Special Protective Colouring of Butterflies.

We will now consider a few cases of special protective colouring in the perfect butterfly or moth. Mr. Mansel Weale states that in South Africa there is a great prevalence of white and silvery foliage or bark, sometimes of dazzling brilliancy, and that many insects and their larvae have brilliant silvery tints which are protective, among them being three species of butterflies whose undersides are silvery, and which are thus effectually protected when at rest.[73] A common African butterfly (Aterica meleagris) always settles on the ground with closed wings, which so closely resemble the soil of the district that it can with difficulty be seen, and the colour varies with the soil in different localities. Thus specimens from Senegambia were dull brown, the soil being reddish sand and iron-clay; those from Calabar and Cameroons were light brown with numerous small white spots, the soil of those countries being light brown clay with small quartz pebbles; while in other localities where the colours of the soil were more varied the colours of the butterfly varied also. Here we have variation in a single species which has become specialised in certain areas to harmonise with the colour of the soil.[74]

Many butterflies, in all parts of the world, resemble dead leaves on their under side, but those in which this form of protection is carried to the greatest perfection are the species of the Eastern genus Kallima. In India K. inachis, and in the larger Malay islands K. paralekta, are very common. They are rather large and showy butterflies, orange and bluish on the upper side, with a very rapid flight, and frequenting dry forests. Their habit is to settle always where there is some dead or decaying foliage, and the shape and colour of the wings (on the under surface), together with the attitude of the insect, is such as to produce an absolutely perfect imitation of a dead leaf. This is effected by the butterfly always settling on a twig, with the short tail of the hind wings just touching it and forming the leaf-stalk. From this a dark curved line runs across to the elongated tip of the upper wings, imitating the midrib, on both sides of which are oblique lines, formed partly by the nervures and partly by markings, which give the effect of the usual veining of a leaf. The head and antennae fit exactly between the closed upper wings so as not to interfere with the outline, which has just that amount of irregular curvature that is seen in dry and withered leaves. The colour is very remarkable for its extreme amount of variability, from deep reddish-brown to olive or pale yellow, hardly two specimens being exactly alike, but all coming within the range of colour of leaves in various stages of decay. Still more curious is the fact that the paler wings, which imitate leaves most decayed, are usually covered with small black dots, often gathered into circular groups, and so exactly resembling the minute fungi on decaying leaves that it is hard at first to believe that the insects themselves are not attacked by some such fungus. The concealment produced by this wonderful imitation is most complete, and in Sumatra I have often seen one enter a bush and then disappear like magic. Once I was so fortunate as to see the exact spot on which the insect settled; but even then I lost sight of it for some time, and only after a persistent search discovered that it was close before my eyes.[75] Here we have a kind of imitation, which is very common in a less developed form, carried to extreme perfection, with the result that the species is very abundant over a considerable area of country.

Protective Resemblance among Marine Animals.

Among marine animals this form of protection is very common. Professor Moseley tells us that all the inhabitants of the Gulf-weed are most remarkably coloured, for purposes of protection and concealment, exactly like the weed itself. "The shrimps and crabs which swarm in the weed are of exactly the same shade of yellow as the weed, and have white markings upon their bodies to represent the patches of Membranipora. The small fish, Antennarius, is in the same way weed-colour with white spots. Even a Planarian worm, which lives in the weed, is similarly yellow-coloured, and also a mollusc, Scyllaea pelagica." The same writer tells us that "a number of little crabs found clinging to the floats of the blue-shelled mollusc, Ianthina, were all coloured of a corresponding blue for concealment."[76]

Professor E.S. Morse of Salem, Mass., found that most of the New England marine mollusca were protectively coloured; instancing among others a little red chiton on rocks clothed with red calcareous algae, and Crepidula plana, living within the apertures of the shells of larger species of Gasteropods and of a pure white colour corresponding to its habitat, while allied species living on seaweed or on the outside of dark shells were dark brown.[77] A still more interesting case has been recorded by Mr. George Brady. He says: "Amongst the Nullipore which matted together the laminaria roots in the Firth of Clyde were living numerous small starfishes (Ophiocoma bellis) which, except when their writhing movements betrayed them, were quite undistinguishable from the calcareous branches of the alga; their rigid angularly twisted rays had all the appearance of the coralline, and exactly assimilated to its dark purple colour, so that though I held in my hand a root in which were half a dozen of the starfishes, I was really unable to detect them until revealed by their movements."[78]

These few examples are sufficient to show that the principle of protective coloration extends to the ocean as well as over the earth; and if we consider how completely ignorant we are of the habits and surroundings of most marine animals, it may well happen that many of the colours of tropical fishes, which seem to us so strange and so conspicuous, are really protective, owing to the number of equally strange and brilliant forms of corals, sea-anemones, sponges, and seaweeds among which they live.

Protection by Terrifying Enemies.

A considerable number of quite defenceless insects obtain protection from some of their enemies by having acquired a resemblance to dangerous animals, or by some threatening or unusual appearance. This is obtained either by a modification of shape, of habits, of colour, or of all combined. The simplest form of this protection is the aggressive attitude of the caterpillars of the Sphingidae, the forepart of the body being erected so as to produce a rude resemblance to the figure of a sphinx, hence the name of the family. The protection is carried further by those species which retract the first three segments and have large ocelli on each side of the fourth segment, thus giving to the caterpillar, when the forepart of its body is elevated, the appearance of a snake in a threatening attitude.

The blood-red forked tentacle, thrown out of the neck of the larvae of the genus Papilio when alarmed, is, no doubt, a protection against the attacks of ichneumons, and may, perhaps, also frighten small birds; and the habit of turning up the tail possessed by the harmless rove-beetles (Staphylinidae), giving the idea that they can sting, has, probably, a similar use. Even an unusual angular form, like a crooked twig or inorganic substance, may be protective; as Mr. Poulton thinks is the case with the curious caterpillar of Notodonta ziczac, which, by means of a few slight protuberances on its body, is able to assume an angular and very unorganic-looking appearance. But perhaps the most perfect example of this kind of protection is exhibited by the large caterpillar of the Royal Persimmon moth (Bombyx regia), a native of the southern states of North America, and known there as the "Hickory-horned devil." It is a large green caterpillar, often six inches long, ornamented with an immense crown of orange-red tubercles, which, if disturbed, it erects and shakes from side to side in a very alarming manner. In its native country the negroes believe it to be as deadly as a rattlesnake, whereas it is perfectly innocuous. The green colour of the body suggests that its ancestors were once protectively coloured; but, growing too large to be effectually concealed, it acquired the habit of shaking its head about in order to frighten away its enemies, and ultimately developed the crown of tentacles as an addition to its terrifying powers. This species is beautifully figured in Abbott and Smith's Lepidopterous Insects of Georgia.

Alluring Coloration.

Besides those numerous insects which obtain protection through their resemblance to the natural objects among which they live, there are some whose disguise is not used for concealment, but as a direct means of securing their prey by attracting them within the enemy's reach. Only a few cases of this kind of coloration have yet been observed, chiefly among spiders and mantidae; but, no doubt, if attention were given to the subject in tropical countries, many more would be discovered. Mr. H.O. Forbes has described a most interesting example of this kind of simulation in Java. While pursuing a large butterfly through the jungle, he was stopped by a dense bush, on a leaf of which he observed one of the skipper butterflies sitting on a bird's dropping. "I had often," he says, "observed small Blues at rest on similar spots on the ground, and have wondered what such a refined and beautiful family as the Lycaenidae could find to enjoy, in food apparently so incongruous for a butterfly. I approached with gentle steps, but ready net, to see if possible how the present species was engaged. It permitted me to get quite close, and even to seize it between my fingers; to my surprise, however, part of the body remained behind, adhering as I thought to the excreta. I looked closely, and finally touched with my finger the excreta to find if it were glutinous. To my delighted astonishment I found that my eyes had been most perfectly deceived, and that what seemed to be the excreta was a most artfully coloured spider, lying on its back with its feet crossed over and closely adpressed to the body." Mr. Forbes then goes on to describe the exact appearance of such excreta, and how the various parts of the spider are coloured to produce the imitation, even to the liquid portion which usually runs a little down the leaf. This is exactly imitated by a portion of the thin web which the spider first spins to secure himself firmly to the leaf; thus producing, as Mr. Forbes remarks, a living bait for butterflies and other insects so artfully contrived as to deceive a pair of human eyes, even when intently examining it.[79]

A native species of spider (Thomisus citreus) exhibits a somewhat similar alluring protection by its close resemblance to buds of the wayfaring tree, Viburnum lantana. It is pure creamy-white, the abdomen exactly resembling in shape and colour the unopened buds of the flowers among which it takes its station; and it has been seen to capture flies which came to the flowers.

