Darwinism (1889) Part 14

_The Constancy of Animal Colour indicates Utility._

That the colours and markings of animals have been acquired under the fundamental law of utility is indicated by a general fact which has received very little attention. As a rule, colour and marking are constant in each species of wild animal, while, in almost every domesticated animal, there arises great variability. We see this in our horses and cattle, our dogs and cats, our pigeons and poultry. Now, the essential difference between the conditions of life of domesticated and wild animals is, that the former are protected by man, while the latter have to protect themselves. The extreme variations in colour that immediately arise under domestication indicate a tendency to vary in this way, and the occasional occurrence of white or piebald or other exceptionally coloured individuals of many species in a state of nature, shows that this tendency exists there also; and, as these exceptionally coloured individuals rarely or never increase, there must be some constant power at work to keep it in check. This power can only be natural selection or the survival of the fittest, which again implies that some colours are useful, some injurious, in each particular case.

With this principle as our guide, let us see how far we can account both for the general and special colours of the animal world.

_Colour and Environment._

The fact that first strikes us in our examination of the colours of animals as a whole, is the close relation that exists between these colours and the general environment. Thus, white prevails among arctic animals; yellow or brown in desert species; while green is only a common colour in tropical evergreen forests. If we consider these cases somewhat carefully we shall find, that they afford us excellent materials for forming a judgment on the various theories that have been suggested to account for the colours of the animal world.

In the arctic regions there are a number of animals which are wholly white all the year round, or which only turn white in winter. Among the former are the polar bear and the American polar hare, the snowy owl and the Greenland falcon; among the latter the arctic fox, the arctic hare, the ermine, and the ptarmigan. Those which are permanently white remain among the snow nearly all the year round, while those which change their colour inhabit regions which are free from snow in summer. The obvious explanation of this style of coloration is, that it is protective, serving to conceal the herbivorous species from their enemies, and enabling carnivorous animals to approach their prey unperceived. Two other explanations have, however, been suggested. One is, that the prevalent white of the arctic regions has a direct effect in producing the white colour in animals, either by some photographic or chemical action on the skin or by a reflex action through vision. The other is, that the white colour is chiefly beneficial as a means of checking radiation and so preserving animal heat during the severity of an arctic winter. The first is part of the general theory that colour is the effect of coloured light on the objects--a pure hypothesis which has, I believe, no facts whatever to support it. The second suggestion is also an hypothesis merely, since it has not been proved by experiment that a white colour, _per se_, independently of the fur or feathers which is so coloured, has any effect whatever in checking the radiation of low-grade heat like that of the animal body. But both alike are sufficiently disproved by the interesting exceptions to the rule of white coloration in the arctic regions, which exceptions are, nevertheless, quite in harmony with the theory of protection.

Whenever we find arctic animals which, from whatever cause, do not require protection by the white colour, then neither the cold nor the snow-glare has any effect upon their coloration. The sable retains its rich brown fur throughout the Siberian winter; but it frequents trees at that season and not only feeds partially on fruits or seeds, but is able to catch birds among the branches of the fir-trees, with the bark of which its colour a.s.similates. Then we have that thoroughly arctic animal, the musk-sheep, which is brown and conspicuous; but this animal is gregarious, and its safety depends on its a.s.sociation in small herds.

It is, therefore, of more importance for it to be able to recognise its kind at a distance than to be concealed from its enemies, against which it can well protect itself so long as it keeps together in a compact body. But the most striking example is that of the common raven, which is a true arctic bird, and is found even in mid-winter as far north as any known bird or mammal. Yet it always retains its black coat, and the reason, from our point of view, is obvious. The raven is a powerful bird and fears no enemy, while, being a carrion-feeder, it has no need for concealment in order to approach its prey. The colour of the raven and of the musk-sheep are, therefore, both inconsistent with any other theory than that the white colour of arctic animals has been acquired for concealment, and to that theory both afford a strong support. Here we have a striking example of the exception proving the rule.

