are usually correlated with variations of form or of colour. Hence, as fixed differences of form and colour, slowly gained by natural selection in adaptation to changed conditions, are what essentially characterise distinct species, some amount of infertility between species is the usual result.

Here the problem was left by Mr. Darwin; but we have shown that its solution may be carried a step further. If we accept the association of some degree of infertility, however slight, as a not unfrequent accompaniment of the external differences which always arise in a state of nature between varieties and incipient species, it has been shown that natural selection has power to increase that infertility just as it has power to increase other favourable variations. Such an increase of infertility will be beneficial, whenever new species arise in the same area with the parent form; and we thus see how, out of the fluctuating and very unequal amounts of infertility correlated with physical variations, there may have arisen that larger and more constant amount which appears usually to characterise well-marked species.

The great body of facts of which a condensed account has been given in the present chapter, although from an experimental point of view very insufficient, all point to the general conclusion we have now reached, and afford us a not unsatisfactory solution of the great problem of hybridism in relation to the origin of species by means of natural selection. Further experimental research is needed in order to complete the elucidation of the subject; but until these additional facts are forthcoming no new theory seems required for the explanation of the phenomena.

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 colourConcluding 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 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 alga 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, how