Darwinism (Wallace)/Chapter VIII

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Darwinism (Wallace) by Alfred Russel Wallace
Chapter VIII - THE ORIGIN AND USES OF COLOUR IN ANIMALS

CHAPTER VIII

THE ORIGIN AND USES OF COLOUR IN ANIMALS

The Darwinian theory threw new light on organic colour—The problem to be solved—The constancy of animal colour indicates utility—Colour and environment—Arctic animals white—Exceptions prove the rule—Desert, forest, nocturnal, and oceanic animals—General theories of animal colour—Variable protective colouring—Mr. Poulton's experiments—Special or local colour adaptations—Imitation of particular objects—How they have been produced—Special protective colouring of butterflies—Protective resemblance among marine animals—Protection by terrifying enemies—Alluring coloration—The coloration of birds' eggs—Colour as a means of recognition—Summary of the preceding exposition—Influence of locality or of climate on colour—Concluding remarks.

Among the numerous applications of the Darwinian theory in the interpretation of the complex phenomena presented by the organic world, none have been more successful, or are more interesting, than those which deal with the colours of animals and plants. To the older school of naturalists colour was a trivial character, eminently unstable and untrustworthy in the determination of species; and it appeared to have, in most cases, no use or meaning to the objects which displayed it. The bright and often gorgeous coloration of insect, bird, or flower, was either looked upon as having been created for the enjoyment of mankind, or as due to unknown and perhaps undiscoverable laws of nature.

But the researches of Mr. Darwin totally changed our point of view in this matter. He showed, clearly, that some of the colours of animals are useful, some hurtful to them; and he believed that many of the most brilliant colours were developed by sexual choice; while his great general principle, that all the fixed characters of organic beings have been developed under the action of the law of utility, led to the inevitable conclusion that so remarkable and conspicuous a character as colour, which so often constitutes the most obvious distinction of species from species, or group from group, must also have arisen from survival of the fittest, and must, therefore, in most cases have some relation to the wellbeing of its possessors. Continuous observation and research, carried on by multitudes of observers during the last thirty years, have shown this to be the case; but the problem is found to be far more complex than was at first supposed. The modes in which colour is of use to different classes of organisms is very varied, and have probably not yet been all discovered; while the infinite variety and marvellous beauty of some of its developments are such as to render it hopeless to arrive at a complete and satisfactory explanation of every individual case. So much, however, has been achieved, so many curious facts have been explained, and so much light has been thrown on some of the most obscure phenomena of nature, that the subject deserves a prominent place in any account of the Darwinian theory.

The Problem to be Solved.

Before dealing with the various modifications of colour in the animal world it is necessary to say a few words on colour in general, on its prevalence in nature, and how it is that the colours of animals and plants require any special explanation. What we term colour is a subjective phenomenon, due to the constitution of our mind and nervous system; while, objectively, it consists of light-vibrations of different wave-lengths emitted by, or reflected from, various objects. Every visible object must be coloured, because to be visible it must send rays of light to our eye. The kind of light it sends is modified by the molecular constitution or the surface texture of the object. Pigments absorb certain rays and reflect the remainder, and this reflected portion has to our eyes a definite colour, according to the portion of the rays constituting white light which are absorbed. Interference colours are produced either by thin films or by very fine striae on the surfaces of bodies, which cause rays of certain wave-lengths to neutralise each other, leaving the remainder to produce the effects of colour. Such are the colours of soap-bubbles, or of steel or glass on which extremely fine lines have been ruled; and these colours often produce the effect of metallic lustre, and are the cause of most of the metallic hues of birds and insects.