In the desert regions of the earth we find an even more general accordance of colour with surroundings. The lion, the camel, and all the desert antelopes have more or less the colour of the sand or rock among which they live. The Egyptian cat and the Pampas cat are sandy or earth coloured. The Australian kangaroos are of similar tints, and the original colour of the wild horse is supposed to have been sandy or clay coloured. Birds are equally well protected by a.s.similative hues; the larks, quails, goatsuckers, and grouse which abound in the North African and Asiatic deserts are all tinted or mottled so as closely to resemble the average colour of the soil in the districts they inhabit. Canon Tristram, who knows these regions and their natural history so well, says, in an often quoted pa.s.sage: "In the desert, where neither trees, brushwood, nor even undulations of the surface afford the slightest protection to its foes, a modification of colour which shall be a.s.similated to that of the surrounding country is absolutely necessary.

Hence, without exception, the upper plumage of every bird, whether lark, chat, sylvain, or sand-grouse, and also the fur of all the smaller mammals, and the skin of all the snakes and lizards, is of one uniform isabelline or sand colour."

Pa.s.sing on to the tropical regions, it is among their evergreen forests alone that we find whole groups of birds whose ground colour is green.

Parrots are very generally green, and in the East we have an extensive group of green fruit-eating pigeons; while the barbets, bee-eaters, turacos, leaf-thrushes (Phyllornis), white-eyes (Zosterops), and many other groups, have so much green in their plumage as to tend greatly to their concealment among the dense foliage. There can be no doubt that these colours have been acquired as a protection, when we see that in all the temperate regions, where the leaves are deciduous, the ground colour of the great majority of birds, especially on the upper surface, is a rusty brown of various shades, well corresponding with the bark, withered leaves, ferns, and bare thickets among which they live in autumn and winter, and especially in early spring when so many of them build their nests.

Nocturnal animals supply another ill.u.s.tration of the same rule, in the dusky colours of mice, rats, bats, and moles, and in the soft mottled plumage of owls and goatsuckers which, while almost equally inconspicuous in the twilight, are such as to favour their concealment in the daytime.

An additional ill.u.s.tration of general a.s.similation of colour to the surroundings of animals, is furnished by the inhabitants of the deep oceans. Professor Moseley of the Challenger Expedition, in his British a.s.sociation lecture on this subject, says: "Most characteristic of pelagic animals is the almost crystalline transparency of their bodies.

So perfect is this transparency that very many of them are rendered almost entirely invisible when floating in the water, while some, even when caught and held up in a gla.s.s globe, are hardly to be seen. The skin, nerves, muscles, and other organs are absolutely hyaline and transparent, but the liver and digestive tract often remain opaque and of a yellow or brown colour, and exactly resemble when seen in the water small pieces of floating seaweed." Such marine organisms, however, as are of larger size, and either occasionally or habitually float on the surface, are beautifully tinged with blue above, thus harmonising with the colour of the sea as seen by hovering birds; while they are white below, and are thus invisible against the wave-foam and clouds as seen by enemies beneath the surface. Such are the tints of the beautiful nudibranchiate mollusc, Glaucus atlanticus, and many others.

_General Theories of Animal Colour._

We are now in a position to test the general theories, or, to speak more correctly, the popular notions, as to the origin of animal coloration, before proceeding to apply the principle of utility to the explanation of some among the many extraordinary manifestations of colour in the animal world. The most generally received theory undoubtedly is, that brilliancy and variety of colour are due to the direct action of light and heat; a theory no doubt derived from the abundance of bright-coloured birds, insects, and flowers which are brought from tropical regions. There are, however, two strong arguments against this theory. We have already seen how generally bright coloration is wanting in desert animals, yet here heat and light are both at a maximum, and if these alone were the agents in the production of colour, desert animals should be the most brilliant. Again, all naturalists who have lived in tropical regions know that the proportion of bright to dull coloured species is little if any greater there than in the temperate zone, while there are many tropical groups in which bright colours are almost entirely unknown. No part of the world presents so many brilliant birds as South America, yet there are extensive families, containing many hundreds of species, which are as plainly coloured as our average temperate birds. Such are the families of the bush-shrikes and ant-thrushes (Formicariidae), the tyrant-shrikes (Tyrannidae), the American creepers (Dendrocolaptidae), together with a large proportion of the wood-warblers (Mniotiltidae), the finches, the wrens, and some other groups. In the eastern hemisphere, also, we have the babbling-thrushes (Timaliidae), the cuckoo-shrikes (Campephagidae), the honey-suckers (Meliphagidae), and several other smaller groups which are certainly not coloured above the average standard of temperate birds.

