Contributions to the Theory of Natural Selection Part 4

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_Mimicry._

It has been long known to entomologists that certain insects bear a strange external resemblance to others belonging to distinct genera, families, or even orders, and with which they have no real affinity whatever. The fact, however, appears to have been generally considered as dependent upon some unknown law of "a.n.a.logy"--some "system of nature," or "general plan," which had guided the Creator in designing the myriads of insect forms, and which we could never hope to understand. In only one case does it appear that the resemblance was thought to be useful, and to have been designed as a means to a definite and intelligible purpose. The flies of the genus Volucella enter the nests of bees to deposit their eggs, so that their larvae may feed upon the larvae of the bees, and these flies are each wonderfully like the bee on which it is parasitic. Kirby and Spence believed that this resemblance or "mimicry" was for the express purpose of protecting the flies from the attacks of the bees, and the connection is so evident that it was hardly possible to avoid this conclusion. The resemblance, however, of moths to b.u.t.terflies or to bees, of beetles to wasps, and of locusts to beetles, has been many times noticed by eminent writers; but scarcely ever till within the last few years does it appear to have been considered that these resemblances had any special purpose, or were of any direct benefit to the insects themselves. In this respect they were looked upon as accidental, as instances of the "curious a.n.a.logies" in nature which must be wondered at but which could not be explained.

Recently, however, these instances have been greatly multiplied; the nature of the resemblances has been more carefully studied, and it has been found that they are often carried out into such details as almost to imply a purpose of deceiving the observer. The phenomena, moreover, have been shown to follow certain definite laws, which again all indicate their dependence on the more general law of the "survival of the fittest," or "the preservation of favoured races in the struggle for life." It will, perhaps, be as well here to state what these laws or general conclusions are, and then to give some account of the facts which support them.

The first law is, that in an overwhelming majority of cases of mimicry, the animals (or the groups) which resemble each other inhabit the same country, the same district, and in most cases are to be found together on the very same spot.

The second law is, that these resemblances are not indiscriminate, but are limited to certain groups, which in every case are abundant in species and individuals, and can often be ascertained to have some special protection.

The third law is, that the species which resemble or "mimic" these dominant groups, are comparatively less abundant in individuals, and are often very rare.

These laws will be found to hold good, in all the cases of true mimicry among various cla.s.ses of animals to which we have now to call the attention of our readers.

_Mimicry among Lepidoptera._

As it is among b.u.t.terflies that instances of mimicry are most numerous and most striking, an account of some of the more prominent examples in this group will first be given. There is in South America an extensive family of these insects, the Heliconidae, which are in many respects very remarkable. They are so abundant and characteristic in all the woody portions of the American tropics, that in almost every locality they will be seen more frequently than any other b.u.t.terflies. They are distinguished by very elongate wings, body, and antennae, and are exceedingly beautiful and varied in their colours; spots and patches of yellow, red, or pure white upon a black, blue, or brown ground, being most general. They frequent the forests chiefly, and all fly slowly and weakly; yet although they are so conspicuous, and could certainly be caught by insectivorous birds more easily than almost any other insects, their great abundance all over the wide region they inhabit shows that they are not so persecuted. It is to be especially remarked also, that they possess no adaptive colouring to protect them during repose, for the under side of their wings presents the same, or at least an equally conspicuous colouring as the upper side; and they may be observed after sunset suspended at the end of twigs and leaves where they have taken up their station for the night, fully exposed to the attacks of enemies if they have any. These beautiful insects possess, however, a strong pungent semi-aromatic or medicinal odour, which seems to pervade all the juices of their system. When the entomologist squeezes the breast of one of them between his fingers to kill it, a yellow liquid exudes which stains the skin, and the smell of which can only be got rid of by time and repeated was.h.i.+ngs. Here we have probably the cause of their immunity from attack, since there is a great deal of evidence to show that certain insects are so disgusting to birds that they will under no circ.u.mstances touch them. Mr. Stainton has observed that a brood of young turkeys greedily devoured all the worthless moths he had ama.s.sed in a night's "sugaring," yet one after another seized and rejected a single white moth which happened to be among them. Young pheasants and partridges which eat many kinds of caterpillars seem to have an absolute dread of that of the common currant moth, which they will never touch, and tomt.i.ts as well as other small birds appear never to eat the same species. In the case of the Heliconidae, however, we have some direct evidence to the same effect. In the Brazilian forests there are great numbers of insectivorous birds--as jacamars, trogons, and puffbirds--which catch insects on the wing, and that they destroy many b.u.t.terflies 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 Heliconidae, while those of the large showy Nymphalidae, 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 b.u.t.terflies, which they brought to their nest to feed their young; yet during half an hour they never brought one of the Heliconidae, which were flying lazily about in great numbers, and which they could have captured more easily than any others. It was this circ.u.mstance that led Mr. Belt to observe them so long, as he could not understand why the most common insects should be altogether pa.s.sed by.