But the most curious and beautiful case of alluring protection is that of a wingless Mantis in India, which is so formed and coloured as to resemble a pink orchis or some other fantastic flower. The whole insect is of a bright pink colour, the large and oval abdomen looking like the labellum of an orchid. On each side, the two posterior legs have immensely dilated and flattened thighs which represent the petals of a flower, while the neck and forelegs imitate the upper sepal and column of an orchid. The insect rests motionless, in this symmetrical attitude, among bright green foliage, being of course very conspicuous, but so exactly resembling a flower that butterflies and other insects settle upon it and are instantly captured. It is a living trap, baited in the most alluring manner to catch the unwary flower-haunting insects.[80]

The Coloration of Birds' Eggs.

The colours of birds' eggs have long been a difficulty on the theory of adaptive coloration, because, in so many cases it has not been easy to see what can be the use of the particular colours, which are often so bright and conspicuous that they seem intended to attract attention rather than to be concealed. A more careful consideration of the subject in all its bearings shows, however, that here too, in a great number of cases, we have examples of protective coloration. When, therefore, we cannot see the meaning of the colour, we may suppose that it has been protective in some ancestral form, and, not being hurtful, has persisted under changed conditions which rendered the protection needless.

We may divide all eggs, for our present purpose, into two great divisions; those which are white or nearly so, and those which are distinctly coloured or spotted. Egg-shells being composed mainly of carbonate of lime, we may assume that the primitive colour of birds' eggs was white, a colour that prevails now among the other egg-bearing vertebrates—lizards, crocodiles, turtles, and snakes; and we might, therefore, expect that this colour would continue where its presence had no disadvantages. Now, as a matter of fact, we find that in all the groups of birds which lay their eggs in concealed places, whether in holes of trees or in the ground, or in domed or covered nests, the eggs are either pure white or of very pale uniform coloration. Such is the case with kingfishers, bee-eaters, penguins, and puffins, which nest in holes in the ground; with the great parrot family, the woodpeckers, the rollers, hoopoes, trogons, owls, and some others, which build in holes in trees or other concealed places; while martins, wrens, willow-warblers, and Australian finches, build domed or covered nests, and usually have white eggs.

There are, however, many other birds which lay their white eggs in open nests; and these afford some very interesting examples of the varied modes by which concealment may be obtained. All the duck tribe, the grebes, and the pheasants belong to this class; but these birds all have the habit of covering their eggs with dead leaves or other material whenever they leave the nest, so as effectually to conceal them. Other birds, as the short-eared owl, the goatsucker, the partridge, and some of the Australian ground pigeons, lay their white or pale eggs on the bare soil; but in these cases the birds themselves are protectively coloured, so that, when sitting, they are almost invisible; and they have the habit of sitting close and almost continuously, thus effectually concealing their eggs.

Pigeons and doves offer a very curious case of the protection of exposed eggs. They usually build very slight and loose nests of sticks and twigs, so open that light can be seen through them from below, while they are generally well concealed by foliage above. Their eggs are white and shining; yet it is a difficult matter to discover, from beneath, whether there are eggs in the nest or not, while they are well hidden by the thick foliage above. The Australian podargihuge goatsuckers—build very similar nests, and their white eggs are protected in the same manner. Some large and powerful birds, as the swans, herons, pelicans, cormorants, and storks, lay white eggs in open nests; but they keep careful watch over them, and are able to drive away intruders. On the whole, then, we see that, while white eggs are conspicuous, and therefore especially liable to attack by egg-eating animals, they are concealed from observation in many and various ways. We may, therefore, assume that, in cases where there seems to be no such concealment, we are too ignorant of the whole of the conditions to form a correct judgment.

We now come to the large class of coloured or richly spotted eggs, and here we have a more difficult task, though many of them decidedly exhibit protective tints or markings. There are two birds which nest on sandy shores—the lesser tern and the ringed plover,—and both lay sand-coloured eggs, the former spotted so as to harmonise with coarse shingle, the latter minutely speckled like fine sand, which are the kinds of ground the two birds choose respectively for their nests. "The common sandpipers' eggs assimilate so closely with the tints around them as to make their discovery a matter of no small difficulty, as every oologist can testify who has searched for them. The pewits' eggs, dark in ground colour and boldly marked, are in strict harmony with the sober tints of moor and fallow, and on this circumstance alone their concealment and safety depend. The divers' eggs furnish another example of protective colour; they are generally laid close to the water's edge, amongst drift and shingle, where their dark tints and black spots conceal them by harmonising closely with surrounding objects. The snipes and the great army of sandpipers furnish innumerable instances of protectively coloured eggs. In all the instances given the sitting-bird invariably leaves the eggs uncovered when it quits them, and consequently their safety depends solely on the colours which adorn them."[81] The wonderful range of colour and marking in the eggs of the guillemot may be imputed to the inaccessible rocks on which it breeds, giving it complete protection from enemies. Thus the pale or bluish ground colour of the eggs of its allies, the auks and puffins, has become intensified and blotched and spotted in the most marvellous variety of patterns, owing to there being no selective agency to prevent individual variation having full sway.

The common black coot (Fulica atra) has eggs which are coloured in a specially protective manner. Dr. William Marshall writes, that it only breeds in certain localities where a large water reed (Phragmites arundinacea) abounds. The eggs of the coot are stained and spotted with black on a yellowish-gray ground, and the dead leaves of the reed are of the same colour, and are stained black by small parasitic fungi of the Uredo family; and these leaves form the bed on which the eggs are laid. The eggs and the leaves agree so closely in colour and markings that it is a difficult thing to distinguish the eggs at any distance. It is to be noted that the coot never covers up its eggs, as its ally the moor-hen usually does.

The beautiful blue or greenish eggs of the hedge-sparrow, the song-thrush, and sometimes those of the blackbird, seem at first sight especially calculated to attract attention, but it is very doubtful whether they are really so conspicuous when seen at a little distance among their usual surroundings. For the nests of these birds are either in evergreens, as holly or ivy, or surrounded by the delicate green tints of our early spring vegetation, and may thus harmonise very well with the colours around them. The great majority of the eggs of our smaller birds are so spotted or streaked with brown or black on variously tinted grounds that, when lying in the shadow of the nest and surrounded by the many colours and tints of bark and moss, of purple buds and tender green or yellow foliage, with all the complex glittering lights and mottled shades produced among these by the spring sunshine and by sparkling raindrops, they must have a quite different aspect from that which they possess when we observe them torn from their natural surroundings. We have here, probably, a similar case of general protective harmony to that of the green caterpillars with beautiful white or purple bands and spots, which, though gaudily conspicuous when seen alone, become practically invisible among the complex lights and shadows of the foliage they feed upon.

In the case of the cuckoo, which lays its eggs in the nests of a variety of other birds, the eggs themselves are subject to considerable variations of colour, the most common type, however, resembling those of the pipits, wagtails, or warblers, in whose nests they are most frequently laid. It also often lays in the nest of the hedge-sparrow, whose bright blue eggs are usually not at all nearly matched, although they are sometimes said to be so on the Continent. It is the opinion of many ornithologists that each female cuckoo lays the same coloured eggs, and that it usually chooses a nest the owners of which lay somewhat similar eggs, though this is by no means universally the case. Although birds which have cuckoos' eggs imposed upon them do not seem to neglect them on account of any difference of colour, yet they probably do so occasionally; and if, as seems probable, each bird's eggs are to some extent protected by their harmony of colour with their surroundings, the presence of a larger and very differently coloured egg in the nest might be dangerous, and lead to the destruction of the whole set. Those cuckoos, therefore, which most frequently placed their eggs among the kinds which they resembled, would in the long run leave most progeny, and thus the very frequent accord in colour might have been brought about.

Some writers have suggested that the varied colours of birds' eggs are primarily due to the effect of surrounding coloured objects on the female bird during the period preceding incubation; and have expended much ingenuity in suggesting the objects that may have caused the eggs of one bird to be blue, another brown, and another pink.[82] But no evidence has been presented to prove that any effects whatever are produced by this cause, while there seems no difficulty in accounting for the facts by individual variability and the action of natural selection. The changes that occur in the conditions of existence of birds must sometimes render the concealment less perfect than it may once have been; and when any danger arises from this cause, it may be met either by some change in the colour of the eggs, or in the structure or position of the nest, or by the increased care which the parents bestow upon the eggs. In this way the various divergences which now so often puzzle us may have arisen.

Colour as a Means of Recognition.

If we consider the habits and life-histories of those animals which are more or less gregarious, comprising a large proportion of the herbivora, some carnivora, and a considerable number of all orders of birds, we shall see that a means of ready recognition of its own kind, at a distance or during rapid motion, in the dusk of twilight or in partial cover, must be of the greatest advantage and often lead to the preservation of life. Animals of this kind will not usually receive a stranger into their midst. While they keep together they are generally safe from attack, but a solitary straggler becomes an easy prey to the enemy; it is, therefore, of the highest importance that, in such a case, the wanderer should have every facility for discovering its companions with certainty at any distance within the range of vision.