Again, there are many families of birds which spread over the whole world, temperate and tropical, and among these the tropical species rarely present any exceptional brilliancy of colour. Such are the thrushes, goatsuckers, hawks, plovers, and ducks; and in the last-named group it is the temperate and arctic zones that afford the most brilliant coloration.

The same general facts are found to prevail among insects. Although tropical insects present some of the most gorgeous coloration in the whole realm of nature, yet there are thousands and tens of thousands of species which are as dull coloured as any in our cloudy land. The extensive family of the carnivorous ground-beetles (Carabidae) attains its greatest brilliancy in the temperate zone; while by far the larger proportion of the great families of the longicorns and the weevils, are of obscure colours even in the tropics. In b.u.t.terflies, there is undoubtedly a larger proportion of brilliant colour in the tropics; but if we compare families which are almost equally developed over the globe--as the Pieridae or whites and yellows, and the Satyridae or ringlets--we shall find no great disproportion in colour between those of temperate and tropical regions.

The various facts which have now briefly been noticed are sufficient to indicate that the light and heat of the sun are not the direct causes of the colours of animals, although they may favour the production of colour when, as in tropical regions, the persistent high temperature favours the development of the maximum of life. We will now consider the next suggestion, that light reflected from surrounding coloured objects tends to produce corresponding colours in the animal world.

This theory is founded on a number of very curious facts which prove, that such a change does sometimes occur and is directly dependent on the colours of surrounding objects; but these facts are comparatively rare and exceptional in their nature, and the same theory will certainly not apply to the infinitely varied colours of the higher animals, many of which are exposed to a constantly varying amount of light and colour during their active existence. A brief sketch of these dependent changes of colour may, however, be advantageously given here.

_Variable Protective Colouring._

There are two distinct kinds of change of colour in animals due to the colouring of the environment. In one case the change is caused by reflex action set up by the animal _seeing_ the colour to be imitated, and the change produced can be altered or repeated as the animal changes its position. In the other case the change occurs but once, and is probably not due to any conscious or sense action, but to some direct influence on the surface tissues while the creature is undergoing a moult or change to the pupa form.

The most striking example of the first cla.s.s is that of the chameleon, which changes to white, brown, yellowish, or green, according to the colour of the object on which it rests. This change is brought about by means of two layers of pigment cells, deeply seated in the skin, and of bluish and yellowish colours. By suitable muscles these cells can be forced upwards so as to modify the colour of the skin, which, when they are not brought into action, is a dirty white. These animals are excessively sluggish and defenceless, and the power of changing their colour to that of their immediate surroundings is no doubt of great service to them. Many of the flatfish are also capable of changing their colour according to the colour of the bottom they rest on; and frogs have a similar power to a limited extent. Some crustacea also change colour, and the power is much developed in the Chameleon shrimp (Mysis Chamaeleon) which is gray when on sand, but brown or green when among brown or green seaweed. It has been proved by experiment that when this animal is blinded the change does not occur. In all these cases, therefore, we have some form of reflex or sense action by which the change is produced, probably by means of pigment cells beneath the skin as in the chameleon.

The second cla.s.s consists of certain larvae, and pupae, which undergo changes of colour when exposed to differently coloured surroundings.

This subject has been carefully investigated by Mr. E.B. Poulton, who has communicated the results of his experiments to the Royal Society.[65] It had been noticed that some species of larvae which fed on several different plants had colours more or less corresponding to the particular plant the individual fed on. Numerous cases are given in Professor Meldola's article on "Variable Protective Colouring" (_Proc.

Zool. Soc._, 1873, p. 153), and while the general green coloration was attributed to the presence of chlorophyll beneath the skin, the particular change in correspondence to each food-plant was attributed to a special function which had been developed by natural selection. Later on, in a note to his translation of Weissmann's _Theory of Descent_, Professor Meldola seemed disposed to think that the variations of colour of some of the species might be phytophagic--that is, due to the direct action of the differently coloured leaves on which the insect fed. Mr.

Poulton's experiments have thrown much light on this question, since he has conclusively proved that, in the case of the sphinx caterpillar of Smerinthus ocellatus, the change of colour is not due to the food but to the coloured light reflected from the leaves.

This was shown by feeding two sets of larvae on the same plant but exposed to differently coloured surroundings, obtained by sewing the leaves together, so that in one case only the dark upper surface, in the other the whitish under surface was exposed to view. The result in each case was a corresponding change of colour in the larvae, confirming the experiments on different individuals of the same batch of larvae which had been supplied with different food-plants or exposed to a different coloured light.

