The Evolution Theory, Vol. 1 of 2

by August Weismann

This point has been disputed from many sides, and even by one of the founders of the whole selection theory, Alfred Russel Wallace. This naturalist doubts whether a choice is exercised among birds by either sex in regard to pairing, and maintains that, even if there could be a choice, this could not have produced such differences in colour and character of the plumage, since that would presuppose the existence of similar taste in the females through many generations. In a similar way it has been doubted whether butterflies can be said to exercise any real power of sexual choice, whether a more beautiful male is as such preferred to a less beautiful suitor.

It must be admitted that direct observation of choosing is difficult, and that as yet there is very little that can be said with certainty on this point. But there are, after all, some precise observations on mammals and birds which prove that the female shows active inclination to, or disinclination for, a particular male. If we hold fast to this fact, and add to it that the distinctive markings of the males are wonderfully developed during the period of courtship, and are displayed before the females, and that they only appear in mammals, birds, amphibians, and fishes at the time of sexual maturity, it seems to me that there can be no doubt that they are intended to fascinate the females, and to induce them to yield themselves to the males. The opponents of the theory of sexual selection attach too much importance to isolated cases; they imagine that each female must make a choice between several males. But the theory of sexual selection does not demand this, any more than the theory of natural selection requires the assumption that every individual of a species which is better equipped for the struggle for existence must necessarily survive and attain to reproduction, or, conversely, that the less well equipped must necessarily perish.

All that the theory requires is, that the selective and eliminative processes do, on an average, secure their ends, and in the same way the theory of sexual selection does not need the assumption that every female is in a position to exercise a scrupulous choice from among a troop of males, but only that, on an average, the males more agreeable to the females are selected, and those less agreeable rejected. If this is the case, it must result in the male characters most attractive to the females gaining preponderance, and becoming more and more firmly established in the species, increasing in intensity, and finally becoming a stable possession of all the males.

When we go more into details we shall see that the particular qualities of the distinctive masculine characters are exactly such as they would be if they owed their existence to processes of selection; in other words, from this point of view the phenomena of the decorative sexual characters can be understood up to a certain point. It seems to me that we are bound to accept the process of sexual selection as really operative, and instead of throwing doubt upon it, because the choice of the females can rarely be directly established, we should rather deduce from the numerous sexual characters of the males, which have a significance only in relation to courtship, that the females of the species are sensitive to these distinguishing characters, and are really capable of exercising a choice.

In my mind at least there remains no doubt that the 'sexual selection' of Darwin is an important factor in the transformation of species, even if I only take into consideration those secondary sexual characters which are related to wooing. We shall see, however, that there are others in regard to whose origin through processes of selection doubt is still less legitimate, and from which, on this account, we can argue back to the courtship characters.

The first beginning of transformation is not, even in ordinary natural selection, to be understood as due to selection, but is to be regarded as a given variation (the causes of which we shall discuss later on); it is only the increase of such incipient variations in a definite direction that can depend on natural selection, and they must depend on it in so far as the transformations are purposeful. Now, all secondary sexual characters can be recognized as useful, save only the decorative distinctions, although these also undoubtedly represent intensifications of originally unimportant variations. Are we then to regard these alone as the mere outcome of the internal impulsive forces of the organism, while in the case of the analogous sexual characters for tracking, catching, and holding the female, and so forth, the augmentation and the directing must be referred to processes of selection? But if there be any utility at all in the decorative sexual characters it can only lie in their greater attractiveness to the females, and it can only be of any account if the females have, in a certain sense, the power of choice. Independently, therefore, of direct observations as to the actual occurrence of choosing, we should be compelled by our chain of reasoning to assume that there was such a power of choice—and I shall immediately discuss it more precisely.

If we consider the decorative, distinctive characters of the males more closely, we find that they are of very diverse kinds. The males of many animals are distinguished from the females chiefly by greater beauty of form, and especially of colour. This is the case in many birds, some amphibians, like the water-salamander, many fishes, many insects, and above all, in diurnal Lepidoptera. Especially among birds the dimorphism between the sexes is in obvious relation to the excess in the number of male individuals, or—what practically comes to the same thing—to polygamy. For when a male attaches to himself four or ten females the result is the same as if the number of female individuals were divided by four or by ten. Thus the fowls and pheasants, which are polygamous, are adorned by magnificent colours in the male sex, while the monogamous partridges and quails exhibit the same colouring in both sexes. Of course 'beautiful' is a relative term, and we must not simply assume that what seems beautiful to us appears so to all animals; yet when we see that all the male birds which are beautifully decorative according to our taste—whether humming-birds, pheasants, birds of Paradise, or rock-cocks (Rupicola crocea)—unfold their 'feather-wheels, 'fans,' 'collars,' and so forth, before the eyes of the females in the breeding season, and display them in all their brilliance, we must conclude that, in these instances at least, human taste accords with that of the animals. That birds have sharp vision and distinguish colours is well known; it is not for nothing that the service berries and many other berries suitable for birds are red, the mistletoe berries white, in contrast to the evergreen foliage of this plant, the juniper berries black so that they stand out amid the snows of winter; in this direction, then, there is no difficulty in the way of sexual selection.

Even among much lower animals, like the butterflies, there seems to me no reason for the assumption that they do not see the gorgeous colours and often very complicated markings, the bars and eye-spots, on the wings of their fellows of the same species. Of course if each facet of the insect eye contributed only a single visual impression, as Johannes Müller supposed, then even an eye with 12,000 facets would give but a rough and ill-defined picture of objects more than a few feet away, and I confess that for a long time I regarded this as an obstacle in the way of referring the sexual dimorphism of butterflies to processes of selection. But we now know, through Exner, that this is not the case; we know that each facet gives a little picture, and not an 'inverted' but an 'upright' one, and experiment with the excised insect eye has directly shown that it throws on a photographic plate a tolerably clear image of even distant objects, such as the frame of a window, a large letter painted on the window, or even a church tower visible through it.

Furthermore, the structure of the eye allows of incomparably clearer vision of near objects, for in that case the eyes act like lenses, and reveal much more minute details than we ourselves are able to make out. Here again, therefore, there is no obstacle to the Darwinian hypothesis of a choice on the part of the females, for although it cannot be demonstrated from the structure of the eye itself that insects see colour, and that colours have a specially exciting influence on them, yet we can deduce this with certainty from the phenomena of their life. The butterflies fly to gaily coloured flowers, and as they find in them their food, the nectar of the flowers, we may take for granted that the sight of the colour of their food-providing plants is associated with an agreeable sensation, and this is an indication that similar colours in their fellows may awaken similar agreeable sensations.