In explaining the mode of action of this protecting resemblance Mr. Wallace observes:[[25]] "Tropical insectivorous birds very frequently sit on dead branches of a lofty tree, or on those which overhang forest paths, gazing intently around, and darting off at intervals to seize an insect at a considerable distance, with which they generally return to their station to devour. If a bird began by capturing the slow-flying conspicuous Heliconidæ, and found them always so disagreeable that it could not eat them, it would after a very few trials leave off catching them at all; and their whole appearance, form, colouring, and mode of flight is so peculiar, that there can be little doubt birds would soon learn to distinguish them at a long distance, and never waste any time in pursuit of them. Under these circumstances, it is evident that any other butterfly of a group which birds were accustomed to devour, would be almost equally well protected by closely resembling a Heliconia externally, as if it acquired also the disagreeable odour; always supposing that there were only a few of them among a great number of Heliconias."
"The approach in colour and form to the Heliconidæ, however, would be at the first a positive, though perhaps a slight, advantage; for although at short distances this variety would be easily distinguished and devoured, yet at a longer distance it might be mistaken for one of the uneatable group, and so be passed by and gain another day's life, which might in many cases be sufficient for it to lay a quantity of eggs and leave a numerous progeny, many of which would inherit the peculiarity which had been the safeguard of their parent."
As a complete example of mimicry Mr. Wallace refers to a common Indian butterfly. He says:[[26]] "But the most wonderful and undoubted case of protective resemblance in a butterfly,
which I have ever seen, is that of the common Indian Kallima inachis, and its Malayan ally, Kallima paralekta. The upper surface of these is very striking and showy, as they are of a
large size, and are adorned with a broad band of rich orange on a deep bluish ground. The under side is very variable in colour, so that out of fifty specimens no two can be found exactly alike, but every one of them will be of some shade of ash, or brown, or ochre, such as are found among dead, dry, or decaying leaves. The apex of the upper wings is produced into an acute point, a very common form in the leaves of tropical shrubs and trees, and the lower wings are also produced into a short narrow tail. Between these two points runs a dark curved line exactly representing the midrib of a leaf, and from this radiate on each side a few oblique lines, which serve to indicate the lateral veins of a leaf. These marks are more clearly seen on the outer portion of the base of the wings, and on the inner side towards the middle and apex, and it is very curious to observe how the usual marginal and transverse striæ of the group are here modified and strengthened so as to become adapted for an imitation of the venation of a leaf." ... "But this resemblance, close as it is, would be of little use if the habits of the insect did not accord with it. If the butterfly sat upon leaves or upon flowers, or opened its wings so as to expose the upper surface, or exposed and moved its head and antennæ as many other butterflies do, its disguise would be of little avail. We might be sure, however, from the analogy of many other cases, that the habits of the insect are such as still further to aid its deceptive garb; but we are not obliged to make any such supposition, since I myself had the good fortune to observe scores of Kallima paralekta, in Sumatra, and to capture many of them, and can vouch for the accuracy of the following details. These butterflies frequent dry forests, and fly very swiftly. They were seen to settle on a flower or a green leaf, but were many times lost sight of in a bush or tree of dead leaves. On such occasions they were generally searched for in vain, for while gazing intently at the very spot where one had disappeared, it would often suddenly dart out, and again vanish twenty or fifty yards
further on. On one or two occasions the insect was detected reposing, and it could then be seen how completely it assimilates itself to the surrounding leaves. It sits on a nearly upright twig, the wings fitting closely back to back, concealing the antennæ and head, which are drawn up between their bases. The little tails of the hind wing touch the branch, and form a perfect stalk to the leaf, which is supported in its place by the claws of the middle pair of feet, which are slender and inconspicuous. The irregular outline of the wings gives exactly the perspective effect of a shrivelled leaf. We thus have size, colour, form, markings, and habits, all combining together to produce a disguise which may be said to be absolutely perfect; and the protection which it affords is sufficiently indicated by the abundance of the individuals that possess it."
Beetles also imitate bees and wasps, as do some Lepidoptera; and objects the most bizarre and unexpected are simulated, such as dung and drops of dew. Some insects, called bamboo and walking-stick insects, have a most remarkable resemblance to pieces of bamboo, to twigs and branches. Of these latter insects Mr. Wallace says:[[27]] "Some of these are a foot long and as thick as one's finger, and their whole colouring, form, rugosity, and the arrangement of the head, legs, and antennæ, are such as to render them absolutely identical in appearance with dry sticks. They hang loosely about shrubs in the forest, and have the extraordinary habit of stretching out their legs unsymmetrically, so as to render the deception more complete." Now let us suppose that the ancestors of these various animals were all destitute of the very special protections they at present possess, as on the Darwinian hypothesis we must do. Let it also be conceded that small deviations from the antecedent colouring or form would tend to make some of their ancestors escape destruction by causing them more or less frequently to
be passed over, or mistaken by their persecutors. Yet the deviation must, as the event has shown, in each case be in some definite direction, whether it be towards some other animal or plant, or towards some dead or inorganic matter. But as, according to Mr. Darwin's theory, there is a constant tendency to indefinite variation, and as the minute incipient variations will be in all directions, they must tend to neutralize each other, and at first to form such unstable modifications that it is difficult, if not impossible, to see how such indefinite oscillations of infinitesimal beginnings can ever build up a sufficiently appreciable resemblance to a leaf, bamboo, or other object, for "Natural Selection" to seize upon and perpetuate. This difficulty is augmented when we consider—a point to be dwelt upon hereafter—how necessary it is that many individuals should be similarly modified simultaneously. This has been insisted on in an able article in the North British Review for June 1867, p. 286, and the consideration of the article has occasioned Mr. Darwin to make an important modification in his views.[[28]]
In these cases of mimicry it seems difficult indeed to imagine a reason why variations tending in an infinitesimal degree in any special direction should be preserved. All variations would be preserved which tended to obscure the perception of an animal by its enemies, whatever direction those variations might take, and the common preservation of conflicting tendencies would greatly favour their mutual neutralization and obliteration if we may rely on the many cases recently brought forward by Mr. Darwin with regard to domestic animals.