An even more interesting series of experiments was made on the colours of pupae, which in many cases were known to be affected by the material on which they underwent their transformations. The late Mr. T.W. Wood proved, in 1867, that the pupae of the common cabbage b.u.t.terflies (Pieris bra.s.sicae and P. rapae) were either light, or dark, or green, according to the coloured boxes they were kept in, or the colours of the fences, walls, etc., against which they were suspended. Mrs. Barber in South Africa found that the pupae of Papilio Nireus underwent a similar change, being deep green when attached to orange leaves of the same tint, pale yellowish-green when on a branch of the bottle-brush tree whose half-dried leaves were of this colour, and yellowish when attached to the wooden frame of a box. A few other observers noted similar phenomena, but nothing more was done till Mr. Poulton's elaborate series of experiments with the larvae of several of our common b.u.t.terflies were the means of clearing up several important points. He showed that the action of the coloured light did not affect the pupa itself but the larva, and that only for a limited period of time. After a caterpillar has done feeding it wanders about seeking a suitable place to undergo its transformation. When this is found it rests quietly for a day or two, spinning the web from which it is to suspend itself; and it is during this period of quiescence, and perhaps also the first hour or two after its suspension, that the action of the surrounding coloured surfaces determines, to a considerable extent, the colour of the pupa.

By the application of various surrounding colours during this period, Mr. Poulton was able to modify the colour of the pupa of the common tortoise-sh.e.l.l b.u.t.terfly from nearly black to pale, or to a brilliant golden; and that of Pieris rapae from dusky through pinkish to pale green. It is interesting to note, that the colours produced were in all cases such only as a.s.similated with the surroundings usually occupied by the species, and also, that colours which did not occur in such surroundings, as dark red or blue, only produced the same effects as dusky or black.

Careful experiments were made to ascertain whether the effect was produced through the sight of the caterpillar. The ocelli were covered with black varnish, but neither this, nor cutting off the spines of the tortoise-sh.e.l.l larva to ascertain whether they might be sense-organs, produced any effect on the resulting colour. Mr. Poulton concludes, therefore, that the colour-action probably occurs over the whole surface of the body, setting up physiological processes which result in the corresponding colour-change of the pupa. Such changes are, however, by no means universal, or even common, in protectively coloured pupae, since in Papilio machaon and some others which have been experimented on, both in this country and abroad, no change can be produced on the pupa by any amount of exposure to differently coloured surroundings. It is a curious point that, with the small tortoise-sh.e.l.l larva, exposure to light from gilded surfaces produced pupae with a brilliant golden l.u.s.tre; and the explanation is supposed to be that mica abounded in the original habitat of the species, and that the pupae thus obtained protection when suspended against micaceous rock. Looking, however, at the wide range of the species and the comparatively limited area in which micaceous rocks occur, this seems a rather improbable explanation, and the occurrence of this metallic appearance is still a difficulty. It does not, however, commonly occur in this country in a natural state.

The two cla.s.ses of variable colouring here discussed are evidently exceptional, and can have little if any relation to the colours of those more active creatures which are continually changing their position with regard to surrounding objects, and whose colours and markings are nearly constant throughout the life of the individual, and (with the exception of s.e.xual differences) in all the individuals of the species. We will now briefly pa.s.s in review the various characteristics and uses of the colours which more generally prevail in nature; and having already discussed those protective colours which serve to harmonise animals with their general environment, we have to consider only those cases in which the colour resemblance is more local or special in its character.

_Special or Local Colour Adaptations._

This form of colour adaptation is generally manifested by markings rather than by colour alone, and is extremely prevalent both among insects and vertebrates, so that we shall be able to notice only a few ill.u.s.trative cases. Among our native birds we have the snipe and woodc.o.c.k, whose markings and tints strikingly accord with the dead marsh vegetation among which they live; the ptarmigan in its summer dress is mottled and tinted exactly like the lichens which cover the stones of the higher mountains; while young unfledged plovers are spotted so as exactly to resemble the beach pebbles among which they crouch for protection, as beautifully exhibited in one of the cases of British birds in the Natural History Museum at South Kensington.

