From our ignorance on several points, the precise manner in which sexual selection acts is to a certain extent uncertain. Nevertheless if those naturalists who already believe in the mutability of species, will read the following chapters, they will, I think, agree with me that sexual selection has played an important part in the history of the organic world. It is certain that with almost all animals there is a struggle between the males for the possession of the female. This fact is so notorious that it would be superfluous to give instances. Hence the females, supposing that their mental capacity sufficed for the exertion of a choice, could select one out of several males. But in numerous cases it appears as if it had been specially arranged that there should be a struggle between many males. Thus with migratory birds, the males generally arrive before the females at their place of breeding, so that many males are ready to contend for each female. The bird-catchers assert that this is invariably the case with the nightingale and blackcap, as I am informed by Mr. Jenner Weir, who confirms the statement with respect to the latter species.
Mr. Swaysland of Brighton, who has been in the habit, during the last forty years, of catching our migratory birds on their first arrival, writes to me that he has260 never known the females of any species to arrive before their males. During one spring he shot thirty-nine males of Ray’s wagtail (Budytes Raii) before he saw a single female. Mr. Gould has ascertained by dissection, as he informs me, that male snipes arrive in this country before the females. In the case of fish, at the period when the salmon ascend our rivers, the males in large numbers are ready to breed before the females. So it apparently is with frogs and toads. Throughout the great class of insects the males almost always emerge from the pupal state before the other sex, so that they generally swarm for a time before any females can be seen.338 The cause of this difference between the males and females in their periods of arrival and maturity is sufficiently obvious. Those males which annually first migrated into any country, or which in the spring were first ready to breed, or were the most eager, would leave the largest number of offspring; and these would tend to inherit similar instincts and constitutions. On the whole there can be no doubt that with almost all animals, in which the sexes are separate, there is a constantly recurrent struggle between the males for the possession of the females.
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 the beaten and less 261attractive males. Unless this result followed, the characters which gave 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 ultimately find females (excepting where polygamy prevails), and leave as many offspring, equally 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 were well developed, the males considerably exceeded the females in number; and this does hold good in some few cases. 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 proportions 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 during many years carefully attended to the habits of birds. There can also be no doubt that the most vigorous, healthy, and best-nourished females would on an average succeed in rearing the largest number of offspring. The males, as we have seen, are generally ready to breed before the females; of the males the262 strongest, and with some species the best armed, drive away the weaker males; and the former would then unite with the more vigorous and best-nourished females, as these are the first to breed. Such vigorous pairs would surely rear a larger number of offspring than the retarded females, which would be compelled, supposing the sexes to be numerically equal, to unite with the conquered and less powerful males; 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.
But in a multitude of cases the males which conquer other males, do not obtain possession of the females, independently of choice on the part 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, as has been actually observed in some cases, that they would at the same time prefer the more vigorous and lively males.339 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. Such early pairs would have the same advantage in rearing offspring on the female side as above explained, and nearly the same advantage on the male side. And this apparently has sufficed during a long course of generations to add not only to the strength and fighting-powers of 263the 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 in providing for the young. The same principles would apply if both sexes mutually preferred and selected certain individuals of the opposite sex; supposing that they selected not only the more attractive, but likewise the more vigorous individuals.
Numerical Proportion of the Two Sexes.—I have remarked that sexual selection would be a simple affair if the males considerably exceeded in number the females. Hence I was led to investigate, as far as I could, the proportions between the two sexes of as many animals as possible; but the materials are scanty. I will here give only a brief abstract of the results, retaining the details for a supplementary discussion, so as not to interfere with the course of my argument. Domesticated animals alone afford the opportunity of ascertaining the proportional numbers at birth; but no records have been specially kept for this purpose. By indirect means, however, I have collected a considerable body of statistical data, from which it appears that with most of our domestic animals the sexes are nearly equal at birth. Thus with race-horses, 25,560 births have been recorded during twenty-one years, and the male births have been to the female births as 99·7 to 100. With greyhounds the264 inequality is greater than with any other animal, for during twelve years, out of 6878 births, the male births have been as 110·1 to 100 female births. It is, however, in some degree doubtful whether it is safe to infer that the same proportional numbers would hold good under natural conditions as under domestication; for slight and unknown differences in the conditions affect to a certain extent the proportion of the sexes. Thus with mankind, the male births in England are as 104·5, in Russia as 108·9, and with the Jews of Livornia as 120 to 100 females. The proportion is also mysteriously affected by the circumstance of the births being legitimate or illegitimate.
