Contributions to the Theory of Natural Selection. Alfred Russel Wallace

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either spoke approvingly of it, or advocated similar principles, and for a good many years it was decidedly in the ascendent. With such a favourable introduction, and with such talented exponents, it must have become established if it had had any germ of truth in it; yet it quite died out in a few short years, its very existence is now a matter of history; and so rapid was its fall that its talented creator, Swainson, perhaps lived to be the last man who believed in it.

      Such is the course of a false theory. That of a true one is very different, as may be well seen by the progress of opinion on the subject of Natural Selection. In less than eight years “The Origin of Species” has produced conviction in the minds of a majority of the most eminent living men of science. New facts, new problems, new difficulties as they arise are accepted, solved or removed by this theory; and its principles are illustrated by the progress and conclusions of every well established branch of human knowledge. It is the object of the present essay to show how it has recently been applied to connect together and explain a variety of curious facts which had long been considered as inexplicable anomalies.

      Importance of the Principle of Utility

      Perhaps no principle has ever been announced so fertile in results as that which Mr. Darwin so earnestly impresses upon us, and which is indeed a necessary deduction from the theory of Natural Selection, namely—that none of the definite facts of organic nature, no special organ, no characteristic form or marking, no peculiarities of instinct or of habit, no relations between species or between groups of species—can exist, but which must now be or once have been useful to the individuals or the races which possess them. This great principle gives us a clue which we can follow out in the study of many recondite phænomena, and leads us to seek a meaning and a purpose of some definite character in minutiæ which we should be otherwise almost sure to pass over as insignificant or unimportant.

      Popular Theories of Colour in Animals

      The adaptation of the external colouring of animals to their conditions of life has long been recognised, and has been imputed either to an originally created specific peculiarity, or to the direct action of climate, soil, or food. Where the former explanation has been accepted, it has completely checked inquiry, since we could never get any further than the fact of the adaptation. There was nothing more to be known about the matter. The second explanation was soon found to be quite inadequate to deal with all the varied phases of the phænomena, and to be contradicted by many well-known facts. For example, wild rabbits are always of grey or brown tints well suited for concealment among grass and fern. But when these rabbits are domesticated, without any change of climate or food, they vary into white or black, and these varieties may be multiplied to any extent, forming white or black races. Exactly the same thing has occurred with pigeons; and in the case of rats and mice, the white variety has not been shown to be at all dependent on alteration of climate, food, or other external conditions. In many cases the wings of an insect not only assume the exact tint of the bark or leaf it is accustomed to rest on, but the form and veining of the leaf or the exact rugosity of the bark is imitated; and these detailed modifications cannot be reasonably imputed to climate or to food, since in many cases the species does not feed on the substance it resembles, and when it does, no reasonable connexion can be shown to exist between the supposed cause and the effect produced. It was reserved for the theory of Natural Selection to solve all these problems, and many others which were not at first supposed to be directly connected with them. To make these latter intelligible, it will be necessary to give a sketch of the whole series of phænomena which may be classed under the head of useful or protective resemblances.

      Importance of Concealment as Influencing Colour

      Concealment, more or less complete, is useful to many animals, and absolutely essential to some. Those which have numerous enemies from which they cannot escape by rapidity of motion, find safety in concealment. Those which prey upon others must also be so constituted as not to alarm them by their presence or their approach, or they would soon die of hunger. Now it is remarkable in how many cases nature gives this boon to the animal, by colouring it with such tints as may best serve to enable it to escape from its enemies or to entrap its prey. Desert animals as a rule are desert-coloured. The lion is a typical example of this, and must be almost invisible when crouched upon the sand or among desert rocks and stones. Antelopes are all more or less sandy-coloured. The camel is pre-eminently so. The Egyptian cat and the Pampas cat are sandy or earth-coloured. The Australian kangaroos are of the same tints, and the original colour of the wild horse is supposed to have been a sandy or clay-colour.

      The desert birds are still more remarkably protected by their assimilative hues. The stonechats, the larks, the quails, the goatsuckers and the grouse, which abound in the North African and Asiatic deserts, are all tinted and mottled so as to resemble with wonderful accuracy the average colour and aspect of the soil in the district they inhabit. The Rev. H. Tristram, in his account of the ornithology of North Africa in the 1st volume of the “Ibis,” says: “In the desert, where neither trees, brush-wood, nor even undulation 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.” After the testimony of so able an observer it is unnecessary to adduce further examples of the protective colours of desert animals.

      Almost equally striking are the cases of arctic animals possessing the white colour that best conceals them upon snowfields and icebergs. The polar bear is the only bear that is white, and it lives constantly among snow and ice. The arctic fox, the ermine and the alpine hare change to white in winter only, because in summer white would be more conspicuous than any other colour, and therefore a danger rather than a protection; but the American polar hare, inhabiting regions of almost perpetual snow, is white all the year round. Other animals inhabiting the same Northern regions do not, however, change colour. The sable is a good example, for throughout the severity of a Siberian winter it retains its rich brown fur. But its habits are such that it does not need the protection of colour, for it is said to be able to subsist on fruits and berries in winter, and to be so active upon the trees as to catch small birds among the branches. So also the woodchuck of Canada has a dark-brown fur; but then it lives in burrows and frequents river banks, catching fish and small animals that live in or near the water.

      Among birds, the ptarmigan is a fine example of protective colouring. Its summer plumage so exactly harmonizes with the lichen-coloured stones among which it delights to sit, that a person may walk through a flock of them without seeing a single bird; while in winter its white plumage is an almost equal protection. The snow-bunting, the jer-falcon, and the snowy owl are also white-coloured birds inhabiting the arctic regions, and there can be little doubt but that their colouring is to some extent protective.

      Nocturnal animals supply us with equally good illustrations. Mice, rats, bats, and moles possess the least conspicuous of hues, and must be quite invisible at times when any light colour would be instantly seen. Owls and goatsuckers are of those dark mottled tints that will assimilate with bark and lichen, and thus protect them during the day, and at the same time be inconspicuous in the dusk.

      It is only in the tropics, among forests which never lose their foliage, that we find whole groups of birds whose chief colour is green. The parrots are the most striking example, but we have also a group of green pigeons in the East; and the barbets, leaf-thrushes, bee-eaters, white-eyes, turacos, and several smaller groups, have so much green in their plumage as to tend greatly to conceal them among the foliage.

      Special Modifications of Colour

      The conformity of tint which has been so far shown to exist between animals and their habitations is of a somewhat general character; we will now consider the cases of more special adaptation. If the lion is enabled by his sandy colour readily to conceal himself

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