In mammalia, we notice the frequency of rounded spots on forest or tree haunting animals of large size, as the forest deer and the forest cats; while those that frequent reedy or gra.s.sy places are striped vertically, as the marsh antelopes and the tiger. I had long been of opinion that the brilliant yellow and black stripes of the tiger were adaptive, but have only recently obtained proof that it is so. An experienced tiger-hunter, Major Walford, states in a letter, that the haunts of the tiger are invariably full of the long gra.s.s, dry and pale yellow for at least nine months of the year, which covers the ground wherever there is water in the rainy season, and he adds: "I once, while following up a wounded tiger, failed for at least a minute to see him under a tree in gra.s.s at a distance of about twenty yards--jungle open--but the natives saw him, and I eventually made him out well enough to shoot him, but even then I could not see at what part of him I was aiming. There can be no doubt whatever that the colour of both the tiger and the panther renders them almost invisible, especially in a strong blaze of light, when among gra.s.s, and one does not seem to notice stripes or spots till they are dead." It is the black shadows of the vegetation that a.s.similate with the black stripes of the tiger; and, in like manner, the spotty shadows of leaves in the forest so harmonise with the spots of ocelots, jaguars, tiger-cats, and spotted deer as to afford them a very perfect concealment.

In some cases the concealment is effected by colours and markings which are so striking and peculiar that no one who had not seen the creature in its native haunts would imagine them to be protective. An example of this is afforded by the banded fruit pigeon of Timor, whose pure white head and neck, black wings and back, yellow belly, and deeply-curved black band across the breast, render it a very handsome and conspicuous bird. Yet this is what Mr. H.O. Forbes says of it: "On the trees the white-headed fruit pigeon (Ptilopus cinctus) sate motionless during the heat of the day in numbers, on well-exposed branches; but it was with the utmost difficulty that I or my sharp-eyed native servant could ever detect them, even in trees where we knew they were sitting."[66] The trees referred to are species of Eucalyptus which abound in Timor. They have whitish or yellowish bark and very open foliage, and it is the intense sunlight casting black curved shadows of one branch upon another, with the white and yellow bark and deep blue sky seen through openings of the foliage, that produces the peculiar combination of colours and shadows to which the colours and markings of this bird have become so closely a.s.similated.

Even such brilliant and gorgeously coloured birds as the sun-birds of Africa are, according to an excellent observer, often protectively coloured. Mrs. M.E. Barber remarks that "A casual observer would scarcely imagine that the highly varnished and magnificently coloured plumage of the various species of Noctarinea could be of service to them, yet this is undoubtedly the case. The most unguarded moments of the lives of these birds are those that are spent amongst the flowers, and it is then that they are less wary than at any other time. The different species of aloes, which blossom in succession, form the princ.i.p.al sources of their winter supplies of food; and a legion of other gay flowering plants in spring and summer, the aloe blossoms especially, are all brilliantly coloured, and they harmonise admirably with the gay plumage of the different species of sun-birds. Even the keen eye of a hawk will fail to detect them, so closely do they resemble the flowers they frequent. The sun-birds are fully aware of this fact, for no sooner have they relinquished the flowers than they become exceedingly wary and rapid in flight, darting arrow-like through the air and seldom remaining in exposed situations. The black sun-bird (Nectarinea amethystina) is never absent from that magnificent forest-tree, the 'Kaffir Boom' (Erythrina caffra); all day long the cheerful notes of these birds may be heard amongst its spreading branches, yet the general aspect of the tree, which consists of a huge ma.s.s of scarlet and purple-black blossoms without a single green leaf, blends and harmonises with the colours of the black sun-bird to such an extent that a dozen of them may be feeding amongst its blossoms without being conspicuous, or even visible."[67]

Some other cases will still further ill.u.s.trate how the colours of even very conspicuous animals may be adapted to their peculiar haunts.

The late Mr. Swinhoe says of the Kerivoula picta, which he observed in Formosa: "The body of this bat was of an orange colour, but the wings were painted with orange-yellow and black. It was caught suspended, head downwards, on a cl.u.s.ter of the fruit of the longan tree (Nephelium longanum). Now this tree is an evergreen, and all the year round some portion of its foliage is undergoing decay, the particular leaves being, in such a stage, partially orange and black. This bat can, therefore, at all seasons suspend from its branches and elude its enemies by its resemblance to the leaves of the tree."[68]