For our present purpose we are concerned with the propor
tion of the sexes, not at birth, but at maturity, and this adds another element of doubt; for it is a well ascertained fact that with man a considerably larger proportion of males than of females die before or during birth, and during the first few years of infancy. So it almost certainly is with male lambs, and so it may be with the males of other animals. The males of some animals kill each other by fighting; or they drive each other about until they become greatly emaciated. They must, also, whilst wandering about in eager search for the females, be often exposed to various dangers. With many kinds of fish the males are much smaller than the females, and they are believed often to be devoured by the latter or by other fishes. With some birds the females appear to die in larger proportion than the males: they are also liable to be destroyed on their nests, or whilst in charge of their young. With insects the female larvæ are often larger than those of the males, and would consequently be more likely to be devoured: in some cases the mature females are less active and less rapid in their movements than the males,265 and would not be so well able to escape from danger. Hence, with animals in a state of nature, in order to judge of the proportions of the sexes at maturity, we must rely on mere estimation; and this, except perhaps when the inequality is strongly marked, is but little trustworthy. Nevertheless, as far as a judgment can be formed, we may conclude from the facts given in the supplement, that the males of some few mammals, of many birds, of some fish and insects, considerably exceed in number the females.
The proportion between the sexes fluctuates slightly during successive years: thus with race-horses, for every 100 females born, the males varied from 1O7.1 in one year to 92.6 in another year, and with greyhounds from 116.3 to 95.3. But had larger numbers been tabulated throughout a more extensive area than England, these fluctuations would probably have disappeared; and such as they are, they would hardly suffice to lead under a state of nature to the effective action of sexual selection. Nevertheless with some few wild animals, the proportions seem, as shewn in the supplement, to fluctuate either during different seasons or in different localities in a sufficient degree to lead to such action. For it should be observed that any advantage gained during certain years or in certain localities by those males which were able to conquer other males, or were the most attractive to the females, would probably be transmitted to the offspring and would not subsequently be eliminated. During the succeeding seasons, when from the equality of the sexes every male was everywhere able to procure a female, the stronger or more attractive males previously produced would still have at least as good a chance of leaving offspring as the less strong or less attractive.
Polygamy.—The practice of polygamy leads to the266 same results as would follow from an actual inequality in the number of the sexes; for if each male secures two or more females, many males will not be able to pair; and the latter assuredly will be the weaker or less attractive individuals. Many mammals and some few birds are polygamous, but with animals belonging to the lower classes I have found no evidence of this habit. The intellectual powers of such animals are, perhaps, not sufficient to lead them to collect and guard a harem of females. That some relation exists between polygamy and the development of secondary sexual characters, appears nearly certain; and this supports the view that a numerical preponderance of males would be eminently favourable to the action of sexual selection. Nevertheless many animals, especially birds, which are strictly monogamous, display strongly-marked secondary sexual characters; whilst some few animals, which are polygamous, are not thus characterised.