Some means of easy recognition must be of vital importance to the young and inexperienced of each flock, and it also enables the sexes to recognise their kind and thus avoid the evils of infertile crosses; and I am inclined to believe that its necessity has had a more widespread influence in determining the diversities of animal coloration than any other cause whatever. To it may probably be imputed the singular fact that, whereas bilateral symmetry of coloration is very frequently lost among domesticated animals, it almost universally prevails in a state of nature; for if the two sides of an animal were unlike, and the diversity of coloration among domestic animals occurred in a wild state, easy recognition would be impossible among numerous closely allied forms.[83] The wonderful diversity of colour and of marking that prevails, especially in birds and insects, may be due to the fact that one of the first needs of a new species would be, to keep separate from its nearest allies, and this could be most readily done by some easily seen external mark of difference. A few illustrations will serve to show how this principle acts in nature.

My attention was first called to the subject by a remark of Mr. Darwin's that, though, "the hare on her form is a familiar instance of concealment through colour, yet the principle partly fails in a closely allied species, the rabbit; for when running to its burrow it is made conspicuous to the sportsman, and no doubt to all beasts of prey, by its upturned white tail."[84] But a little consideration of the habits of the animal will show that the white upturned tail is of the greatest value, and is really, as it has been termed by a writer in The Field, a "signal flag of danger." For the rabbit is usually a crepuscular animal, feeding soon after sunset or on moonlight nights. When disturbed or alarmed it makes for its burrow, and the white upturned tails of those in front serve as guides and signals to those more remote from home, to the young and the feeble; and thus each following the one or two before it, all are able with the least possible delay to regain a place of comparative safety. The apparent danger, therefore, becomes a most important means of security.

The same general principle enables us to understand the singular, and often conspicuous, markings on so many gregarious herbivora which are yet, on the whole, protectively coloured. Thus, the American prong-buck has a white patch behind and a black muzzle. The Tartarian antelope, the Ovis poli of High Asia, the Java wild ox, several species of deer, and a large number of antelopes have a similar conspicuous white patch behind, which, in contrast to the dusky body, must enable them to be seen and followed from a distance by their fellows. Where there are many species of nearly the same general size and form inhabiting the same region—as with the antelopes of Africa—we find many distinctive markings of a similar kind. The gazelles have variously striped and banded faces, besides white patches behind and on the flanks, as shown in the woodcut. The spring-bok has a white patch on the face and one on the sides, with a curiously distinctive white stripe above the tail, which is nearly concealed when the animal is at rest by a fold of skin but comes into full view when it is in motion, being thus quite analogous to the upturned white tail of the rabbit. In the pallah the white rump-mark is bordered with black, and the peculiar shape of the horns distinguishes it when seen from the front. The sable-antelope, the gems-bok, the oryx, the hart-beest, the bonte-bok, and the addax have each peculiar white markings; and they are besides characterised by horns so remarkably different in each species and so conspicuous, that it seems probable that the peculiarities in length, twist, and curvature have been differentiated for the purpose of recognition, rather than for any speciality of defence in species whose general habits are so similar.



It is interesting to note that these markings for recognition are very slightly developed in the antelopes of the woods and marshes. Thus, the grys-bok is nearly uniform in colour, except the long black-tipped ears; and it frequents the wooded mountains. The duyker-bok and the rhoode-bok are wary bush-haunters, and have no marks but the small white patch behind. The wood-haunting bosch-bok goes in pairs, and has hardly any distinctive marks on its dusky chestnut coat, but the male alone is horned. The large and handsome koodoo frequents brushwood, and its vertical white stripes are no doubt protective, while its magnificent spiral horns afford easy recognition. The eland, which is an inhabitant of the open country, is uniformly coloured, being sufficiently recognisable by its large size and distinctive form; but the Derbyan eland is a forest animal, and has a protectively striped coat. In like manner, the fine Speke's antelope, which lives entirely in the swamps and among reeds, has pale vertical stripes on the sides (protective), with white markings on face and breast for recognition. An inspection of the figures of antelopes and other animals in Wood's Natural History, or in other illustrated works, will give a better idea of the peculiarities of recognition markings than any amount of description.

Other examples of such coloration are to be seen in the dusky tints of the musk-sheep and the reindeer, to whom recognition at a distance on the snowy plains is of more importance than concealment from their few enemies. The conspicuous stripes and bands of the zebra and the quagga are probably due to the same cause, as may be the singular crests and face-marks of several of the monkeys and lemurs.[85]



Among birds, these recognition marks are especially numerous and suggestive. Species which inhabit open districts are usually protectively coloured; but they generally possess some distinctive markings for the purpose of being easily recognised by their kind, both when at rest and during flight. Such are, the white bands or patches on the breast or belly of many birds, but more especially the head and neck markings in the form of white or black caps, collars, eye-marks or frontal patches, examples of which are seen in the three species of African plovers figured on page 221.

Recognition marks during flight are very important for all birds which congregate in flocks or which migrate together; and it is essential that, while being as conspicuous as possible, the marks shall not interfere with the general protective tints of the species when at rest. Hence they usually consist of well-contrasted markings on the wings and tail, which are concealed during repose but become fully visible when the bird takes flight. Such markings are well seen in our four British species of shrikes, each having quite different white marks on the expanded wings and on the tail feathers; and the same is the case with our three species of Saxicola—the stone-chat, whin-chat, and wheat-ear—which are thus easily recognisable on the wing, especially when seen from above, as they would be by stragglers looking out for their companions. The figures opposite, of the wings of two African species of stone-curlew which are sometimes found in the same districts, well illustrates these specific recognition marks. Though not very greatly different to our eyes, they are no doubt amply so to the sharp vision of the birds themselves.

Besides the white patches on the primaries here shown, the secondary feathers are, in some cases, so coloured as to afford very distinctive markings during flight, as seen in the central secondary quills of two African coursers (Fig. 21).



Most characteristic of all, however, are the varied markings of the outer tail-feathers, whose purpose is so well shown by their being almost always covered during repose by the two middle feathers, which are themselves quite unmarked and protectively tinted like the rest of the upper surface of the body. The figures of the expanded tails of two species of East Asiatic snipe, whose geographical ranges overlap each other, will serve to illustrate this difference; which is frequently much greater and modified in an endless variety of ways (Fig. 22).

Numbers of species of pigeons, hawks, finches, warblers, ducks, and innumerable other birds possess this class of markings; and they correspond so exactly in general character with those of the mammalia, already described, that we cannot doubt they serve a similar purpose.[86]



Those birds which are inhabitants of tropical forests, and which need recognition marks that shall be at all times visible among the dense foliage, and not solely or chiefly during flight, have usually small but brilliant patches of colour on the head or neck, often not interfering with the generally protective character of their plumage. Such are the bright patches of blue, red, or yellow, by which the usually green Eastern barbets are distinguished; and similar bright patches of colour characterise the separate species of small green fruit-doves. To this necessity for specialisation in colour, by which each bird may easily recognise its kind, is probably due that marvellous variety in the peculiar beauties of some groups of birds. The Duke of Argyll, speaking of the humming birds, made the objection that "A crest of topaz is no better in the struggle for existence than a crest of sapphire. A frill ending in spangles of the emerald is no better in the battle of life than a frill ending in spangles of the ruby. A tail is not affected for the purposes of flight, whether its marginal or its central feathers are decorated with white;" and he goes on to urge that mere beauty and variety for their own sake are the only causes of these differences. But, on the principles here suggested, the divergence itself is useful, and must have been produced pari passu with the structural differences on which the differentiation of species depends; and thus we have explained the curious fact that prominent differences of colour often distinguish species otherwise very closely allied to each other.

Among insects, the principle of distinctive coloration for recognition has probably been at work in the production of the wonderful diversity of colour and marking we find everywhere, more especially among the butterflies and moths; and here its chief function may have been to secure the pairing together of individuals of the same species. In some of the moths this has been secured by a peculiar odour, which attracts the males to the females from a distance; but there is no evidence that this is universal or even general, and among butterflies, especially, the characteristic colour and marking, aided by size and form, afford the most probable means of recognition. That this is so is shown by the fact that "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;" while, according to Mr. Collingwood, in the Malay Archipelago, "a dead butterfly 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."[87] In a great number of insects, no doubt, form, motions, stridulating sounds, or peculiar odours, serve to distinguish allied species from each other, and this must be especially the case with nocturnal insects, or with those whose colours are nearly uniform and are determined by the need of protection; but by far the larger number of day-flying and active insects exhibit varieties of colour and marking, forming the most obvious distinction between allied species, and which have, therefore, in all probability been acquired in the process of differentiation for the purpose of checking the intercrossing of closely allied forms.[88]

Whether this principle extends to any of the less highly organised animals is doubtful, though it may perhaps have affected the higher mollusca. But in marine animals it seems probable that the colours, however beautiful, varied, and brilliant they may often be, are in most cases protective, assimilating them to the various bright-coloured seaweeds, or to some other animals which it is advantageous for them to imitate.[89]

Summary of the Preceding Exposition.