Mr. Bates also tells us that he never saw them molested by lizards or predacious flies, which often pounce on other b.u.t.terflies.

If, therefore, we accept it as highly probable (if not proved) that the Heliconidae 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 b.u.t.terflies 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 a.s.similate 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 Heliconidae, 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 circ.u.mstances, it is evident that any other b.u.t.terfly 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 b.u.t.terfly 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 b.u.t.terflies 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 Heliconidae, 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 Heliconidae, 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 pa.s.sed by and gain another day's life, which might in many cases be sufficient for it to lay a quant.i.ty 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 b.u.t.terflies forming the family Pieridae (many of which do not greatly differ in appearance from our own cabbage b.u.t.terflies) is a genus of rather small size (Leptalis), some species of which are white like their allies, while the larger number exactly resemble the Heliconidae 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 Heliconidae 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 ident.i.ty in cause and purpose with protective resemblance to vegetable or inorganic forms.

The imitation of the Heliconidae 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 antennae 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 Heliconidae. 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 Heliconidae 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 Heliconidae;--a genus of quite another family of most lovely small American b.u.t.terflies, the Erycinidae, 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 Heliconidae 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 Heliconidae. 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 Heliconidae, 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 Heliconidae themselves mimic other groups. Species of Heliconia mimic Mechanitis, and every species of Napeogenes mimics some other Heliconideous b.u.t.terfly. This would seem to indicate that the distasteful secretion is not produced alike by all members of the family, and that where it is deficient protective imitation comes into play. It is this, perhaps, that has caused such a general resemblance among the Heliconidae, such a uniformity of type with great diversity of colouring, since any aberration causing an insect to cease to look like one of the family would inevitably lead to its being attacked, wounded, and exterminated, even although it was not eatable.

In other parts of the world an exactly parallel series of facts have been observed. The Danaidae and the Acraeidae of the Old World tropics form in fact one great group with the Heliconidae. They have the same general form, structure, and habits: they possess the same protective odour, and are equally abundant in individuals, although not so varied in colour, blue and white spots on a black ground being the most general pattern.

The insects which mimic these are chiefly Papilios, and Diadema, a genus allied to our peac.o.c.k and tortoisesh.e.l.l b.u.t.terflies. In tropical Africa there is a peculiar group of the genus Danais, characterized by dark-brown and bluish-white colours, arranged in bands or stripes. One of these, Danais niavius, is exactly imitated both by Papilio hippoc.o.o.n and by Diadema anthedon; another, Danais echeria, by Papilio cenea; and in Natal a variety of the Danais is found having a white spot at the tip of wings, accompanied by a variety of the Papilio bearing a corresponding white spot. Acraea gea is copied in its very peculiar style of colouration by the female of Papilio cynorta, by Panopaea hirce, and by the female of Elymnias phegea. Acraea euryta of Calabar has a female variety of Panopea hirce from the same place which exactly copies it; and Mr. Trimen, in his paper on Mimetic a.n.a.logies among African b.u.t.terflies, published in the Transactions of the Linnaean Society for 1868, gives a list of no less than sixteen species and varieties of Diadema and its allies, and ten of Papilio, which in their colour and markings are perfect mimics of species or varieties of Danais or Acraea which inhabit the same districts.