As colour thus depends on molecular or chemical constitution or on the minute surface texture of bodies, and, as the matter of which organic beings are composed consists of chemical compounds of great complexity and extreme instability, and is also subject to innumerable changes during growth and development, we might naturally expect the phenomena of colour to be more varied here than in less complex and more stable compounds. Yet even in the inorganic world we find abundant and varied colours; in the earth and in the water; in metals, gems, and minerals; in the sky and in the ocean; in sunset clouds and in the many-tinted rainbow. Here we can have no question of use to the coloured object, and almost as little perhaps in the vivid red of blood, in the brilliant colours of red snow and other low algae and fungi, or even in the universal mantle of green which clothes so large a portion of the earth's surface. The presence of some colour, or even of many brilliant colours, in animals and plants would require no other explanation than does that of the sky or the ocean, of the ruby or the emerald—that is, it would require a purely physical explanation only. It is the wonderful individuality of the colours of animals and plants that attracts our attention—the fact that the colours are localised in definite patterns, sometimes in accordance with structural characters, sometimes altogether independent of them; while often differing in the most striking and fantastic manner in allied species. We are thus compelled to look upon colour not merely as a physical but also as a biological characteristic, which has been differentiated and specialised by natural selection, and must, therefore, find its explanation in the principle of adaptation or utility.

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 assimilates. 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 association 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 assimilative 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 passage: "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 assimilated 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."

Passing 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 illustration 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 illustration of general assimilation 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 Association 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 glass 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 butterflies, 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 class 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 class 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.[1] 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 butterflies (Pieris brassicae 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 butterflies 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-shell butterfly 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 assimilated 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-shell 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-shell larva, exposure to light from gilded surfaces produced pupae with a brilliant golden lustre; 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 classes 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 sexual differences) in all the individuals of the species. We will now briefly pass 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.

{{C|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 illustrative cases. Among our native birds we have the snipe and woodcock, 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 grassy 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 grass, 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 grass 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 grass, and one does not seem to notice stripes or spots till they are dead." It is the black shadows of the vegetation that assimilate 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."[2] 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 assimilated.

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 principal 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 mass 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."[3]

Some other cases will still further illustrate 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 cluster 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."[4]

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 coarse 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.[5]

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."[6]

Caterpillars also exhibit a considerable amount of detailed resemblance to the plants on which they live. Grass-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 grass-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.[7]

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 space of four inches in which it was to be seen, but none of them could perceive that it was a caterpillar."[8]

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 masses 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 illustrate 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 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.[9] 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.[10]

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.[11] 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."[12]

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.[13] 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."[14]

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 ziezae, 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.[15]

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.[16]

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 podargi— huge 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."[17] 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, the blackbird, and the lesser redpole 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.[18] 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.[19] 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."[20] 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.

FIG. 18.—Gazella soemmerringi.
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.[21]

FIG. 19—Recognition marks of three African plovers.
C. forbesi Charadrius bifrontatus C. tricollaris
Fig. 19.—Recognition marks of three African plovers
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,

FIG. 20. CEdicnemus venniculatus (above). CE. senegalensis (below).
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.[22]

FIG. 21.—Secondary quills.

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

FIG. 22.—Scolopax megala (upper). S. stenura (lower).
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."[23] 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.[24]

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.[25]

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.[26] 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."[27] 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.

  1. Proceedings of the Royal Society, No. 243, 1886; Transactions of the Royal Society, vol. clxxviii. B. pp. 311-441.
  2. A Naturalist's Wanderings in the Eastern Archipelago, p. 460.
  3. Trans. Phil. Soc. (? of S. Africa), 1878, part iv, p. 27.
  4. Proc. Zool. Soc., 1862 p. 357.
  5. 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.
  6. The Naturalist in Nicaragua, p. 19.
  7. R. Meldola, in Proc. Zool. Soc., 1873, p. 155.
  8. Nature, vol. iii. p. 166.
  9. Trans. Ent. Soc. Lond., 1878, p. 185.
  10. Ibid. (Proceedings, p. xlii.)
  11. Wallace's Malay Archipelago, vol. i. p. 204 (fifth edition, p. 130), with figure.
  12. Moseley's Notes by a Naturalist on the Challenger.
  13. Proceedings of the Boston Soc. of Nat. Hist., vol. xiv. 1871.
  14. Nature, 1870, p. 376.
  15. A Naturalist's Wanderings in the Eastern Archipelago, p. 63.
  16. 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.
  17. 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.
  18. See A.H.S. Lucas, in Proceedings of Royal Society of Victoria, 1887, p. 56.
  19. 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.
  20. Descent of Man, p. 542.
  21. 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.
  22. 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.
  23. Quoted by Darwin in Descent of Man, p. 317.
  24. 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.
  25. 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.
  26. See Origin of Species, p. 107.
  27. 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.