Before proceeding to discuss some of the more recondite phenomena of animal coloration, it will be well to consider for a moment the extent of the ground we have already covered. Protective coloration, in some of its varied forms, has not improbably modified the appearance of one-half of the animals living on the globe. The white of arctic animals, the yellowish tints of the desert forms, the dusky hues of crepuscular and nocturnal species, the transparent or bluish tints of oceanic creatures, represent a vast host in themselves; but we have an equally numerous body whose tints are adapted to tropical foliage, to the bark of trees, or to the soil or dead leaves on or among which they habitually live. Then we have the innumerable special adaptations to the tints and forms of leaves, or twigs, or flowers; to bark or moss; to rock or pebble; by which such vast numbers of the insect tribes obtain protection; and we have seen that these various forms of coloration are equally prevalent in the waters of the seas and oceans, and are thus coextensive with the domain of life upon the earth. The comparatively small numbers which possess "terrifying" or "alluring" coloration may be classed under the general head of the protectively coloured.

But under the next head—colour for recognition—we have a totally distinct category, to some extent antagonistic or complementary to the last, since its essential principle is visibility rather than concealment. Yet it has been shown, I think, that this mode of coloration is almost equally important, since it not only aids in the preservation of existing species and in the perpetuation of pure races, but was, perhaps, in its earlier stages, a not unimportant factor in their development. To it we owe most of the variety and much of the beauty in the colours of animals; it has caused at once bilateral symmetry and general permanence of type; and its range of action has been perhaps equally extensive with that of coloration for concealment.

Influence of Locality or of Climate on Colour.

Certain relations between locality and coloration have long been noticed. Mr. Gould observed that birds from inland or continental localities were more brightly coloured than those living near the sea-coast or on islands, and he supposed that the more brilliant atmosphere of the inland stations was the explanation of the phenomenon.[90] Many American naturalists have observed similar facts, and they assert that the intensity of the colours of birds and mammals increases from north to south, and also with the increase of humidity. This change is imputed by Mr. J.A. Allen to the direct action of the environment. He says: "In respect to the correlation of intensity of colour in animals with the degree of humidity, it would perhaps be more in accordance with cause and effect to express the law of correlation as a decrease of intensity of colour with a decrease of humidity, the paleness evidently resulting from exposure and the blanching effect of intense sunlight, and a dry, often intensely heated atmosphere. With the decrease of the aqueous precipitation the forest growth and the protection afforded by arborescent vegetation gradually also decreases, as of course does also the protection afforded by clouds, the excessively humid regions being also regions of extreme cloudiness, while the dry regions are comparatively cloudless districts."[91] Almost identical changes occur in birds, and are imputed by Mr. Allen to similar causes.

It will be seen that Mr. Gould and Mr. Allen impute opposite effects to the same cause, brilliancy or intensity of colour being due to a brilliant atmosphere according to the former, while paleness of colour is imputed by the latter to a too brilliant sun. According to the principles which have been established by the consideration of arctic, desert, and forest animals respectively, we shall be led to conclude that there has been no direct action in this case, but that the effects observed are due to the greater or less need of protection. The pale colour that is prevalent in arid districts is in harmony with the general tints of the surface; while the brighter tints or more intense coloration, both southward and in humid districts, are sufficiently explained by the greater shelter due to a more luxuriant vegetation and a shorter winter. The advocates of the theory that intensity of light directly affects the colours of organisms, are led into perpetual inconsistencies. At one time the brilliant colours of tropical birds and insects are imputed to the intensity of a tropical sun, while the same intensity of sunlight is now said to have a "bleaching" effect. The comparatively dull and sober hues of our northern fauna were once supposed to be the result of our cloudy skies; but now we are told that cloudy skies and a humid atmosphere intensify colour.

In my Tropical Nature (pp. 257-264) I have called attention to what is perhaps the most curious and decided relation of colour to locality which has yet been observed—the prevalence of white markings in the butterflies and birds of islands.

So many cases are adduced from so many different islands, both in the eastern and western hemisphere, that it is impossible to doubt the existence of some common cause; and it seems probable to me now, after a fuller consideration of the whole subject of colour, that here too we have one of the almost innumerable results of the principle of protective coloration. White is, as a rule, an uncommon colour in animals, but probably only because it is so conspicuous. Whenever it becomes protective, as in the case of arctic animals and aquatic birds, it appears freely enough; while we know that white varieties of many species occur occasionally in the wild state, and that, under domestication, white or parti-coloured breeds are freely produced. Now in all the islands in which exceptionally white-marked birds and butterflies have been observed, we find two features which would tend to render the conspicuous white markings less injurious—a luxuriant tropical vegetation, and a decided scarcity of rapacious mammals and birds. White colours, therefore, would not be eliminated by natural selection; but variations in this direction would bear their part in producing the recognition marks which are everywhere essential, and which, in these islands, need not be so small or so inconspicuous as elsewhere.

Concluding Remarks.

On a review of the whole subject, then, we must conclude that there is no evidence of the individual or prevalent colours of organisms being directly determined by the amount of light, or heat, or moisture, to which they are exposed; while, on the other hand, the two great principles of the need of concealment from enemies or from their prey, and of recognition by their own kind, are so wide-reaching in their application that they appear at first sight to cover almost the whole ground of animal coloration. But, although they are indeed wonderfully general and have as yet been very imperfectly studied, we are acquainted with other modes of coloration which have a different origin. These chiefly appertain to the very singular class of warning colours, from which arise the yet more extraordinary phenomena of mimicry; and they open up so curious a field of inquiry and present so many interesting problems, that a chapter must be devoted to them. Yet another chapter will be required by the subject of sexual differentiation of colour and ornament, as to the origin and meaning of which I have arrived at different conclusions from Mr. Darwin. These various forms of coloration having been discussed and illustrated, we shall be in a position to attempt a brief sketch of the fundamental laws which have determined the general coloration of the animal world.

FOOTNOTES:

[Footnote 65: Proceedings of the Royal Society, No. 243, 1886; Transactions of the Royal Society, vol. clxxviii. B. pp. 311-441.]

[Footnote 66: A Naturalist's Wanderings in the Eastern Archipelago, p. 460.]

[Footnote 67: Trans. Phil. Soc. (? of S. Africa), 1878, part iv, p. 27.]

[Footnote 68: Proc. Zool. Soc., 1862 p. 357.]

[Footnote 69: With reference to this general resemblance of insects to their environment the following remarks by Mr. Poulton are very instructive. He says: "Holding the larva of Sphinx ligustri in one hand and a twig of its food-plant in the other, the wonder we feel is, not at the resemblance but at the difference; we are surprised at the difficulty experienced in detecting so conspicuous an object. And yet the protection is very real, for the larvae will be passed over by those who are not accustomed to their appearance, although the searcher may be told of the presence of a large caterpillar. An experienced entomologist may also fail to find the larvae till after a considerable search. This is general protective resemblance, and it depends upon a general harmony between the appearance of the organism and its whole environment. It is impossible to understand the force of this protection for any larva, without seeing it on its food-plant and in an entirely normal condition. The artistic effect of green foliage is more complex than we often imagine; numberless modifications are wrought by varied lights and shadows upon colours which are in themselves far from uniform. In the larva of Papilio machaon the protection is very real when the larva is on the food-plant, and can hardly be appreciated at all when the two are apart." Numerous other examples are given in the chapter on "Mimicry and other Protective Resemblances among Animals," in my Contributions to the Theory of Natural Selection.]

[Footnote 70: The Naturalist in Nicaragua, p. 19.]

[Footnote 71: R. Meldola, in Proc. Zool. Soc., 1873, p. 155.]

[Footnote 72: Nature, vol. iii. p. 166.]

[Footnote 73: Trans. Ent. Soc. Lond., 1878, p. 185.]

[Footnote 74: Ibid. (Proceedings, p. xlii.)]

[Footnote 75: Wallace's Malay Archipelago, vol. i. p. 204 (fifth edition, p. 130), with figure.]

[Footnote 76: Moseley's Notes by a Naturalist on the Challenger.]

[Footnote 77: Proceedings of the Boston Soc. of Nat. Hist., vol. xiv. 1871.]

[Footnote 78: Nature, 1870, p. 376.]

[Footnote 79: A Naturalist's Wanderings in the Eastern Archipelago, p. 63.]

[Footnote 80: A beautiful drawing of this rare insect, Hymenopus bicornis (in the nymph or active pupa state), was kindly sent me by Mr. Wood-Mason, Curator of the Indian Museum at Calcutta. A species, very similar to it, inhabits Java, where it is said to resemble a pink orchid. Other Mantidae, of the genus Gongylus, have the anterior part of the thorax dilated and coloured either white, pink, or purple; and they so closely resemble flowers that, according to Mr. Wood-Mason, one of them, having a bright violet-blue prothoracic shield, was found in Pegu by a botanist, and was for a moment mistaken by him for a flower. See Proc. Ent. Soc. Lond., 1878, p. liii.]

[Footnote 81: C. Dixon, in Seebohm's History of British Birds, vol. ii. Introduction, p. xxvi. Many of the other examples here cited are taken from the same valuable work.]

[Footnote 82: See A.H.S. Lucas, in Proceedings of Royal Society of Victoria, 1887, p. 56.]