Pa.s.sing on to India, we have Danais tytia, a b.u.t.terfly with semi-transparent bluish wings and a border of rich reddish brown. This remarkable style of colouring is exactly reproduced in Papilio agestor and in Diadema nama, and all three insects not unfrequently come together in collections made at Darjeeling. In the Philippine Islands the large and curious Idea leuconoe with its semi-transparent white wings, veined and spotted with black, is copied by the rare Papilio idaeoides from the same islands.

In the Malay archipelago the very common and beautiful Euploea midamus is so exactly mimicked by two rare Papilios (P. paradoxa and P. aenigma) that I generally caught them under the impression that they were the more common species; and the equally common and even more beautiful Euploea rhadamanthus, with its pure white bands and spots on a ground of glossy blue and black, is reproduced in the Papilio caunus. Here also there are species of Diadema imitating the same group in two or three instances; but we shall have to adduce these further on in connexion with another branch of the subject.

It has been already mentioned that in South America there is a group of Papilios which have all the characteristics of a protected race, and whose peculiar colours and markings are imitated by other b.u.t.terflies not so protected. There is just such a group also in the East, having very similar colours and the same habits, and these also are mimicked by other species in the same genus not closely allied to them, and also by a few of other families. Papilio hector, a common Indian b.u.t.terfly of a rich black colour spotted with crimson, is so closely copied by Papilio romulus, that the latter insect has been thought to be its female. A close examination shows, however, that it is essentially different, and belongs to another section of the genus. Papilio antiphus and P.

diphilus, black swallow-tailed b.u.t.terflies with cream-coloured spots, are so well imitated by varieties of P. theseus, that several writers have cla.s.sed them as the same species. Papilio liris, found only in the island of Timor, is accompanied there by P. aenomaus, the female of which so exactly resembles it that they can hardly be separated in the cabinet, and on the wing are quite undistinguishable. But one of the most curious cases is the fine yellow-spotted Papilio c.o.o.n, which is unmistakeably imitated by the female tailed form of Papilio memnon.

These are both from Sumatra; but in North India P. c.o.o.n is replaced by another species, which has been named P. doubledayi, having red spots instead of yellow; and in the same district the corresponding female tailed form of Papilio androgeus, sometimes considered a variety of P.

memnon, is similarly red-spotted. Mr. Westwood has described some curious day-flying moths (Epicopeia) from North India, which have the form and colour of Papilios of this section, and two of these are very good imitations of Papilio polydorus and Papilio varuna, also from North India.

Almost all these cases of mimicry are from the tropics, where the forms of life are more abundant, and where insect development especially is of unchecked luxuriance; but there are also one or two instances in temperate regions. In North America, the large and handsome red and black b.u.t.terfly Danais erippus is very common; and the same country is inhabited by Limenitis archippus, which closely resembles the Danais, while it differs entirely from every species of its own genus.

The only case of probable mimicry in our own country is the following:--A very common white moth (Spilosoma menthastri) was found by Mr. Stainton to be rejected by young turkeys among hundreds of other moths on which they greedily fed. Each bird in succession took hold of this moth and threw it down again, as if too nasty to eat. Mr. Jenner Weir also found that this moth was refused by the Bullfinch, Chaffinch, Yellow Hammer, and Red Bunting, but eaten after much hesitation by the Robin. We may therefore fairly conclude that this species would be disagreeable to many other birds, and would thus have an immunity from attack, which may be the cause of its great abundance and of its conspicuous white colour. Now it is a curious thing that there is another moth, Diaphora mendica, which appears about the same time, and whose female only is white. It is about the same size as Spilosoma menthastri, and sufficiently resembles it in the dusk, and this moth is much less common. It seems very probable, therefore, that these species stand in the same relation to each other as the mimicking b.u.t.terflies of various families do to the Heliconidae and Danaidae. It would be very interesting to experiment on all white moths, to ascertain if those which are most common are generally rejected by birds. It may be antic.i.p.ated that they would be so, because white is the most conspicuous of all colours for nocturnal insects, and had they not some other protection would certainly be very injurious to them.

