Darwinism (1889) / An exposition of the theory of natural selection, with some of its applications

by Alfred Russel Wallace

In another case which has been adduced as one of special difficulty, a more complete explanation is given. Soles, turbots, and other flatfish are, as is well known, unsymmetrical. They live and move on their sides, the under side being usually differently coloured from that which is kept uppermost. Now the eyes of these fish are curiously distorted in order that both eyes may be on the upper side, where alone they would be of any use. It was objected by Mr. Mivart that a sudden transformation of the eye from one side to the other was inconceivable, while, if the transit were gradual the first step could be of no use, since this would not remove the eye from the lower side. But, as Mr. Darwin shows by reference to the researches of Malm and others, the young of these fish are quite symmetrical, and during their growth exhibit to us the whole process of change. This begins by the fish (owing to the increasing depth of the body) being unable to maintain the vertical position, so that it falls on one side. It then twists the lower eye as much as possible towards the upper side; and, the whole bony structure of the head being at this time soft and flexible, the constant repetition of this effort causes the eye gradually to move round the head till it comes to the upper side. Now if we suppose this process, which in the young is completed in a few days or weeks, to have been spread over thousands of generations during the development of these fish, those usually surviving whose eyes retained more and more of the position into which the young fish tried to twist them, the change becomes intelligible; though it still remains one of the most extraordinary cases of degeneration, by which symmetry—which is so universal a characteristic of the higher animals—is lost, in order that the creature may be adapted to a new mode of life, whereby it is enabled the better to escape danger and continue its existence.

The most difficult case of all, that of the eye—the thought of which even to the last, Mr. Darwin says, "gave him a cold shiver"—is nevertheless shown to be not unintelligible; granting of course the sensitiveness to light of some forms of nervous tissue. For he shows that there are, in several of the lower animals, rudiments of eyes, consisting merely of pigment cells covered with a translucent skin, which may possibly serve to distinguish light from darkness, but nothing more. Then we have an optic nerve and pigment cells; then we find a hollow filled with gelatinous substance of a convex form—the first rudiment of a lens. Many of the succeeding steps are lost, as would necessarily be the case, owing to the great advantage of each modification which gave increased distinctness of vision, the creatures possessing it inevitably surviving, while those below them became extinct. But we can well understand how, after the first step was taken, every variation tending to more complete vision would be preserved till we reached the perfect eye of birds and mammals. Even this, as we know, is not absolutely, but only relatively, perfect. Neither the chromatic nor the spherical aberration is absolutely corrected; while long-and short-sightedness, and the various diseases and imperfections to which the eye is liable, may be looked upon as relics of the imperfect condition from which the eye has been raised by variation and natural selection.

These few examples of difficulties as to the origin of remarkable or complex organs must suffice here; but the reader who wishes further information on the matter may study carefully the whole of the sixth and seventh chapters of the last edition of The Origin of Species, in which these and many other cases are discussed in considerable detail.

Useless or non-adaptive Characters.

Many naturalists seem to be of opinion that a considerable number of the characters which distinguish species are of no service whatever to their possessors, and therefore cannot have been produced or increased by natural selection. Professors Bronn and Broca have urged this objection on the continent. In America, Dr. Cope, the well-known palaeontologist, has long since put forth the same objection, declaring that non-adaptive characters are as numerous as those which are adaptive; but he differs completely from most who hold the same general opinion in considering that they occur chiefly "in the characters of the classes, orders, families, and other higher groups;" and the objection, therefore, is quite distinct from that in which it is urged that "specific characters" are mostly useless. More recently, Professor G.J. Romanes has urged this difficulty in his paper on "Physiological Selection" (Journ. Linn. Soc., vol. xix. pp. 338, 344). He says that the characters "which serve to distinguish allied species are frequently, if not usually, of a kind with which natural selection can have had nothing to do," being without any utilitarian significance. Again he speaks of "the enormous number," and further on of "the innumerable multitude" of specific peculiarities which are useless; and he finally declares that the question needs no further arguing, "because in the later editions of his works Mr. Darwin freely acknowledges that a large proportion of specific distinctions must be conceded to be useless to the species presenting them."