[Footnote 83: Professor Wm.H. Brewer of Yale College has shown that the white marks or the spots of domesticated animals are rarely symmetrical, but have a tendency to appear more frequently on the left side. This is the case with horses, cattle, dogs, and swine. Among wild animals the skunk varies considerably in the amount of white on the body, and this too was found to be usually greatest on the left side. A close examination of numerous striped or spotted species, as tigers, leopards, jaguars, zebras, etc., showed that the bilateral symmetry was not exact, although the general effect of the two sides was the same. This is precisely what we should expect if the symmetry is not the result of a general law of the organisation, but has been, in part at least, produced and preserved for the useful purpose of recognition by the animal's fellows of the same species, and especially by the sexes and the young. See Proc. of the Am. Ass. for Advancement of Science, vol. xxx. p. 246.]

[Footnote 84: Descent of Man, p. 542.]

[Footnote 85: It may be thought that such extremely conspicuous markings as those of the zebra would be a great danger in a country abounding with lions, leopards, and other beasts of prey; but it is not so. Zebras usually go in bands, and are so swift and wary that they are in little danger during the day. It is in the evening, or on moonlight nights, when they go to drink, that they are chiefly exposed to attack; and Mr. Francis Galton, who has studied these animals in their native haunts, assures me, that in twilight they are not at all conspicuous, the stripes of white and black so merging together into a gray tint that it is very difficult to see them at a little distance. We have here an admirable illustration of how a glaringly conspicuous style of marking for recognition may be so arranged as to become also protective at the time when protection is most needed; and we may also learn how impossible it is for us to decide on the inutility of any kind of coloration without a careful study of the habits of the species in its native country.]

[Footnote 86: The principle of colouring for recognition was, I believe, first stated in my article on "The Colours of Animals and Plants" in Macmillan's Magazine, and more fully in my volume on Tropical Nature. Subsequently Mrs. Barber gave a few examples under the head of "Indicative or Banner Colours," but she applied it to the distinctive colours of the males of birds, which I explain on another principle, though this may assist.]

[Footnote 87: Quoted by Darwin in Descent of Man, p. 317.]

[Footnote 88: In the American Naturalist of March 1888, Mr. J.E. Todd has an article on "Directive Coloration in Animals," in which he recognises many of the cases here referred to, and suggests a few others, though I think he includes many forms of coloration—as "paleness of belly and inner side of legs"—which do not belong to this class.]

[Footnote 89: For numerous examples of this protective colouring of marine animals see Moseley's Voyage of the Challenger, and Dr. E.S. Morse in Proc. of Bost. Soc. of Nat. Hist., vol. xiv. 1871.]

[Footnote 90: See Origin of Species, p. 107.]

[Footnote 91: The "Geographical Variation of North American Squirrels," Proc. Bost. Soc. of Nat. Hist., 1874, p. 284; and Mammals and Winter Birds of Florida, pp. 233-241.]



CHAPTER IX

WARNING COLORATION AND MIMICRY

The skunk as an example of warning coloration—Warning colours among insects—Butterflies—Caterpillars—Mimicry—How mimicry has been produced—Heliconidae—Perfection of the imitation—Other cases of mimicry among Lepidoptera—Mimicry among protected groups—Its explanation—Extension of the principle—Mimicry in other orders of insects—Mimicry among the vertebrata—Snakes—The rattlesnake and the cobra—Mimicry among birds—Objections to the theory of mimicry—Concluding remarks on warning colours and mimicry.



We have now to deal with a class of colours which are the very opposite of those we have hitherto considered, since, instead of serving to conceal the animals that possess them or as recognition marks to their associates, they are developed for the express purpose of rendering the species conspicuous. The reason of this is that the animals in question are either the possessors of some deadly weapons, as stings or poison fangs, or they are uneatable, and are thus so disagreeable to the usual enemies of their kind that they are never attacked when their peculiar powers or properties are known. It is, therefore, important that they should not be mistaken for defenceless or eatable species of the same class or order, since in that case they might suffer injury, or even death, before their enemies discovered the danger or the uselessness of the attack. They require some signal or danger-flag which shall serve as a warning to would-be enemies not to attack them, and they have usually obtained this in the form of conspicuous or brilliant coloration, very distinct from the protective tints of the defenceless animals allied to them.

The Skunk as illustrating Warning Coloration.

While staying a few days, in July 1887, at the Summit Hotel on the Central Pacific Railway, I strolled out one evening after dinner, and on the road, not fifty yards from the house, I saw a pretty little white and black animal with a bushy tail coming towards me. As it came on at a slow pace and without any fear, although it evidently saw me, I thought at first that it must be some tame creature, when it suddenly occurred to me that it was a skunk. It came on till within five or six yards of me, then quietly climbed over a dwarf wall and disappeared under a small outhouse, in search of chickens, as the landlord afterwards told me. This animal possesses, as is well known, a most offensive secretion, which it has the power of ejecting over its enemies, and which effectually protects it from attack. The odour of this substance is so penetrating that it taints, and renders useless, everything it touches, or in its vicinity. Provisions near it become uneatable, and clothes saturated with it will retain the smell for several weeks, even though they are repeatedly washed and dried. A drop of the liquid in the eyes will cause blindness, and Indians are said not unfrequently to lose their sight from this cause. Owing to this remarkable power of offence the skunk is rarely attacked by other animals, and its black and white fur, and the bushy white tail carried erect when disturbed, form the danger-signals by which it is easily distinguished in the twilight or moonlight from unprotected animals. Its consciousness that it needs only to be seen to be avoided gives it that slowness of motion and fearlessness of aspect which are, as we shall see, characteristic of most creatures so protected.

Warning Colours among Insects.

It is among insects that warning colours are best developed, and most abundant. We all know how well marked and conspicuous are the colours and forms of the stinging wasps and bees, no one of which in any part of the world is known to be protectively coloured like the majority of defenceless insects. Most of the great tribe of Malacoderms among beetles are distasteful to insect-eating animals. Our red and black Telephoridae, commonly called "soldiers and sailors," were found, by Mr. Jenner Weir, to be refused by small birds. These and the allied Lampyridae (the fireflies and glow-worms) in Nicaragua, were rejected by Mr. Belt's tame monkey and by his fowls, though most other insects were greedily eaten by them. The Coccinellidae or lady-birds are another uneatable group, and their conspicuous and singularly spotted bodies serve to distinguish them at a glance from all other beetles.

These uneatable insects are probably more numerous than is supposed, although we already know immense numbers that are so protected. The most remarkable are the three families of butterflies—Heliconidae, Danaidae, and Acraeidae—comprising more than a thousand species, and characteristic respectively of the three great tropical regions—South America, Southern Asia, and Africa. All these butterflies have peculiarities which serve to distinguish them from every other group in their respective regions. They all have ample but rather weak wings, and fly slowly; they are always very abundant; and they all have conspicuous colours or markings, so distinct from those of other families that, in conjunction with their peculiar outline and mode of flight, they can usually be recognised at a glance. Other distinctive features are, that their colours are always nearly the same on the under surface of their wings as on the upper; they never try to conceal themselves, but rest on the upper surfaces of leaves or flowers; and, lastly, they all have juices which exhale a powerful scent, so that when one kills them by pinching the body, the liquid that exudes stains the fingers yellow, and leaves an odour that can only be removed by repeated washings.

Now, there is much direct evidence to show that this odour, though not very offensive to us, is so to most insect-eating creatures. Mr. Bates observed that, when set out to dry, specimens of Heliconidae were less subject to the attacks of vermin; while both he and I noticed that they were not attacked by insect-eating birds or dragonflies, and that their wings were not found in the forest paths among the numerous wings of other butterflies whose bodies had been devoured. Mr. Belt once observed a pair of birds capturing insects for their young; and although the Heliconidae swarmed in the vicinity, and from their slow flight could have been easily caught, not one was ever pursued, although other butterflies did not escape. His tame monkey also, which would greedily munch up other butterflies, would never eat the Heliconidae. It would sometimes smell them, but always rolled them up in its hand and then dropped them.

We have also some corresponding evidence as to the distastefulness of the Eastern Danaidae. The Hon. Mr. Justice Newton, who assiduously collected and took notes upon the Lepidoptera of Bombay, informed Mr. Butler of the British Museum that the large and swift-flying butterfly Charaxes psaphon, was continually persecuted by the bulbul, so that he rarely caught a specimen of this species which had not a piece snipped out of the hind wings. He offered one to a bulbul which he had in a cage, and it was greedily devoured, whilst it was only by repeated persecution that he succeeded in inducing the bird to touch a Danais.[92]

Besides these three families of butterflies, there are certain groups of the great genus Papilio—the true swallow-tailed butterflies—which have all the characteristics of uneatable insects. They have a special coloration, usually red and black (at least in the females), they fly slowly, they are very abundant, and they possess a peculiar odour somewhat like that of the Heliconidae. One of these groups is common in tropical America, another in tropical Asia, and it is curious that, although not very closely allied, they have each the same red and black colours, and are very distinct from all the other butterflies of their respective countries. There is reason to believe also that many of the brilliantly coloured and weak-flying diurnal moths, like the fine tropical Agaristidae and burnet-moths, are similarly protected, and that their conspicuous colours serve as a warning of inedibility. The common burnet-moth (Anthrocera filipendula) and the equally conspicuous ragwort-moth (Euchelia jacobeae) have been proved to be distasteful to insect-eating creatures.

