There is one genus of diurnal butterflies which seems to contradict the law that all the surface that is visible in the resting position exhibits the protective coloration—the South American wood-butterflies of the genus Ageronia. They have on the upper surface a very complicated bark-like pattern of mingled grey on grey, and this confirms the usual rule, for we know that these butterflies—a striking exception among all the other diurnal forms—settle with outspread wings on the trunk of a tree in exactly the same attitude as many of the nocturnal Lepidoptera of the family of the Loopers or Geometridæ, in which the upper surface is also deceptively like the bark of the tree on which they rest.
Fig. 9. Hebomoja glaucippe, from India; under surface. A, in flight. B, in resting attitude.
In all the nocturnal Lepidoptera it is the upper side of the wing which is sympathetically coloured, if protective coloration has been developed at all. In all the Sphingidæ, many 'Owls' and Bombycidæ, the anterior wings are grey banded with darker zigzag lines, and mottled with many shades of black, grey, yellow, red, and even violet. As the anterior wings cover the body and the posterior wings like a roof, they make the resting insect very inconspicuous when it has settled on wooden fences, trunks of trees, or even old timber. When bright colours—red, yellow, or blue—occur in these moths it is always on the posterior wings, which are covered when at rest. This can best be observed in the species of the genus Catocala.
Let us now, however, interrupt our survey of the facts for a moment, and let us inquire whether all the cases of protective colouring in Lepidoptera we have considered can be referred to natural selection, or whether it is not conceivable that other causes may have evoked them.
Fig. 10. Xylina vetusta, after Rösel. A, in flight. B, at rest.
The first thing to be said is that the Lamarckian principle of the inherited effects of use and disuse cannot here be taken into account, as the colours of the surface of the body do not exercise any active function at all; their effect is due simply to their presence, and it is for them quite indifferent whether and how often they have opportunity to protect their bearers from enemies, or whether no enemies ever chance to appear. It has frequently been suggested, too, that these colorations are associated with the differences in the strength of the illumination to which the different parts and surfaces are exposed. But this again is untenable, as is proved even by the dimorphism frequently occurring in caterpillars, for the green and the brown individuals are exposed to precisely the same light; and still more clearly by the sympathetic colouring, which is so exactly defined and yet so different on the under surface of the diurnal butterflies. Yet there are isolated cases in which it seems as if the direct influence of the light had brought about certain striking differences in the colouring of the parts of an insect, and I shall describe perhaps the prettiest of these cases, to which Brunner von Wattenwyl directed attention. It concerns one of the Orthoptera of Australia, a Phasmid, Tropidoderus childreni, Grey, which has a general colouring of leaf-green, but with singular deviations from it on certain areas of the body. In this insect the anterior wings which form the wing covers or elytra ([Fig. 11], V) are so short that they scarcely cover the half of the long abdomen. Their place is taken by the anterior margin of the posterior wing (H. horn), which is hard and horny like the elytra, and in the resting position protects the whole abdomen. All these covering parts are grass-green, except at the places where they overlap; on these areas they have a faded look, and are yellowish instead of green. Brunner says of this: 'The phenomenon gives the impression that the more brilliant colour is a character due to daylight. If several sheets of white paper of unequal dimensions be placed one above the other, ... and exposed to the sun, after a short time silhouettes of the smaller sheets will appear on the larger ones, either in a lighter or in a darker colour. Probably this "fading" of the covered parts in the Phasmid also belongs to this "category of photographs."' This seems convincing, but analogous phenomena in other insects prevent our regarding the pretty comparison with the photograph as a sufficient explanation. If it were a question of a diurnal butterfly, such an assumption would have to be rejected on this ground alone, that the wing colouring is developed in the pupa, and appears perfect and unalterable as soon as the perfect insect emerges. But in the pupa the position of the wings is exactly the reverse of that seen in the resting attitude of a butterfly, that is, the protectively coloured under side of the wing is not turned towards the light but away from it. Moreover, in the pupa the anterior wings cover the posterior ones completely, no matter what the wing position may be later in the perfect insect. Furthermore, the thick and often darkly coloured sheath of the pupa prevents the light having any effect, and not a few species pass their pupal stage in such dark places—for instance, under stones, as in the case of many 'Blues'—that the light can hardly reach them. And if the light did exercise an influence, how could it produce such diverse coloration as the protective colours of diurnal butterflies, on the one side dark, even to blackness, on the other side, yellow, reddish, and even white and pure green; and how should the same rays of light call forth complicated colour patterns on one and the same surface, for instance, the white, sprinkled with green, of the Aurora butterfly (Anthocharis cardaminis)? Finally, we have only to remember that numerous nocturnal Lepidoptera pass through their pupa stage underground, although they exhibit brilliant as well as protective colours in the most appropriate distribution, to reject once for all the hypothesis that the influence of light plays any decisive rôle in determining the distribution of the colours on the wings of Lepidoptera.
But it is otherwise with Tropidoderus. In this case the wings grow gradually during the slow growth of the animal, which takes place in full light, and the wings of the young insect probably lie one above the other, in exactly the same position, and cover the same places as in the full-grown form; we might, therefore, from the facts of the case, admit the possibility that the yellow of the covered portions is due to the exclusion of light.