_Lepidoptera mimicking other Insects._

In the preceding cases we have found Lepidoptera imitating other species of the same order, and such species only as we have good reason to believe were free from the attacks of many insectivorous creatures; but there are other instances in which they altogether lose the external appearance of the order to which they belong, and take on the dress of bees or wasps--insects which have an undeniable protection in their stings. The Sesiidae and aegeriidae, two families of day-flying moths, are particularly remarkable in this respect, and a mere inspection of the names given to the various species shows how the resemblance has struck everyone. We have apiformis, vespiforme, ichneumoniforme, scoliaeforme, sphegiforme (bee-like, wasp-like, ichneumon-like, &c.) and many others, all indicating a resemblance to stinging Hymenoptera. In Britain we may particularly notice Sesia bombiliformis, which very closely resembles the male of the large and common humble bee, Bombus hortorum; Sphecia craboniforme, which is coloured like a hornet, and is (on the authority of Mr. Jenner Weir) much more like it when alive than when in the cabinet, from the way in which it carries its wings; and the currant clear-wing, Trochilium tipuliforme, which resembles a small black wasp (Odynerus sinuatus) very abundant in gardens at the same season. It has been so much the practice to look upon these resemblances as mere curious a.n.a.logies playing no part in the economy of nature, that we have scarcely any observations of the habits and appearance when alive of the hundreds of species of these groups in various parts of the world, or how far they are accompanied by Hymenoptera, which they specifically resemble. There are many species in India (like those figured by Professor Westwood in his "Oriental Entomology") which have the hind legs very broad and densely hairy, so as exactly to imitate the brush-legged bees (Scopulipedes) which abound in the same country. In this case we have more than mere resemblance of colour, for that which is an important functional structure in the one group is imitated in another whose habits render it perfectly useless.

_Mimicry among Beetles._

It may fairly be expected that if these imitations of one creature by another really serve as a protection to weak and decaying species, instances of the same kind will be found among other groups than the Lepidoptera; and such is the case, although they are seldom so prominent and so easily recognised as those already pointed out as occurring in that order. A few very interesting examples may, however, be pointed out in most of the other orders of insects. The Coleoptera or beetles that imitate other Coleoptera of distinct groups are very numerous in tropical countries, and they generally follow the laws already laid down as regulating these phenomena. The insects which others imitate always have a special protection, which leads them to be avoided as dangerous or uneatable by small insectivorous animals; some have a disgusting taste (a.n.a.logous to that of the Heliconidae); others have such a hard and stony covering that they cannot be crushed or digested; while a third set are very active, and armed with powerful jaws, as well as having some disagreeable secretion. Some species of Eumorphidae and Hispidae, small flat or hemispherical beetles which are exceedingly abundant, and have a disagreeable secretion, are imitated by others of the very distinct group of Longicornes (of which our common musk-beetle may be taken as an example). The extraordinary little Cyclopeplus batesii, belongs to the same sub-family of this group as the Onychocerus scorpio and O. concentricus, which have already been adduced as imitating with such wonderful accuracy the bark of the trees they habitually frequent; but it differs totally in outward appearance from every one of its allies, having taken upon itself the exact shape and colouring of a globular Corynomalus, a little stinking beetle with clubbed antennae. It is curious to see how these clubbed antennae are imitated by an insect belonging to a group with long slender antennae. The sub-family Anisocerinae, to which Cyclopeplus belongs, is characterised by all its members possessing a little k.n.o.b or dilatation about the middle of the antennae. This k.n.o.b is considerably enlarged in C. batesii, and the terminal portion of the antennae beyond it is so small and slender as to be scarcely visible, and thus an excellent subst.i.tute is obtained for the short clubbed antennae of the Corynomalus. Erythroplatis corallifer is another curious broad flat beetle, that no one would take for a Longicorn, since it almost exactly resembles Cephalodonta spinipes, one of the commonest of the South American Hispidae; and what is still more remarkable, another Longicorn of a distinct group, Streptolabis hispoides, was found by Mr. Bates, which resembles the same insect with equal minuteness,--a case exactly parallel to that among b.u.t.terflies, where species of two or three distinct groups mimicked the same Heliconia. Many of the soft-winged beetles (Malacoderms) are excessively abundant in individuals, and it is probable that they have some similar protection, more especially as other species often strikingly resemble them. A Longicorn beetle, Paeciloderma terminale, found in Jamaica, is coloured exactly in the same way as a Lycus (one of the Malacoderms) from the same island. Eroschema poweri, a Longicorn from Australia, might certainly be taken for one of the same group, and several species from the Malay Islands are equally deceptive. In the Island of Celebes I found one of this group, having the whole body and elytra of a rich deep blue colour, with the head only orange; and in company with it an insect of a totally different family (Eucnemidae) with identically the same colouration, and of so nearly the same size and form as to completely puzzle the collector on every fresh occasion of capturing them. I have been recently informed by Mr. Jenner Weir, who keeps a variety of small birds, that none of them will touch our common "soldiers and sailors"