The most interesting and most conclusive example of warning coloration is, however, furnished by caterpillars, because in this case the facts have been carefully ascertained experimentally by competent observers. In the year 1866, when Mr. Darwin was collecting evidence as to the supposed effect of sexual selection in bringing about the brilliant coloration of the higher animals, he was struck by the fact that many caterpillars have brilliant and conspicuous colours, in the production of which sexual selection could have no place. We have numbers of such caterpillars in this country, and they are characterised not only by their gay colours but by not concealing themselves. Such are the mullein and the gooseberry caterpillars, the larvae of the spurge hawk-moth, of the buff-tip, and many others. Some of these caterpillars are wonderfully conspicuous, as in the case of that noticed by Mr. Bates in South America, which was four inches long, banded across with black and yellow, and with bright red head, legs, and tail. Hence it caught the eye of any one who passed by, even at the distance of many yards.

Mr. Darwin asked me to try and suggest some explanation of this coloration; and, having been recently interested in the question of the warning coloration of butterflies, I suggested that this was probably a similar case,—that these conspicuous caterpillars were distasteful to birds and other insect-eating creatures, and that their bright non-protective colours and habit of exposing themselves to view, enabled their enemies to distinguish them at a glance from the edible kinds and thus learn not to touch them; for it must be remembered that the bodies of caterpillars while growing are so delicate, that a wound from a bird's beak would be perhaps as fatal as if they were devoured.[93] At this time not a single experiment or observation had been made on the subject, but after I had brought the matter before the Entomological Society, two gentlemen, who kept birds and other tame animals, undertook to make experiments with a variety of caterpillars.

Mr. Jenner Weir was the first to experiment with ten species of small birds in his aviary, and he found that none of them would eat the following smooth-skinned conspicuous caterpillars—Abraxas grossulariata, Diloba caeruleocephala, Anthrocera filipendula, and Cucullia verbasci. He also found that they would not touch any hairy or spiny larvae, and he was satisfied that it was not the hairs or the spines, but the unpleasant taste that caused them to be rejected, because in one case a young smooth larva of a hairy species, and in another case the pupa of a spiny larva, were equally rejected. On the other hand, all green or brown caterpillars as well as those that resemble twigs were greedily devoured.[94]

Mr. A.G. Butler also made experiments with some green lizards (Lacerta viridis), which greedily ate all kinds of food, including flies of many kinds, spiders, bees, butterflies, and green caterpillars; but they would not touch the caterpillar of the gooseberry-moth (Abraxas grossulariata), or the imago of the burnet-moth (Anthrocera filipendula). The same thing happened with frogs. When the gooseberry caterpillars were first given to them, "they sprang forward and licked them eagerly into their mouths; no sooner, however, had they done so, than they seemed to become aware of the mistake that they had made, and sat with gaping mouths, rolling their tongues about, until they had got quit of the nauseous morsels, which seemed perfectly uninjured, and walked off as briskly as ever." Spiders seemed equally to dislike them. This and another conspicuous caterpillar (Halia wavaria) were rejected by two species—the geometrical garden spider (Epeira diadema) and a hunting spider.[95]

Some further experiments with lizards were made by Professor Weismann, quite confirming the previous observations; and in 1886 Mr. E.B. Poulton of Oxford undertook a considerable series of experiments, with many other species of larvae and fresh kinds of lizards and frogs. Mr. Poulton then reviewed the whole subject, incorporating all recorded facts, as well as some additional observations made by Mr. Jenner Weir in 1886. More than a hundred species of larvae or of perfect insects of various orders have now been made the subject of experiment, and the results completely confirm my original suggestion. In almost every case the protectively coloured larvae have been greedily eaten by all kinds of insectivorous animals, while, in the immense majority of cases, the conspicuous, hairy, or brightly coloured larvae have been rejected by some or all of them. In some instances the inedibility of the larvae extends to the perfect insect, but not in others. In the former cases the perfect insect is usually adorned with conspicuous colours, as the burnet and ragwort moths; but in the case of the buff-tip, the moth resembles a broken piece of rotten stick, yet it is partly inedible, being refused by lizards. It is, however, very doubtful whether these are its chief enemies, and its protective form and colour may be needed against insectivorous birds or mammals.

Mr. Samuel H. Scudder, who has largely bred North American butterflies, has found so many of the eggs and larvae destroyed by hymenopterous and dipterous parasites that he thinks at least nine-tenths, perhaps a greater proportion, never reach maturity. Yet he has never found any evidence that such parasites attack either the egg or the larva of the inedible Danais archippus, so that in this case the insect is distasteful to its most dangerous foes in all the stages of its existence, a fact which serves to explain its great abundance and its extension over almost the whole world.[96]

One case has been found of a protectively coloured larva,—one, moreover, which in all its habits shows that it trusts to concealment to escape its enemies—which was yet always rejected by lizards after they had seized it, evidently under the impression that from its colour it would be eatable. This is the caterpillar of the very common moth Mania typica; and Mr. Poulton thinks that, in this case, the unpleasant taste is an incidental result of some physiological processes in the organism, and is itself a merely useless character. It is evident that the insect would not conceal itself so carefully as it does if it had not some enemies, and these are probably birds or small mammals, as its food-plants are said to be dock and willow-herb, not suggestive of places frequented by lizards; and it has been found by experiment that lizards and birds have not always the same likes and dislikes. The case is interesting, because it shows that nauseous fluids sometimes occur sporadically, and may thus be intensified by natural selection when required for the purpose of protection. Another exceptional case is that of the very conspicuous caterpillar of the spurge hawk-moth (Deilephila euphorbiae), which was at once eaten by a lizard, although, as it exposes itself on its food-plant in the daytime and is very abundant in some localities, it must almost certainly be disliked by birds or by some animals who would otherwise devour it. If disturbed while feeding it is said to turn round with fury and eject a quantity of green liquid, of an acid and disagreeable smell similar to that of the spurge milk, only worse.[97]

These facts, and Mr. Poulton's evidence that some larvae rejected by lizards at first will be eaten if the lizards are very hungry, show that there are differences in the amount of the distastefulness, and render it probable that if other food were wanting many of these conspicuous insects would be eaten. It is the abundance of the eatable kinds that gives value to the inedibility of the smaller number; and this is probably the reason why so many insects rely on protective colouring rather than on the acquisition of any kind of defensive weapons. In the long run the powers of attack and defence must balance each other. Hence we see that even the powerful stings of bees and wasps only protect them against some enemies, since a tribe of birds, the bee-eaters, have been developed which feed upon them, and some frogs and lizards do so occasionally.

The preceding outline will sufficiently explain the characteristics of "warning coloration" and the end it serves in nature. There are many other curious modifications of it, but these will be best appreciated after we have discussed the remarkable phenomenon of "mimicry," which is bound up with and altogether depends upon "warning colour," and is in some cases the chief indication we have of the possession of some offensive weapon to secure the safety of the species imitated.

Mimicry.

This term has been given to a form of protective resemblance, in which one species so closely resembles another in external form and colouring as to be mistaken for it, although the two may not be really allied and often belong to distinct families or orders. One creature seems disguised in order to be made like another; hence the terms "mimic" and mimicry, which imply no voluntary action on the part of the imitator. It has long been known that such resemblances do occur, as, for example, the clear-winged moths of the families Sesiidae and Aegeriidae, many of which resemble bees, wasps, ichneumons, or saw-flies, and have received names expressive of the resemblance; and the parasitic flies (Volucella) which closely resemble bees, on whose larvae the larvae of the flies feed.

The great bulk of such cases remained, however, unnoticed, and the subject was looked upon as one of the inexplicable curiosities of nature, till Mr. Bates studied the phenomenon among the butterflies of the Amazon, and, on his return home, gave the first rational explanation of it.[98] The facts are, briefly, these. Everywhere in that fertile region for the entomologist the brilliantly coloured Heliconidae abound, with all the characteristics which I have already referred to when describing them as illustrative of "warning coloration." But along with them other butterflies were occasionally captured, which, though often mistaken for them, on account of their close resemblance in form, colour, and mode of flight, were found on examination to belong to a very distinct family, the Pieridae. Mr. Bates notices fifteen distinct species of Pieridae, belonging to the genera Leptalis and Euterpe, each of which closely imitates some one species of Heliconidae, inhabiting the same region and frequenting the same localities. It must be remembered that the two families are altogether distinct in structure. The larvae of the Heliconidae are tubercled or spined, the pupae suspended head downwards, and the imago has imperfect forelegs in the male; while the larvae of the Pieridae are smooth, the pupae are suspended with a brace to keep the head erect, and the forefeet are fully developed in both sexes. These differences are as large and as important as those between pigs and sheep, or between swallows and sparrows; while English entomologists will best understand the case by supposing that a species of Pieris in this country was coloured and shaped like a small tortoise-shell, while another species on the Continent was equally like a Camberwell beauty—so like in both cases as to be mistaken when on the wing, and the difference only to be detected by close examination. As an example of the resemblance, woodcuts are given of one pair in which the colours are simple, being olive, yellow, and black, while the very distinct neuration of the wings and form of the head and body can be easily seen.