I have looked in vain in Mr. Darwin's works to find any such acknowledgment, and I think Mr. Romanes has not sufficiently distinguished between "useless characters" and "useless specific distinctions." On referring to all the passages indicated by him I find that, in regard to specific characters, Mr. Darwin is very cautious in admitting inutility. His most pronounced "admissions" on this question are the following: "But when, from the nature of the organism and of the conditions, modifications have been induced which are unimportant for the welfare of the species, they may be, and apparently often have been, transmitted in nearly the same state to numerous, otherwise modified, descendants" (Origin, p. 175). The words I have here italicised clearly show that such characters are usually not "specific," in the sense that they are such as distinguish species from each other, but are found in numerous allied species. Again: "Thus a large yet undefined extension may safely be given to the direct and indirect results of natural selection; but I now admit, after reading the essay of Nägeli on plants, and the remarks by various authors with respect to animals, more especially those recently made by Professor Broca, that in the earlier editions of my Origin of Species I perhaps attributed too much to the action of natural selection or the survival of the fittest. I have altered the fifth edition of the Origin so as to confine my remarks to adaptive changes of structure, but I am convinced, from the light gained during even the last few years, that very many structures which now appear to us useless, will hereafter be proved to be useful, and will therefore come within the range of natural selection. Nevertheless I did not formerly consider sufficiently the existence of structures which, as far as we can at present judge, are neither beneficial nor injurious; and this I believe to be one of the greatest oversights as yet detected in my work." Now it is to be remarked that neither in these passages nor in any of the other less distinct expressions of opinion on this question, does Darwin ever admit that "specific characters"—that is, the particular characters which serve to distinguish one species from another—are ever useless, much less that "a large proportion of them" are so, as Mr. Romanes makes him "freely acknowledge." On the other hand, in the passage which I have italicised he strongly expresses his view that much of what we suppose to be useless is due to our ignorance; and as I hold myself that, as regards many of the supposed useless characters, this is the true explanation, it may be well to give a brief sketch of the progress of knowledge in transferring characters from the one category to the other.

We have only to go back a single generation, and not even the most acute botanist could have suggested a reasonable use, for each species of plant, of the infinitely varied forms, sizes, and colours of the flowers, the shapes and arrangement of the leaves, and the numerous other external characters of the whole plant. But since Mr. Darwin showed that plants gained both in vigour and in fertility by being crossed with other individuals of the same species, and that this crossing was usually effected by insects which, in search of nectar or pollen, carried the pollen from one plant to the flowers of another plant, almost every detail is found to have a purpose and a use. The shape, the size, and the colour of the petals, even the streaks and spots with which they are adorned, the position in which they stand, the movements of the stamens and pistil at various times, especially at the period of, and just after, fertilisation, have been proved to be strictly adaptive in so many cases that botanists now believe that all the external characters of flowers either are or have been of use to the species.

It has also been shown, by Kerner and other botanists, that another set of characteristics have relation to the prevention of ants, slugs, and other animals from reaching the flowers, because these creatures would devour or injure them without effecting fertilisation. The spines, hairs, or sticky glands on the stem or flower-stalk, the curious hairs or processes shutting up the flower, or sometimes even the extreme smoothness and polish of the outside of the petals so that few insects can hang to the part, have been shown to be related to the possible intrusion of these "unbidden guests."[42] And, still more recently, attempts have been made by Grant Allen and Sir John Lubbock to account for the innumerable forms, textures, and groupings of leaves, by their relation to the needs of the plants themselves; and there can be little doubt that these attempts will be ultimately successful. Again, just as flowers have been adapted to secure fertilisation or cross-fertilisation, fruits have been developed to assist in the dispersal of seeds; and their forms, sizes, juices, and colours can be shown to be specially adapted to secure such dispersal by the agency of birds and mammals; while the same end is secured in other cases by downy seeds to be wafted through the air, or by hooked or sticky seed-vessels to be carried away, attached to skin, wool, or feathers.

Here, then, we have an enormous extension of the region of utility in the vegetable kingdom, and one, moreover, which includes almost all the specific characters of plants. For the species of plants are usually characterised either by differences in the form, size, and colour of the flowers, or of the fruits; or, by peculiarities in the shape, size, dentation, or arrangement of the leaves; or by peculiarities in the spines, hairs, or down with which various parts of the plant are clothed. In the case of plants it must certainly be admitted that "specific" characters are pre-eminently adaptive; and though there may be some which are not so, yet all those referred to by Darwin as having been adduced by various botanists as useless, either pertain to genera or higher groups, or are found in some plants of a species only—that is, are individual variations not specific characters.

In the case of animals, the most recent wide extension of the sphere of utility has been in the matter of their colours and markings. It was of course always known that certain creatures gained protection by their resemblance to their normal surroundings, as in the case of white arctic animals, the yellow or brown tints of those living in deserts, and the green hues of many birds and insects surrounded by tropical vegetation. But of late years these cases have been greatly increased both in number and variety, especially in regard to those which closely imitate special objects among which they live; and there are other kinds of coloration which long appeared to have no use. Large numbers of animals, more especially insects, are gaudily coloured, either with vivid hues or with striking patterns, so as to be very easily seen. Now it has been found, that in almost all these cases the creatures possess some special quality which prevents their being attacked by the enemies of their kind whenever the peculiarity is known; and the brilliant or conspicuous colours or markings serve as a warning or signal flag against attack. Large numbers of insects thus coloured are nauseous and inedible; others, like wasps and bees, have stings; others are too hard to be eaten by small birds; while snakes with poisonous fangs often have some characteristic either of rattle, hood, or unusual colour, which indicates that they had better be left alone.