Our difficulty in regard to sexual selection lies in understanding how it is that the males which conquer other males, or those which prove the most attractive to the females, leave a greater number of offspring to inherit their superiority than their beaten and less attractive rivals. Unless this result does follow, the characters which give to certain males an advantage over others could not be perfected and augmented through sexual selection. When the sexes exist in exactly equal numbers, the worst-endowed males will (except where polygamy prevails) ultimately find females, and leave as many offspring, as well fitted for their general habits of life, as the best-endowed males. From various facts and considerations, I formerly inferred that with most animals, in which secondary sexual characters are well developed, the males considerably exceeded the females in number; but this is not by any means always true. If the males were to the females as two to one, or as three to two, or even in a somewhat lower ratio, the whole affair would be simple; for the better-armed or more attractive males would leave the largest number of offspring. But, after investigating, as far as possible, the numerical proportion of the sexes, I do not believe that any great inequality in number commonly exists. In most cases sexual selection appears to have been effective in the following manner:
Let us take any species, a bird for instance, and divide the females inhabiting a district into two equal bodies, the one consisting of the more vigorous and better-nourished individuals, and the other of the less vigorous and healthy. The former, there can be little doubt, would be ready to breed in the spring before the others; and this is the opinion of Mr. Jenner Weir, who has carefully attended to the habits of birds during many years. There can also be no doubt that the most vigorous, best-nourished, and earliest breeders would on an average succeed in rearing the largest number of fine offspring. The males, as we have seen, are generally ready to breed before the females; the strongest, and with some species the best armed of the males, drive away the weaker; and the former would then unite with the more vigorous and better-nourished females, because they are the first to breed. Such vigorous pairs would surely rear a larger number of offspring than the retarded females, which would be compelled to unite with the conquered and less powerful males, supposing the sexes to be numerically equal; and this is all that is wanted to add, in the course of successive generations, to the size, strength, and courage of the males, or to improve their weapons.
COURTSHIP AMONG THE LOWER ANIMALS.
Page 214.
But in very many cases the males which conquer their rivals do not obtain possession of the females, independently of the choice of the latter. The courtship of animals is by no means so simple and short an affair as might be thought. The females are most excited by, or prefer pairing with, the more ornamented males, or those which are the best songsters, or play the best antics; but it is obviously probable that they would at the same time prefer the more vigorous and lively males, and this has in some cases been confirmed by actual observation. Thus, the more vigorous females, which are the first to breed, will have the choice of many males; and, though they may not always select the strongest or best armed, they will select those which are vigorous and well armed, and in other respects the most attractive. Both sexes, therefore, of such early pairs would, as above explained, have an advantage over others in rearing offspring; and this apparently has sufficed, during a long course of generations, to add not only to the strength and fighting powers of the males, but likewise to their various ornaments or other attractions.
In the converse and much rarer case, of the males selecting particular females, it is plain that those which were the most vigorous, and had conquered others, would have the freest choice; and it is almost certain that they would select vigorous as well as attractive females. Such pairs would have an advantage in rearing offspring, more especially if the male had the power to defend the female during the pairing-season, as occurs with some of the higher animals, or aided her in providing for the young. The same principles would apply if each sex preferred and selected certain individuals of the opposite sex; supposing that they selected not only the more attractive but likewise the more vigorous individuals.
WHY THE MALE PLAYS THE MORE ACTIVE PART IN COURTING.
Page 222.
We are naturally led to inquire why the male, in so many and such distinct classes, has become more eager than the female, so that he searches for her, and plays the more active part in courtship. It would be no advantage, and some loss of power, if each sex searched for the other; but why should the male almost always be the seeker? The ovules of plants after fertilization have to be nourished for a time; hence the pollen is necessarily brought to the female organs—being placed on the stigma by means of insects or the wind, or by the spontaneous movements of the stamens; and, in the Algæ, etc., by the locomotive power of the antherozoöids. With lowly-organized aquatic animals, permanently affixed to the same spot, and having their sexes separate, the male element is invariably brought to the female; and of this we can see the reason, for even if the ova were detached before fertilization, and did not require subsequent nourishment or protection, there would yet be greater difficulty in transporting them than the male element, because, being larger than the latter, they are produced in far smaller numbers. So that many of the lower animals are, in this respect, analogous with plants. The males of affixed and aquatic animals, having been led to emit their fertilizing element in this way, it is natural that any of their descendants, which rose in the scale and became locomotive, should retain the same habit; and they would approach the female as closely as possible, in order not to risk the loss of the fertilizing element in a long passage of it through the water. With some few of the lower animals, the females alone are fixed, and the males of these must be the seekers. But it is difficult to understand why the males of species, of which the progenitors were primordially free, should invariably have acquired the habit of approaching the females, instead of being approached by them. But, in all cases, in order that the males should seek efficiently, it would be necessary that they should be endowed with strong passions; and the acquirement of such passions would naturally follow from the more eager leaving a larger number of offspring than the less eager.