and 392; also Harris, ‘Treatise on the Insects of New England,’ 1842, p. 315.)

There is evidence of another kind in regard to display. Butterflies, as before remarked, elevate their wings when at rest, but whilst basking in the sunshine often alternately raise and depress them, thus exposing both surfaces to full view; and although the lower surface is often coloured in an obscure manner as a protection, yet in many species it is as highly decorated 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. (15. Such differences between the upper and lower surfaces of the wings of several species of Papilio may be seen in the beautiful plates to Mr. Wallace’s ‘Memoir on the Papilionidae of the Malayan Region,’ in ‘Transactions of the Linnean Society,’ vol. xxv. part i. 1865.) In the English fritillaries (Argynnis) the lower surface alone is ornamented with shining silver. Nevertheless, as a general rule, the upper surface, which is probably more fully exposed, is coloured more brightly and diversely than the lower. Hence the lower surface generally affords to entomologists the more useful character for detecting the affinities of the various species. Fritz Muller informs me that three species of Castnia are found near his house in S. Brazil: of two of them the hind-wings are obscure, and are always covered by the front-wings when these butterflies are at rest; but the third species has black hind-wings, beautifully spotted with red and white, and these are fully expanded and displayed whenever the butterfly rests. Other such cases could be added.

If we now turn to the enormous group of moths, which, as I hear from Mr. Stainton, do not habitually expose the under surface of their wings to full view, we find this side very rarely coloured with a brightness greater than, or even equal to, that of the upper side. Some exceptions to the rule, either real or apparent, must be noticed, as the case of Hypopyra. (16. See Mr. Wormald on this moth: ‘Proceedings of the Entomological Society,’ March 2, 1868.) Mr. Trimen informs me that in Guenee’s great work, three moths are figured, in which the under surface is much the more brilliant. For instance, in the Australian 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 Geometrae (17. See also an account of the S. American genus Erateina (one of the Geometrae) in ‘Transactions, Ent. Soc.’ new series, vol. v. pl. xv. and xvi.) and quadrifid Noctuae 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 the under surface to view. Other species, when settled on the ground or herbage, now and then suddenly and slightly lift up their wings. Hence the lower surface of the wings being brighter than the upper surface in certain moths is not so anomalous as it at first appears. The Saturniidae 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. Wood (18. ‘Proc Ent. Soc. of London,’ July 6, 1868, p. xxvii.) 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 which are brilliantly coloured, and, as far as I can discover, hardly any foreign species, 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. (19. Harris, ‘Treatise,’ etc., edited by Flint, 1862, p. 395.) The British moths which differ sexually in colour are all brown, or of various dull yellow tints, or nearly white. In several species the males are much darker than the females (20. For instance, I observe in my son’s cabinet that the males are darker than the females in the Lasiocampa quercus, Odonestis potatoria, Hypogymna dispar, Dasychira pudibunda, and Cycnia mendica. In this latter species the difference in colour between the two sexes is strongly marked; and Mr. Wallace informs me that we here have, as he believes, an instance of protective mimicry confined to one sex, as will hereafter be more fully explained. The white female of the Cycnia resembles the very common Spilosoma menthrasti, both sexes of which are white; and Mr. Stainton observed that this latter moth was rejected with utter disgust by a whole brood of young turkeys, which were fond of eating other moths; so that if the Cycnia was commonly mistaken by British birds for the Spilosoma, it would escape being devoured, and its white deceptive colour would thus be highly beneficial.), and these belong to groups which generally fly about during the afternoon. On the other hand, in many genera, as Mr. Stainton informs me, the males have the hind-wings whiter than those of the female–of which fact Agrotis exclamationis offers a good instance. In the Ghost Moth (Hepialus humuli) the difference is more strongly marked; the males being white, and the females yellow with darker markings. (21. It is remarkable, that in the Shetland Islands the male of this moth, instead of differing widely from the female, frequently resembles her closely in colour (see Mr. MacLachlan, ‘Transactions, Entomological Society,’ vol. ii. 1866, p. 459). Mr. G. Fraser suggests (‘Nature,’ April 1871, p. 489) that at the season of the year when the ghost-moth appears in these northern islands, the whiteness of the males would not be needed to render them visible to the females in the twilight night.) It is probable that in these cases the males are thus rendered more conspicuous, and more easily seen by the females whilst flying about in the dusk.

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 exhibited as if for display. Hence 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. 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. 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; and I have been assured by two persons abroad, that these moths repeatedly visit flowers painted on the walls of a room, and vainly endeavour to insert their proboscis into them. Fritz Muller informs me that several kinds of butterflies in S. Brazil shew an unmistakable preference for certain colours over others: he observed that they very often visited the brilliant red flowers of five or six genera of plants, but never the white or yellow flowering species of the same and other genera, growing in the same garden; and I have received other accounts to the same effect. As I hear from Mr. Doubleday, the common white butterfly often flies down to a bit of paper on the ground, no doubt mistaking it for one of its own species. Mr. Collingwood (22. ‘Rambles of a Naturalist in the Chinese Seas,’ 1868, p. 182.) in speaking of the difficulty in 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, as before remarked, a prolonged affair. The males sometimes fight together in rivalry; and many may be seen pursuing or crowding round the same female. Unless, then, the females prefer one male to another, the pairing must be left to mere chance, and this does not appear probable. If, on the other band, 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 the law of inheritance which has prevailed. The process of sexual selection will have been much facilitated, if the conclusion can be trusted, arrived at from various kinds of evidence in the supplement to the ninth chapter; namely, that the males of many Lepidoptera, at least in the imago state, greatly exceed the females in number.

Some facts, however, are opposed to the belief that female butterflies prefer the more beautiful males; thus, as I have been assured by several collectors, fresh females may frequently be seen paired with battered, faded, or dingy 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 Bombycidae, 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 great experience in breeding Bombyx cynthia, is convinced that the females evince no choice or preference. He has kept above 300 of these moths together, and has often found the most vigorous females mated with stunted males. The reverse appears to occur seldom; for, as he believes, the more vigorous males pass over the weakly females, and are attracted by those endowed with most vitality. Nevertheless, the Bombycidae, though obscurely-coloured, are often beautiful to our eyes from their elegant and mottled shades.

I have as yet only referred to the species in which the males are brighter coloured than the females, and I have attributed their beauty to the females for many generations having chosen and paired with the more attractive males. But converse cases occur, though rarely, in which the females are more brilliant than the males; and here, as I believe, the males have selected the more beautiful females, and have thus slowly added to their beauty. We do not know why in various classes of animals the males of some few species have selected the more beautiful females instead of having gladly accepted any female, as seems to be the general rule in the animal kingdom: but if, contrary to what generally occurs with the Lepidoptera, the females were much more numerous than the males, the latter would be likely to pick out the more beautiful females. Mr. Butler shewed me several species of Callidryas in the British Museum, in some of which the females equalled, and in others greatly surpassed the males in beauty; for the females alone have the borders of their wings suffused with crimson and orange, and spotted with black. The plainer males of these species closely resemble each other, shewing that here the females have been modified; whereas in those cases, where the males are the more ornate, it is these which have been modified, the females remaining closely alike.