Besides these Pieridae, Mr. Bates found four true Papilios, seven Erycinidae, three Castnias (a genus of day-flying moths), and fourteen species of diurnal Bombycidae, all imitating some species of Heliconidae which inhabited the same district; and it is to be especially noted that none of these insects were so abundant as the Heliconidae they resembled, generally they were far less common, so that Mr. Bates estimated the proportion in some cases as not one to a thousand. Before giving an account of the numerous remarkable cases of mimicry in other parts of the world, and between various groups of insects and of higher animals, it will be well to explain briefly the use and purport of the phenomenon, and also the mode by which it has been brought about.

How Mimicry has been Produced.

The fact has been now established that the Heliconidae possess an offensive odour and taste, which lead to their being almost entirely free from attack by insectivorous creatures; they possess a peculiar form and mode of flight, and do not seek concealment; while their colours—although very varied, ranging from deep blue-black, with white, yellow, or vivid red bands and spots, to the most delicate semitransparent wings adorned with pale brown or yellow markings—are yet always very distinctive, and unlike those of all the other families of butterflies in the same country. It is, therefore, clear that if any other butterflies in the same region, which are eatable and suffer great persecution from insectivorous animals, should come to resemble any of these uneatable species so closely as to be mistaken for them by their enemies, they will obtain thereby immunity from persecution. This is the obvious and sufficient reason why the imitation is useful, and therefore why it occurs in nature. We have now to explain how it has probably been brought about, and also why a still larger number of persecuted groups have not availed themselves of this simple means of protection.

From the great abundance of the Heliconidae[99] all over tropical America, the vast number of their genera and species, and their marked distinctions from all other butterflies, it follows that they constitute a group of high antiquity, which in the course of ages has become more and more specialised, and owing to its peculiar advantages has now become a dominant and aggressive race. But when they first arose from some ancestral species or group which, owing to the food of the larvae or some other cause, possessed disagreeable juices that caused them to be disliked by the usual enemies of their kind, they were in all probability not very different either in form or coloration from many other butterflies. They would at that time be subject to repeated attacks by insect-eaters, and, even if finally rejected, would often receive a fatal injury. Hence arose the necessity for some distinguishing mark, by which the devourers of butterflies in general might learn that these particular butterflies were uneatable; and every variation leading to such distinction, whether by form, colour, or mode of flight, was preserved and accumulated by natural selection, till the ancestral Heliconoids became well distinguished from eatable butterflies, and thenceforth comparatively free from persecution. Then they had a good time of it. They acquired lazy habits, and flew about slowly. They increased abundantly and spread all over the country, their larvae feeding on many plants and acquiring different habits; while the butterflies themselves varied greatly, and colour being useful rather than injurious to them, gradually diverged into the many coloured and beautifully varied forms we now behold.

But, during the early stages of this process, some of the Pieridae, inhabiting the same district, happened to be sufficiently like some of the Heliconidae to be occasionally mistaken for them. These, of course, survived while their companions were devoured. Those among their descendants that were still more like Heliconidae again survived, and at length the imitation would become tolerably perfect. Thereafter, as the protected group diverged into distinct species of many different colours, the imitative group would occasionally be able to follow it with similar variations,—a process that is going on now, for Mr. Bates informs us that in each fresh district he visited he found closely allied representative species or varieties of Heliconidae, and along with them species of Leptalis (Pieridae), which had varied in the same way so as still to be exact imitations. But this process of imitation would be subject to check by the increasing acuteness of birds and other animals which, whenever the eatable Leptalis became numerous, would surely find them out, and would then probably attack both these and their friends the Heliconidae in order to devour the former and reject the latter. The Pieridae would, however, usually be less numerous, because their larvae are often protectively coloured and therefore edible, while the larvae of the Heliconidae are adorned with warning colours, spines, or tubercles, and are uneatable. It seems probable that the larvae and pupae of the Heliconidae were the first to acquire the protective distastefulness, both because in this stage they are more defenceless and more liable to fatal injury, and also because we now find many instances in which the larvae are distasteful while the perfect insects are eatable, but I believe none in which the reverse is the case. The larvae of the Pieridae are now beginning to acquire offensive juices, but have not yet obtained the corresponding conspicuous colours; while the perfect insects remain eatable, except perhaps in some Eastern groups, the under sides of whose wings are brilliantly coloured although this is the part which is exposed when at rest.

It is clear that if a large majority of the larvae of Lepidoptera, as well as the perfect insects, acquired these distasteful properties, so as seriously to diminish the food supply of insectivorous and nestling birds, these latter would be forced by necessity to acquire corresponding tastes, and to eat with pleasure what some of them now eat only under pressure of hunger; and variation and natural selection would soon bring about this change.

Many writers have denied the possibility of such wonderful resemblances being produced by the accumulation of fortuitous variations, but if the reader will call to mind the large amount of variability that has been shown to exist in all organisms, the exceptional power of rapid increase possessed by insects, and the tremendous struggle for existence always going on, the difficulty will vanish, especially when we remember that nature has the same fundamental groundwork to act upon in the two groups, general similarity of forms, wings of similar texture and outline, and probably some original similarity of colour and marking. Yet there is evidently considerable difficulty in the process, or with these great resources at her command nature would have produced more of these mimicking forms than she has done. One reason of this deficiency probably is, that the imitators, being always fewer in number, have not been able to keep pace with the variations of the much more numerous imitated form; another reason may be the ever-increasing acuteness of the enemies, which have again and again detected the imposture and exterminated the feeble race before it has had time to become further modified. The result of this growing acuteness of enemies has been, that those mimics that now survive exhibit, as Mr. Bates well remarks, "a palpably intentional likeness that is perfectly staggering," and also "that those features of the portrait are most attended to by nature which produce the most effective deception when the insects are seen in nature." No one, in fact, can understand the perfection of the imitation who has not seen these species in their native wilds. So complete is it in general effect that in almost every box of butterflies, brought from tropical America by amateurs, are to be found some species of the mimicking Pieridae, Erycinidae, or moths, and the mimicked Heliconidae, placed together under the impression that they are the same species. Yet more extraordinary, it sometimes deceives the very insects themselves. Mr. Trimen states that the male Danais chrysippus is sometimes deceived by the female Diadema bolina which mimics that species. Dr. Fritz Mueller, writing from Brazil to Professor Meldola, says, "One of the most interesting of our mimicking butterflies is Leptalis melite. The female alone of this species imitates one of our common white Pieridae, which she copies so well that even her own male is often deceived; for I have repeatedly seen the male pursuing the mimicked species, till, after closely approaching and becoming aware of his error, he suddenly returned."[100] This is evidently not a case of true mimicry, since the species imitated is not protected; but it may be that the less abundant Leptalis is able to mingle with the female Pieridae and thus obtain partial immunity from attack. Mr. Kirby of the insect department of the British Museum informs me that there are several species of South American Pieridae which the female Leptalis melite very nearly resembles. The case, however, is interesting as showing that the butterflies are themselves deceived by a resemblance which is not so great as that of some mimicking species.

Other Examples of Mimicry among Lepidoptera.

In tropical Asia, and eastward to the Pacific Islands, the Danaidae take the place of the Heliconidae of America, in their abundance, their conspicuousness, their slow flight, and their being the subjects of mimicry. They exist under three principal forms or genera. The genus Euploea is the most abundant both in species and individuals, and consists of fine broad-winged butterflies of a glossy or metallic blue-black colour, adorned with pure white, or rich blue, or dusky markings situated round the margins of the wings. Danais has generally more lengthened wings, of a semitransparent greenish or a rich brown colour, with radial or marginal pale spots; while the fine Hestias are of enormous size, of a papery or semitransparent white colour, with dusky or black spots and markings. Each of these groups is mimicked by various species of the genus Papilio, usually with such accuracy that it is impossible to distinguish them on the wing.[101] Several species of Diadema, a genus of butterflies allied to our Vanessas, also mimic species of Danais, but in this case the females only are affected, a subject which will be discussed in another chapter.

Another protected group in the Eastern tropics is that of the beautiful day-flying moths forming the family Agaristidae. These are usually adorned with the most brilliant colours or conspicuous markings, they fly slowly in forests among the butterflies and other diurnal insects, and their great abundance sufficiently indicates their possession of some distastefulness which saves them from attack. Under these conditions we may expect to find other moths which are not so protected imitating them, and this is the case. One of the common and wide-ranging species (Opthalmis lincea), found in the islands from Amboyna to New Ireland, is mimicked in a wonderful manner by one of the Liparidae (the family to which our common "tussock" and "vapourer" moths belong). This is a new species collected at Amboyna during the voyage of the Challenger, and has been named Artaxa simulans. Both insects are black, with the apex of the fore wings ochre coloured, and the outer half of the hind wings bright orange. The accompanying woodcuts (for the use of which I am indebted to Mr. John Murray of the Challenger Office) well exhibit their striking resemblance to each other.