(species of Malacoderms), thus confirming my belief that they were a protected group, founded on the fact of their being at once very abundant, of conspicuous colours, and the objects of mimicry.

There are a number of the larger tropical weevils which have the elytra and the whole covering of the body so hard as to be a great annoyance to the entomologist, because in attempting to transfix them the points of his pins are constantly turned. I have found it necessary in these cases to drill a hole very carefully with the point of a sharp penknife before attempting to insert a pin. Many of the fine long-antennaed Anthribidae (an allied group) have to be treated in the same way. We can easily understand that after small birds have in vain attempted to eat these insects, they should get to know them by sight, and ever after leave them alone, and it will then be an advantage for other insects which are comparatively soft and eatable, to be mistaken for them. We need not be surprised, therefore, to find that there are many Longicorns which strikingly resemble the "hard beetles" of their own district. In South Brazil, Acanthotritus dorsalis is strikingly like a Curculio of the hard genus Heiliplus, and Mr. Bates a.s.sures me that he found Gymnocerus cratosomoides (a Longicorn) on the same tree with a hard Cratosomus (a weevil), which it exactly mimics. Again, the pretty Longicorn, Phacellocera batesii, mimics one of the hard Anthribidae of the genus Ptychoderes, having long slender antennae. In the Moluccas we find Cacia anthriboides, a small Longicorn which might be easily mistaken for a very common species of Anthribidae found in the same districts; and the very rare Capnolymma stygium closely imitates the common Mecocerus gazella, which abounded where it was taken. Doliops curculionoides and other allied Longicorns from the Philippine Islands most curiously resemble, both in form and colouring, the brilliant Pachyrhynchi,--Curculionidae, which are almost peculiar to that group of islands. The remaining family of Coleoptera most frequently imitated is the Cicindelidae. The rare and curious Longicorn, Collyrodes lacordairei, has exactly the form and colouring of the genus Collyris, while an undescribed species of Heteromera is exactly like a Therates, and was taken running on the trunks of trees, as is the habit of that group.

There is one curious example of a Longicorn mimicking a Longicorn, like the Papilios and Heliconidae which mimic their own allies. Agnia fasciata, belonging to the sub-family Hypselominae, and Nemophas grayi, belonging to the Lamiinae, were taken in Amboyna on the same fallen tree at the same time, and were supposed to be the same species till they were more carefully examined, and found to be structurally quite different. The colouring of these insects is very remarkable, being rich steel-blue black, crossed by broad hairy bands of orange buff, and out of the many thousands of known species of Longicorns they are probably the only two which are so coloured. The Nemophas grayi is the larger, stronger, and better armed insect, and belongs to a more widely spread and dominant group, very rich in species and individuals, and is therefore most probably the subject of mimicry by the other species.

_Beetles mimicking other Insects._

We will now adduce a few cases in which beetles imitate other insects, and insects of other orders imitate beetles.

Charis melipona, a South American Longicorn of the family Necydalidae, has been so named from its resemblance to a small bee of the genus Melipona. It is one of the most remarkable cases of mimicry, since the beetle has the thorax and body densely hairy like the bee, and the legs are tufted in a manner most unusual in the order Coleoptera. Another Longicorn, Odontocera odyneroides, has the abdomen banded with yellow, and constricted at the base, and is altogether so exactly like a small common wasp of the genus Odynerus, that Mr. Bates informs us he was afraid to take it out of his net with his fingers for fear of being stung. Had Mr. Bates's taste for insects been less omnivorous than it was, the beetle's disguise might have saved it from his pin, as it had no doubt often done from the beak of hungry birds. A larger insect, Sphecomorpha chalybea, is exactly like one of the large metallic blue wasps, and like them has the abdomen connected with the thorax by a pedicel, rendering the deception most complete and striking. Many Eastern species of Longicorns of the genus Oberea, when on the wing exactly resemble Tenthredinidae, and many of the small species of Hesthesis run about on timber, and cannot be distinguished from ants.

