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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 mollifications 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 colow's 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 ly, 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 reflectei 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 striæ 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 algæ 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 permanently white or nearly so, 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 isa belline 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, bec-caters, 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 colow's 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

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