In Africa exactly similar phenomena recur, species of Papilio and of Diadema mimicking Danaidae or Acraeidae with the most curious accuracy. Mr. Trimen, who studied this subject in South Africa, has recorded eight species or varieties of Diadema, and eight of Papilio, which each mimic some species of Danais; while eight species or varieties of Panopaea (another genus of Nymphalidae), three of Melanitis (Eurytelidae), and two of Papilio, resemble with equal accuracy some species of Acraea.[102] He has also independently observed the main facts on which the explanation of the phenomenon rests,—the unpleasant odour of the Danais and Acraea, extending to their larvae and pupae; their great abundance, slow flight, and disregard of concealment; and he states that while lizards, mantidae, and dragonflies all hunt butterflies, and the rejected wings are to be found abundantly at some of their feeding-places, those of the two genera Danais and Acraea were never among them.

The two groups of the great genus Papilio (the true swallow-tailed butterflies) which have been already referred to as having the special characteristics of uneatable insects, have also their imitators in other groups; and thus, the belief in their inedibility—derived mainly from their style of warning coloration and their peculiar habits—is confirmed. In South America, several species of the "Aeneas" group of these butterflies are mimicked by Pieridae and by day-flying moths of the genera Castnia and Pericopis. In the East, Papilio hector, P. diphilus, and P. liris, all belonging to the inedible group, are mimicked by the females of other species of Papilio belonging to very distinct groups; while in Northern India and China, many fine day-flying moths (Epicopeia) have acquired the strange forms and peculiar colours of some of the large inedible Papilios of the same regions.

In North America, the large and handsome Danais archippus, with rich reddish-brown wings, is very common; and it is closely imitated by Limenitis misippus, a butterfly allied to our "white admiral," but which has acquired a colour quite distinct from that of the great bulk of its allies. In the same country there is a still more interesting case. The beautiful dark bronzy green butterfly, Papilio philenor, is inedible both in larva and perfect insect, and it is mimicked by the equally dark Limenitis ursula. There is also in the Southern and Western States a dark female form of the yellow Papilio turnus, which in all probability obtains protection from its general resemblance to P. philenor. Mr. W.H. Edwards has found, by extensive experiment, that both the dark and yellow females produce their own kinds, with very few exceptions; and he thinks that the dark form has the advantage in the more open regions and in the prairies, where insectivorous birds abound. But in open country the dark form would be quite as conspicuous as the yellow form, if not more so, so that the resemblance to an inedible species would be there more needed.[103]

The only probable case of mimicry in this country is that of the moth, Diaphora mendica, whose female only is white, while the larva is of protective colours, and therefore almost certainly edible. A much more abundant moth, of about the same size and appearing about the same time, is Spilosoma menthrasti, also white, but in this case both it and its larva have been proved to be inedible. The white colour of the female Diaphora, although it must be very conspicuous at night, may, therefore, have been acquired in order to resemble the uneatable Spilosoma, and thus gain some protection.[104]

Mimicry among Protected (Uneatable) Genera.

Before giving some account of the numerous other cases of warning colours and of mimicry that occur in the animal kingdom, it will be well to notice a curious phenomenon which long puzzled entomologists, but which has at length received a satisfactory explanation.

We have hitherto considered, that mimicry could only occur when a comparatively scarce and much persecuted species obtained protection by its close external resemblance to a much more abundant uneatable species inhabiting its own district; and this rule undoubtedly prevails among the great majority of mimicking species all over the world. But Mr. Bates also found a number of pairs of species of different genera of Heliconidae, which resembled each other quite as closely as did the other mimicking species he has described; and since all these insects appear to be equally protected by their inedibility, and to be equally free from persecution, it was not easy to see why this curious resemblance existed, or how it had been brought about. That it is not due to close affinity is shown by the fact that the resemblance occurs most frequently between the two distinct sub-families into which (as Mr. Bates first pointed out) the Heliconidae are naturally divided on account of very important structural differences. One of these sub-families (the true Heliconinae) consists of two genera only, Heliconius and Eueides, the other (the Danaoid Heliconinae) of no less than sixteen genera; and, in the instances of mimicry we are now discussing, one of the pairs or triplets that resemble each other is usually a species of the large and handsome genus Heliconius, the others being species of the genera Mechanitis, Melinaea, or Tithorea, though several species of other Danaoid genera also imitate each other. The following lists will give some idea of the number of these curious imitative forms, and of their presence in every part of the Neotropical area. The bracketed species are those that resemble each other so closely that the difference is not perceptible when they are on the wing.

In the Lower Amazon region are found—

{ Heliconius sylvana. { Melinaea egina.

{ Heliconius numata. { Melinaea mneme. { Tithorea harmonia.

{ Methona psidii. { Thyridia ino.

{ Ceratina ninonia. { Melinaea mnasias.

In Central America are found—

{ Heliconius zuleika. Nicaragua { Melinaea hezia. { Mechanitis sp.

{ Heliconius formosus. { Tithorea penthias.

Guatemala { Heliconius telchina. { Melinaea imitata.



In the Upper Amazon region—

{ Heliconius pardalinus. { Melinaea pardalis.

{ Heliconius aurora. { Melinaea lucifer.

In New Grenada—

{ Heliconius ismenius. { Melinaea messatis.

{ Heliconius messene. { Melinaea mesenina. { (?) Mechanitis sp.

{ Heliconius hecalesia. { Tithorea hecalesina.

{ Heliconius hecuba. { Tithorea bonplandi.

In Eastern Peru and Bolivia—

{ Heliconius aristona. { Melinaea cydippe. { (?) Mechanitis mothone.

In Pernambuco—

{ Heliconius ethra. { Mechanitis nesaea.

In Rio Janeiro—

{ Helieonius eucrate. { Mechanitis lysimnia.

In South Brazil—

{ Thyridia megisto. { Ituna ilione.

{ Acraea thalia. { Eueides pavana.

Besides these, a number of species of Ithomia and Napeogenes, and of Napeogenes and Mechanitis, resemble each other with equal accuracy, so that they are liable to be mistaken for each other when on the wing; and no doubt many other equally remarkable cases are yet unnoticed.



The figures above of the fore and hind wings of two of these mimicking species, from Dr. Fritz Mueller's original paper in Kosmos, will serve to show the considerable amount of difference, in the important character of the neuration of the wings, between these butterflies, which really belong to very distinct and not at all closely allied genera. Other important characters are—(1) The existence of a small basal cell in the hind wings of Ituna which is wanting in Thyridia; (2) the division of the cell between the veins 1b and 2 of the hind wings in the former genus, while it is undivided in the latter; and (3) the existence in Thyridia of scent-producing tufts of hair on the upper edge of the hind wing, while in Ituna these are wanting; but in place of them are extensible processes at the end of the abdomen, also emitting a powerful scent. These differences characterise two marked subdivisions of the Danaoid Heliconinae, each containing several distinct genera; and these subdivisions are further distinguished by very different forms of larvae, that to which Ituna belongs having from two to four long threadlike tentacles on the back, while in that containing Thyridia these are always absent. The former usually feed on Asclepiadeae, the latter on Solanaceae or Scrophulariaceae.

The two species figured, though belonging to such distinct and even remote genera, have acquired almost identical tints and markings so as to be deceptively alike. The surface of the wings is, in both, transparent yellowish, with black transverse bands and white marginal spots, while both have similar black-and white-marked bodies and long yellow antennae. Dr. Mueller states that they both show a preference for the same flowers growing on the edges of the forest paths.[105]

We will now proceed to give the explanation of these curious similarities, which have remained a complete puzzle for twenty years. Mr. Bates, when first describing them, suggested that they might be due to some form of parallel variation dependent on climatic influences; and I myself adduced other cases of coincident local modifications of colour, which did not appear to be explicable by any form of mimicry.[106] But we neither of us hit upon the simple explanation given by Dr. Fritz Mueller in 1879.

His theory is founded on the assumed, but probable, fact, that insect-eating birds only learn by experience to distinguish the edible from the inedible butterflies, and in doing so necessarily sacrifice a certain number of the latter. The quantity of insectivorous birds in tropical America is enormous; and the number of young birds which every year have to learn wisdom by experience, as regards the species of butterflies to be caught or to be avoided, is so great that the sacrifice of life of the inedible species must be considerable, and, to a comparatively weak or scarce species, of vital importance. The number thus sacrificed will be fixed by the quantity of young birds, and by the number of experiences requisite to cause them to avoid the inedible species for the future, and not at all by the numbers of individuals of which each species consists. Hence, if two species are so much alike as to be mistaken for one another, the fixed number annually sacrificed by inexperienced birds will be divided between them, and both will benefit. But if the two species are very unequal in numbers, the benefit will be comparatively slight for the more abundant species, but very great for the rare one. To the latter it may make all the difference between safety and destruction.