There is one genus of South American Longicorns that appears to mimic the s.h.i.+elded bugs of the genus Scutellera. The Gymnocerous capucinus is one of these, and is very like Pachyotris fabricii, one of the Scutelleridae. The beautiful Gymnocerous dulcissimus is also very like the same group of insects, though there is no known species that exactly corresponds to it; but this is not to be wondered at, as the tropical Hemiptera have been comparatively so little cared for by collectors.

_Insects mimicking Species of other Orders._

The most remarkable case of an insect of another order mimicking a beetle is that of the Condylodera tricondyloides, one of the cricket family from the Philippine Islands, which is so exactly like a Tricondyla (one of the tiger beetles), that such an experienced entomologist as Professor Westwood placed it among them in his cabinet, and retained it there a long time before he discovered his mistake! Both insects run along the trunks of trees, and whereas Tricondylas are very plentiful, the insect that mimics it is, as in all other cases, very rare. Mr. Bates also informs us that he found at Santarem on the Amazon, a species of locust which mimicked one of the tiger beetles of the genus Odontocheila, and was found on the same trees which they frequented.

There are a considerable number of Diptera, or two-winged flies, that closely resemble wasps and bees, and no doubt derive much benefit from the wholesome dread which those insects excite. The Midas dives, and other species of large Brazilian flies, have dark wings and metallic blue elongate bodies, resembling the large stinging Sphegidae of the same country; and a very large fly of the genus Asilus has black-banded wings and the abdomen tipped with rich orange, so as exactly to resemble the fine bee Euglossa dimidiata, and both are found in the same parts of South America. We have also in our own country species of Bombylius which are almost exactly like bees. In these cases the end gained by the mimicry is no doubt freedom from attack, but it has sometimes an altogether different purpose. There are a number of parasitic flies whose larvae feed upon the larvae of bees, such as the British genus Volucella and many of the tropical Bombylii, and most of these are exactly like the particular species of bee they prey upon, so that they can enter their nests unsuspected to deposit their eggs. There are also bees that mimic bees. The cuckoo bees of the genus Nomada are parasitic on the Andrenidae, and they resemble either wasps or species of Andrena; and the parasitic humble-bees of the genus Apathus almost exactly resemble the species of humble-bees in whose nests they are reared. Mr.

Bates informs us that he found numbers of these "cuckoo" bees and flies on the Amazon, which all wore the livery of working bees peculiar to the same country.

There is a genus of small spiders in the tropics which feed on ants, and they are exactly like ants themselves, which no doubt gives them more opportunity of seizing their prey; and Mr. Bates found on the Amazon a species of Mantis which exactly resembled the white ants which it fed upon, as well as several species of crickets (Scaphura), which resembled in a wonderful manner different sand-wasps of large size, which are constantly on the search for crickets with which to provision their nests.

Perhaps the most wonderful case of all is the large caterpillar mentioned by Mr. Bates, which startled him by its close resemblance to a small snake. The first three segments behind the head were dilatable at the will of the insect, and had on each side a large black pupillated spot, which resembled the eye of the reptile. Moreover, it resembled a poisonous viper, not a harmless species of snake, as was proved by the imitation of keeled scales on the crown produced by the rec.u.mbent feet, as the caterpillar threw itself backward!

The att.i.tudes of many of the tropical spiders are most extraordinary and deceptive, but little attention has been paid to them. They often mimic other insects, and some, Mr. Bates a.s.sures us, are exactly like flower buds, and take their station in the axils of leaves, where they remain motionless waiting for their prey.

