Contributions to the Theory of Natural Selection. Alfred Russel Wallace

birds—as jacamars, trogons, and puffbirds—which catch insects on the wing, and that they destroy many butterflies is indicated by the fact that the wings of these insects are often found on the ground where their bodies have been devoured. But among these there are no wings of Heliconidæ, while those of the large showy Nymphalidæ, which have a much swifter flight, are often met with. Again, a gentleman who had recently returned from Brazil stated at a meeting of the Entomological Society that he once observed a pair of puffbirds catching butterflies, which they brought to their nest to feed their young; yet during half an hour they never brought one of the Heliconidæ, which were flying lazily about in great numbers, and which they could have captured more easily than any others. It was this circumstance that led Mr. Belt to observe them so long, as he could not understand why the most common insects should be altogether passed by. Mr. Bates also tells us that he never saw them molested by lizards or predacious flies, which often pounce on other butterflies.

      If, therefore, we accept it as highly probable (if not proved) that the Heliconidæ are very greatly protected from attack by their peculiar odour and taste, we find it much more easy to understand their chief characteristics—their great abundance, their slow flight, their gaudy colours, and the entire absence of protective tints on their under surfaces. This property places them somewhat in the position of those curious wingless birds of oceanic islands, the dodo, the apteryx, and the moas, which are with great reason supposed to have lost the power of flight on account of the absence of carnivorous quadrupeds. Our butterflies have been protected in a different way, but quite as effectually; and the result has been that as there has been nothing to escape from, there has been no weeding out of slow flyers, and as there has been nothing to hide from, there has been no extermination of the bright-coloured varieties, and no preservation of such as tended to assimilate with surrounding objects.

      Now let us consider how this kind of protection must act. 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, 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 the Heliconias. If the birds could not distinguish the two kinds externally, and there were on the average only one eatable among fifty uneatable, they would soon give up seeking for the eatable ones, even if they knew them to exist. If, on the other hand, any particular butterfly of an eatable group acquired the disagreeable taste of the Heliconias while it retained the characteristic form and colouring of its own group, this would be really of no use to it whatever; for the birds would go on catching it among its eatable allies (compared with which it would rarely occur), it would be wounded and disabled, even if rejected, and its increase would thus be as effectually checked as if it were devoured. It is important, therefore, to understand that if any one genus of an extensive family of eatable butterflies were in danger of extermination from insect-eating birds, and if two kinds of variation were going on among them, some individuals possessing a slightly disagreeable taste, others a slight resemblance to the Heliconidæ, this latter quality would be much more valuable than the former. The change in flavour would not at all prevent the variety from being captured as before, and it would almost certainly be thoroughly disabled before being rejected. The approach in colour and form to the Heliconidæ, however, would be at the very 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.

      Now, this hypothetical case is exactly realized in South America. Among the white butterflies forming the family Pieridæ (many of which do not greatly differ in appearance from our own cabbage butterflies) is a genus of rather small size (Leptalis), some species of which are white like their allies, while the larger number exactly resemble the Heliconidæ in the form and colouring of the wings. It must always be remembered that these two families are as absolutely distinguished from each other by structural characters as are the carnivora and the ruminants among quadrupeds, and that an entomologist can always distinguish the one from the other by the structure of the feet, just as certainly as a zoologist can tell a bear from a buffalo by the skull or by a tooth. Yet the resemblance of a species of the one family to another species in the other family was often so great, that both Mr. Bates and myself were many times deceived at the time of capture, and did not discover the distinctness of the two insects till a closer examination detected their essential differences. During his residence of eleven years in the Amazon valley, Mr. Bates found a number of species or varieties of Leptalis, each of which was a more or less exact copy of one of the Heliconidæ of the district it inhabited; and the results of his observations are embodied in a paper published in the Linnean Transactions, in which he first explained the phenomena of “mimicry” as the result of natural selection, and showed its identity in cause and purpose with protective resemblance to vegetable or inorganic forms.

      The imitation of the Heliconidæ by the Leptalides is carried out to a wonderful degree in form as well as in colouring. The wings have become elongated to the same extent, and the antennæ and abdomen have both become lengthened, to correspond with the unusual condition in which they exist in the former family. In colouration there are several types in the different genera of Heliconidæ. The genus Mechanitis is generally of a rich semi-transparent brown, banded with black and yellow; Methona is of large size, the wings transparent like horn, and with black transverse bands; while the delicate Ithomias are all more or less transparent, with black veins and borders, and often with marginal and transverse bands of orange red. These different forms are all copied by the various species of Leptalis, every band and spot and tint of colour, and the various degrees of transparency, being exactly reproduced. As if to derive all the benefit possible from this protective mimicry, the habits have become so modified that the Leptalides generally frequent the very same spots as their models, and have the same mode of flight; and as they are always very scarce (Mr. Bates estimating their numbers at about one to a thousand of the group they resemble), there is hardly a possibility of their being found out by their enemies. It is also very remarkable that in almost every case the particular Ithomias and other species of Heliconidæ which they resemble, are noted as being very common species, swarming in individuals, and found over a wide range of country. This indicates antiquity and permanence in the species, and is exactly the condition most essential both to aid in the development of the resemblance, and to increase its utility.

      But the Leptalides are not the only insects who have prolonged their existence by imitating the great protected group of Heliconidæ;—a genus of quite another family of most lovely small American butterflies, the Erycinidæ, and three genera of diurnal moths, also present species which often mimic the same dominant forms, so that some, as Ithomia ilerdina of St. Paulo, for instance, have flying with them a few individuals of three widely different insects, which are yet disguised with exactly the same form, colour, and markings, so as to be quite undistinguishable when upon the wing. Again, the Heliconidæ are not the only group that are imitated, although they are the most frequent models. The black and red group of South American Papilios, and the handsome Erycinian genus Stalachtis, have also a few who copy them; but this fact offers no difficulty, since these two groups are almost as dominant as the Heliconidæ. They both fly very slowly, they are both conspicuously coloured, and they both abound in individuals; so that there is every reason to believe that they possess a protection of a similar kind to the Heliconidæ, and that it is therefore equally an advantage to other insects to be mistaken for them. There is also another extraordinary fact that we are not yet in a position clearly to comprehend: some groups of the Heliconidæ themselves mimic other groups. Species