Even more curious is the case of the sloths--defenceless animals which feed upon leaves, and hang from the branches of trees with their back downwards. Most of the species have a curious buff-coloured spot on the back, rounded or oval in shape and often with a darker border, which seems placed there on purpose to make them conspicuous; and this was a great puzzle to naturalists, because the long coa.r.s.e gray or greenish hair was evidently like tree-moss and therefore protective. But an old writer, Baron von Slack, in his _Voyage_ _to Surinam_ (1810), had already explained the matter. He says: "The colour and even the shape of the hair are much like withered moss, and serve to hide the animal in the trees, but particularly when it has that orange-coloured spot between the shoulders and lies close to the tree; it looks then exactly like a piece of branch where the rest has been broken off, by which the hunters are often deceived." Even such a huge animal as the giraffe is said to be perfectly concealed by its colour and form when standing among the dead and broken trees that so often occur on the outskirts of the thickets where it feeds. The large blotch-like spots on the skin and the strange shape of the head and horns, like broken branches, so tend to its concealment that even the keen-eyed natives have been known to mistake trees for giraffes or giraffes for trees.

Innumerable examples of this kind of protective colouring occur among insects; beetles mottled like the bark of trees or resembling the sand or rock or moss on which they live, with green caterpillars of the exact general tints of the foliage they feed on; but there are also many cases of detailed imitation of particular objects by insects that must be briefly described.[69]

_Protective Imitation of Particular Objects._

The insects which present this kind of imitation most perfectly are the Phasmidae, or stick and leaf insects. The well-known leaf-insects of Ceylon and of Java, species of Phyllium, are so wonderfully coloured and veined, with leafy expansions on the legs and thorax, that not one person in ten can see them when resting on the food-plant close beneath their eyes. Others resemble pieces of stick with all the minutiae of knots and branches, formed by the insects' legs, which are stuck out rigidly and unsymmetrically. I have often been unable to distinguish between one of these insects and a real piece of stick, till I satisfied myself by touching it and found it to be alive. One species, which was brought me in Borneo, was covered with delicate semitransparent green foliations, exactly resembling the hepaticae which cover pieces of rotten stick in the damp forests. Others resemble dead leaves in all their varieties of colour and form; and to show how perfect is the protection obtained and how important it is to the possessors of it, the following incident, observed by Mr. Belt in Nicaragua, is most instructive. Describing the armies of foraging ants in the forest which devour every insect they can catch, he says: "I was much surprised with the behaviour of a green leaf-like locust. This insect stood immovably among a host of ants, many of which ran over its legs without ever discovering there was food within their reach. So fixed was its instinctive knowledge that its safety depended on its immovability, that it allowed me to pick it up and replace it among the ants without making a single effort to escape. This species closely resembles a green leaf."[70]

Caterpillars also exhibit a considerable amount of detailed resemblance to the plants on which they live. Gra.s.s-feeders are striped longitudinally, while those on ordinary leaves are always striped obliquely. Some very beautiful protective resemblances are shown among the caterpillars figured in Smith and Abbott's _Lepidopterous Insects of Georgia_, a work published in the early part of the century, before any theories of protection were started. The plates in this work are most beautifully executed from drawings made by Mr. Abbott, representing the insects, in every case, on the plants which they frequented, and no reference is made in the descriptions to the remarkable protective details which appear upon the plates. We have, first, the larva of Sphinx fuciformis feeding on a plant with linear gra.s.s-like leaves and small blue flowers; and we find the insect of the same green as the leaves, striped longitudinally in accordance with the linear leaves, and with the head blue corresponding both in size and colour with the flowers. Another species (Sphinx tersa) is represented feeding on a plant with small red flowers situated in the axils of the leaves; and the larva has a row of seven red spots, unequal in size, and corresponding very closely with the colour and size of the flowers. Two other figures of sphinx larvae are very curious. That of Sphinx pampinatrix feeds on a wild vine (Vitis indivisa), having green tendrils, and in this species the curved horn on the tail is green, and closely imitates in its curve the tip of the tendril. But in another species (Sphinx cranta), which feeds on the fox-grape (Vitis vulpina), the horn is very long and red, corresponding with the long red-tipped tendrils of the plant. Both these larvae are green with oblique stripes, to harmonise with the veined leaves of the vines; but a figure is also given of the last-named species after it has done feeding, when it is of a decided brown colour and has entirely lost its horn. This is because it then descends to the ground to bury itself, and the green colour and red horn would be conspicuous and dangerous; it therefore loses both at the last moult. Such a change of colour occurs in many species of caterpillars. Sometimes the change is seasonal; and, in those which hibernate with us, the colour of some species, which is brownish in autumn in adaptation to the fading foliage, becomes green in spring to harmonise with the newly-opened leaves at that season.[71]