_Cases of Mimicry among the Vertebrata._

Having thus shown how varied and extraordinary are the modes in which mimicry occurs among insects, we have now to enquire if anything of the same kind is to be observed among vertebrated animals. When we consider all the conditions necessary to produce a good deceptive imitation, we shall see at once that such can very rarely occur in the higher animals, since they possess none of those facilities for the almost infinite modifications of external form which exist in the very nature of insect organization. The outer covering of insects being more or less solid and h.o.r.n.y, they are capable of almost any amount of change of form and appearance without any essential modification internally. In many groups the wings give much of the character, and these organs may be much modified both in form and colour without interfering with their special functions. Again, the number of species of insects is so great, and there is such diversity of form and proportion in every group, that the chances of an accidental approximation in size, form, and colour, of one insect to another of a different group, are very considerable; and it is these chance approximations that furnish the basis of mimicry, to be continually advanced and perfected by the survival of those varieties only which tend in the right direction.

In the Vertebrata, on the contrary, the skeleton being internal the external form depends almost entirely on the proportions and arrangement of that skeleton, which again is strictly adapted to the functions necessary for the well-being of the animal. The form cannot therefore be rapidly modified by variation, and the thin and flexible integument will not admit of the development of such strange protuberances as occur continually in insects. The number of species of each group in the same country is also comparatively small, and thus the chances of that first accidental resemblance which is necessary for natural selection to work upon are much diminished. We can hardly see the possibility of a mimicry by which the elk could escape from the wolf, or the buffalo from the tiger. There is, however, in one group of Vertebrata such a general similarity of form, that a very slight modification, if accompanied by ident.i.ty of colour, would produce the necessary amount of resemblance; and at the same time there exist a number of species which it would be advantageous for others to resemble, since they are armed with the most fatal weapons of offence. We accordingly find that reptiles furnish us with a very remarkable and instructive case of true mimicry.

_Mimicry among Snakes._

There are in tropical America a number of venomous snakes of the genus Elaps, which are ornamented with brilliant colours disposed in a peculiar manner. The ground colour is generally bright red, on which are black bands of various widths and sometimes divided into two or three by yellow rings. Now, in the same country are found several genera of harmless snakes, having no affinity whatever with the above, but coloured exactly the same. For example, the poisonous Elaps fulvius often occurs in Guatemala with simple black bands on a coral-red ground; and in the same country is found the harmless snake Pliocerus equalis, coloured and banded in identically the same manner. A variety of Elaps corallinus has the black bands narrowly bordered with yellow on the same red ground colour, and a harmless snake, Homalocranium semicinctum, has exactly the same markings, and both are found in Mexico. The deadly Elaps lemniscatus has the black bands very broad, and each of them divided into three by narrow yellow rings; and this again is exactly copied by a harmless snake, Pliocerus elapoides, which is found along with its model in Mexico.

But, more remarkable still, there is in South America a third group of snakes, the genus Oxyrhopus, doubtfully venomous, and having no immediate affinity with either of the preceding, which has also the same curious distribution of colours, namely, variously disposed rings of red, yellow, and black; and there are some cases in which species of all three of these groups similarly marked inhabit the same district. For example, Elaps mipart.i.tus has single black rings very close together. It inhabits the west side of the Andes, and in the same districts occur Pliocerus euryzonus and Oxyrhopus petolarius, which exactly copy its pattern. In Brazil Elaps lemniscatus is copied by Oxyrhopus trigeminus, both having black rings disposed in threes. In Elaps hemiprichii the ground colour appears to be black, with alternations of two narrow yellow bands and a broader red one; and of this pattern again we have an exact double in Oxyrhopus formosus, both being found in many localities of tropical South America.

What adds much to the extraordinary character of these resemblances is the fact, that nowhere in the world but in America are there any snakes at all which have this style of colouring. Dr. Gunther, of the British Museum, who has kindly furnished some of the details here referred to, a.s.sures me that this is the case; and that red, black, and yellow rings occur together on no other snakes in the world but on Elaps and the species which so closely resemble it. In all these cases, the size and form as well as the colouration, are so much alike, that none but a naturalist would distinguish the harmless from the poisonous species.

Contributions to the Theory of Natural Selection Part 4

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