On the other hand, how remarkable it is that the only small and brightly coloured birds of our own country in which the male and female are alike—the tits and starlings—either build in holes or construct covered nests; while the beautiful hangnests (Icteridae) of South America, which always build covered or purse-shaped nests, are equally showy in both sexes, in striking contrast with the chatterers and tanagers of the same country, whose females are invariably less conspicuous than the males. On a rough estimate, there are about 1200 species of birds in the class of showy males and females, with concealed nidification; while there are probably, from an equally rough estimate, about the same number in the contrasted class of showy males and dull females, with open nests. This will leave the great bulk of known birds in the classes of those which are more or less protectively coloured in both sexes; or which, from their organisation and habits, do not require special protective coloration, such as many of the birds of prey, the larger waders, and the oceanic birds.
There are a few very curious cases in which the female bird is actually more brilliant than the male, and which yet have open nests. Such are the dotterel (Eudromias morinellus), several species of phalarope, an Australian creeper (Climacteris erythropus), and a few others; but in every one of these cases the relation of the sexes in regard to nidification is reversed, the male performing the duties of incubation, while the female is the stronger and more pugnacious. This curious case, therefore, quite accords with the general law of coloration.[124]
Sexual Colours of other Vertebrates.
We may consider a few of the cases of sexual colouring of other classes of vertebrates, as given by Mr. Darwin. In fishes, though the sexes are usually alike, there are several species in which the males are more brightly coloured, and have more elongated fins, spines, or other appendages, and in some few cases the colours are decidedly different. The males often fight together, and are altogether more vivacious and excitable than the females during the breeding season; and with this we may connect a greater intensity of coloration.
In frogs and toads the colours are usually alike, or a little more intense in the males, and the same may be said of most snakes. It is in lizards that we first meet with considerable sexual differences, many of the species having gular pouches, frills, dorsal crests, or horns, either confined to the males, or more developed in them than in the females, and these ornaments are often brightly coloured. In most cases, however, the tints of lizards are protective, the male being usually a little more intense in coloration; and the difference in extreme cases may be partly due to the need of protection for the female, which, when laden with eggs, must be less active and less able to escape from enemies than the male, and may, therefore, have retained more protective colours, as so many insects and birds have certainly done.[125]
In mammalia there is often a somewhat greater intensity of colour in the male, but rarely a decided difference. The female of the great red kangaroo, however, is a delicate gray; while in the Lemur macaco of Madagascar the male is jet-black and the female brown. In many monkeys also there are some differences of colour, especially on the face. The sexual weapons and ornaments of male mammalia, as horns, crests, manes, and dewlaps, are well known, and are very numerous and remarkable. Having thus briefly reviewed the facts, we will now consider the theories to which they have given rise.
Sexual Selection by the Struggles of Males.
Among the higher animals it is a very general fact that the males fight together for the possession of the females. This leads, in polygamous animals especially, to the stronger or better armed males becoming the parents of the next generation, which inherits the peculiarities of the parents; and thus vigour and offensive weapons are continually increased in the males, resulting in the strength and horns of the bull, the tusks of the boar, the antlers of the stag, and the spurs and fighting instinct of the gamecock. But almost all male animals fight together, though not specially armed; even hares, moles, squirrels, and beavers fight to the death, and are often found to be scarred and wounded. The same rule applies to almost all male birds; and these battles have been observed in such different groups as humming-birds, finches, goatsuckers, woodpeckers, ducks, and waders. Among reptiles, battles of the males are known to occur in the cases of crocodiles, lizards, and tortoises; among fishes, in those of salmon and sticklebats. Even among insects the same law prevails; and male spiders, beetles of many groups, crickets, and butterflies often fight together.
From this very general phenomenon there necessarily results a form of natural selection which increases the vigour and fighting power of the male animal, since, in every case, the weaker are either killed, wounded, or driven away. This selection would be more powerful if males were always in excess of females, but after much research Mr. Darwin could not obtain any satisfactory evidence that this was the case. The same effect, however, is produced in some cases by constitution or habits; thus male insects usually emerge first from the pupa, and among migrating birds the males arrive first both in this country and in North America. The struggle is thus intensified, and the most vigorous males are the first to have offspring. This in all probability is a great advantage, as the early breeders have the start in securing food, and the young are strong enough to protect themselves while the later broods are being produced.
It is to this form of male rivalry that Mr. Darwin first applied the term "sexual selection." It is evidently a real power in nature; and to it we must impute the development of the exceptional strength, size, and activity of the male, together with the possession of special offensive and defensive weapons, and of all other characters which arise from the development of these or are correlated with them. But he has extended the principle into a totally different field of action, which has none of that character of constancy and of inevitable result that attaches to natural selection, including male rivalry; for by far the larger portion of the phenomena, which he endeavours to explain by the direct action of sexual selection, can only be so explained on the hypothesis that the immediate agency is female choice or preference. It is to this that he imputes the origin of all secondary sexual characters other than weapons of offence and defence, of all the ornamental crests and accessory plumes of birds, the stridulating sounds of insects, the crests and beards of monkeys and other mammals, and the brilliant colours and patterns of male birds and butterflies. He even goes further, and imputes to it a large portion of the brilliant colour that occurs in both sexes, on the principle that variations occurring in one sex are sometimes transmitted to the same sex only, sometimes to both, owing to peculiarities in the laws of inheritance. In this extension of sexual selection to include the action of female choice or preference, and in the attempt to give to that choice such wide-reaching effects, I am unable to follow him more than a very little way; and I will now state some of the reasons why I think his views are unsound.