Some of the most curious examples of minute imitation are afforded by the caterpillars of the geometer moths, which are always brown or reddish, and resemble in form little twigs of the plant on which they feed. They have the habit, when at rest, of standing out obliquely from the branch, to which they hold on by their hind pair of prolegs or claspers, and remain motionless for hours. Speaking of these protective resemblances Mr. Jenner Weir says: "After being thirty years an entomologist I was deceived myself, and took out my pruning scissors to cut from a plum tree a spur which I thought I had overlooked. This turned out to be the larva of a geometer two inches long. I showed it to several members of my family, and defined a s.p.a.ce of four inches in which it was to be seen, but none of them could perceive that it was a caterpillar."[72]

One more example of a protected caterpillar must be given. Mr. A.

Everett, writing from Sarawak, Borneo, says: "I had a caterpillar brought me, which, being mixed by my boy with some other things, I took to be a bit of moss with two exquisite pinky-white seed-capsules; but I soon saw that it moved, and examining it more closely found out its real character: it is covered with hair, with two little pink spots on the upper surface, the general hue being more green. Its motions are very slow, and when eating the head is withdrawn beneath a fleshy mobile hood, so that the action of feeding does not produce any movement externally. It was found in the limestone hills at Busan, the situation of all others where mosses are most plentiful and delicate, and where they partially clothe most of the protruding ma.s.ses of rock."

_How these Imitations have been Produced._

To many persons it will seem impossible that such beautiful and detailed resemblances as those now described--and these are only samples of thousands that occur in all parts of the world--can have been brought about by the preservation of accidental useful variations. But this will not seem so surprising if we keep in mind the facts set forth in our earlier chapters--the rapid multiplication, the severe struggle for existence, and the constant variability of these and all other organisms. And, further, we must remember that these delicate adjustments are the result of a process which has been going on for millions of years, and that we now see the small percentage of successes among the myriads of failures. From the very first appearance of insects and their various kinds of enemies the need of protection arose, and was usually most easily met by modifications of colour. Hence, we may be sure that the earliest leaf-eating insects acquired a green colour as one of the necessities of their existence; and, as the species became modified and specialised, those feeding on particular species of plants would rapidly acquire the peculiar tints and markings best adapted to conceal them upon those plants. Then, every little variation that, once in a hundred years perhaps, led to the preservation of some larva which was thereby rather better concealed than its fellows, would form the starting-point of a further development, leading ultimately to that perfection of imitation in details which now astonishes us. The researches of Dr. Weismann ill.u.s.trate this progressive adaptation. The very young larvae of several species are green or yellowish without any markings; they then, in subsequent moults, obtain certain markings, some of which are often lost again before the larva is fully grown. The early stages of those species which, like elephant hawk-moths (Chaerocampa), have the anterior segments elongated and retractile, with large eye-like spots to imitate the head of a vertebrate, are at first like those of non-retractile species, the anterior segments being as large as the rest. After the first moult they become smaller, comparatively; but it is only after the second moult that the ocelli begin to appear, and these are not fully defined till after the third moult. This progressive development of the individual--the ontogeny--gives us a clue to the ancestral development of the whole race--the phylogeny; and we are enabled to picture to ourselves the very slow and gradual steps by which the existing perfect adaptation has been brought about. In many larvae great variability still exists, and in some there are two or more distinctly-coloured forms--usually a dark and a light or a brown and a green form. The larva of the humming-bird hawk-moth (Macroglossa stellatarum) varies in this manner, and Dr. Weismann raised five varieties from a batch of eggs from one moth. It feeds on species of bedstraw (Galium verum and G. mollugo), and as the green forms are less abundant than the brown, it has probably undergone some recent change of food-plant or of habits which renders brown the more protective colour.

_Special Protective Colouring of b.u.t.terflies._

We will now consider a few cases of special protective colouring in the perfect b.u.t.terfly 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 b.u.t.terflies whose undersides are silvery, and which are thus effectually protected when at rest.[73] A common African b.u.t.terfly (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 b.u.t.terfly 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]