We will first briefly run through the class of mammals, and then turn to birds. The gorilla seems to be a polygamist, and the male differs considerably from the female; so it is with some baboons which live in herds containing twice as many adult females as males. In South America the Mycetes caraya presents well-marked sexual differences in colour, beard, and vocal organs, and the male generally lives with two or three wives: the male of the Cebus capucinus differs somewhat from the female, and appears to be polygamous.340 Little is known on this head with respect to most other monkeys, but some species are strictly monogamous. The ruminants are eminently polygamous, and they 267more frequently present sexual differences than almost any other group of mammals, especially in their weapons, but likewise in other characters. Most deer, cattle, and sheep are polygamous; as are most antelopes, though some of the latter are monogamous. Sir Andrew Smith, in speaking of the antelopes of South Africa, says that in herds of about a dozen there was rarely more than one mature male. The Asiatic Antilope saiga appears to be the most inordinate polygamist in the world; for Pallas341 states that the male drives away all rivals, and collects a herd of about a hundred, consisting of females and kids: the female is hornless and has softer hair, but does not otherwise differ much from the male. The horse is polygamous, but, except in his greater size and in the proportions of his body, differs but little from the mare. The wild boar, in his great tusks and some other characters, presents well-marked sexual characters; in Europe and in India he leads a solitary life, except during the breeding-season; but at this season he consorts in India with several females, as Sir W. Elliot, who has had large experience in observing this animal, believes: whether this holds good in Europe is doubtful, but is supported by some statements. The adult male Indian elephant, like the boar, passes much of his time in solitude; but when associating with others, “it is rare to find,” as Dr. Campbell states, “more than one male with a whole herd of females.” The larger males expel or kill the smaller and weaker ones. The male differs from the female by his immense tusks and greater size, strength, and endurance; so great is the difference in these latter 268respects, that the males when caught are valued at twenty per cent. above the females.342 With other pachydermatous animals the sexes differ very little or not at all, and they are not, as far as known, polygamists. Hardly a single species amongst the Cheiroptera and Edentata, or in the great Orders of the Rodents and Insectivora, presents well-developed secondary sexual differences; and I can find no account of any species being polygamous, excepting, perhaps, the common rat, the males of which, as some rat-catchers affirm, live with several females.
The lion in South Africa, as I hear from Sir Andrew Smith, sometimes lives with a single female, but generally with more than one, and, in one case, was found with as many as five females, so that he is polygamous. He is, as far as I can discover, the sole polygamist in the whole group of the terrestrial Carnivora, and he alone presents well-marked sexual characters. If, however, we turn to the marine Carnivora, the case is widely different; for many species of seals offer, as we shall hereafter see, extraordinary sexual differences, and they are eminently polygamous. Thus the male sea-elephant of the Southern Ocean, always possesses, according to Péron, several females, and the sea-lion of Forster is said to be surrounded by from twenty to thirty females. In the North, the male sea-bear of Steller is accompanied by even a greater number of females.
With respect to birds, many species, the sexes of which differ greatly from each other, are certainly monogamous. In Great Britain we see well-marked sexual differences in, for instance, the wild-duck which pairs with a single female, with the common blackbird, 269and with the bullfinch which is said to pair for life. So it is, as I am informed by Mr. Wallace, with the Chatterers or Cotingidæ of South America, and numerous other birds. In several groups I have not been able to discover whether the species are polygamous or monogamous. Lesson says that birds of paradise, so remarkable for their sexual differences, are polygamous, but Mr. Wallace doubts whether he had sufficient evidence. Mr. Salvin informs me that he has been led to believe that humming-birds are polygamous. The male widow-bird; remarkable for his caudal plumes, certainly seems to be a polygamist.343 I have been assured by Mr. Jenner Weir and by others, that three starlings not rarely frequent the same nest; but whether this is a case of polygamy or polyandry has not been ascertained.
The Gallinaceæ present almost as strongly marked sexual differences as birds of paradise or humming-bir
ds, and many of the species are, as is well known, polygamous; others being strictly monogamous. What a contrast is presented between the sexes by the polygamous peacock or pheasant, and the monogamous guinea-fowl or partridge! Many similar cases could be given, as in the grouse tribe, in which the males of the polygamous capercailzie and black-cock differ greatly from the females; whilst the sexes of the monogamous red grouse and ptarmigan differ very little. Amongst the Cursores, no great number of species offer strongly-marked sexual differences, except the bustards, and the great bustard (Otis tarda), is said to 270be polygamous. With the Grallatores, extremely few species differ sexually, but the ruff (Machetes pugnax) affords a strong exception, and this species is believed by Montagu to be a polygamist. Hence it appears that with birds there often exists a close relation between polygamy and the development of strongly-marked sexual differences. On asking Mr. Bartlett, at the Zoological Gardens, who has had such large experience with birds, whether the male tragopan (one of the Gallinaceæ) was polygamous, I was struck by his answering, “I do not know, but should think so from his splendid colours.”