Secondary sexual characters--Sexual selection--Manner of action--Excess of males-Polygamy--The male alone generally modified through sexual selection--Eagerness of the male--Variability of the male--Choice exerted by the female--Sexual compared with natural selection--Inheritance, at corresponding periods of life, at corresponding seasons of the year, and as limited by sex--Relations between the several forms of inheritance-Causes why one sex and the young are not modified through sexual selection-- Supplement on the proportional numbers of the two sexes throughout the animal kingdom--The proportion of the sexes in relation to natural selection.
With animals which have their sexes separated, the males necessarily differ from the females in their organs of reproduction; and these are the primary sexual characters. But the sexes often differ in what Hunter has called secondary sexual characters, which are not directly connected with the act of reproduction; for instance, the male possesses certain organs of sense or locomotion, of which the female is quite destitute, or has them more highly-developed, in order that he may readily find or reach her; or again the male has special organs of prehension for holding her securely. These latter organs, of infinitely diversified kinds, graduate into those which are commonly ranked as primary, and in some cases can hardly be distinguished from them; we see instances of this in the complex appendages at the apex of the abdomen in male insects. Unless indeed we confine the term "primary" to the reproductive glands, it is scarcely possible to decide which ought to be called primary and which secondary.
The female often differs from the male in having organs for the nourishment or protection of her young, such as the mammary glands of mammals, and the abdominal sacks of the marsupials. In some few cases also the male possesses similar organs, which are wanting in the female, such as the receptacles for the ova in certain male fishes, and those temporarily developed in certain male frogs. The females of most bees are provided with a special apparatus for collecting and carrying pollen, and their ovipositor is modified into a sting for the defence of the larvae and the community. Many similar cases could be given, but they do not here concern us. There are, however, other sexual differences quite unconnected with the primary reproductive organs, and it is with these that we are more especially concerned--such as the greater size, strength, and pugnacity of the male, his weapons of offence or means of defence against rivals, his gaudy colouring and various ornaments, his power of song, and other such characters.
Besides the primary and secondary sexual differences, such as the foregoing, the males and females of some animals differ in structures related to different habits of life, and not at all, or only indirectly, to the reproductive functions. Thus the females of certain flies (Culicidae and Tabanidae) are blood-suckers, whilst the males, living on flowers, have mouths destitute of mandibles. (1. Westwood, 'Modern Classification of Insects,' vol. ii. 1840, p. 541. For the statement about Tanais, mentioned below, I am indebted to Fritz Muller.) The males of certain moths and of some crustaceans (e.g. Tanais) have imperfect, closed mouths, and cannot feed. The complemental males of certain Cirripedes live like epiphytic plants either on the female or the hermaphrodite form, and are destitute of a mouth and of prehensile limbs. In these cases it is the male which has been modified, and has lost certain important organs, which the females possess. In other cases it is the female which has lost such parts; for instance, the female glow-worm is destitute of wings, as also are many female moths, some of which never leave their cocoons. Many female parasitic crustaceans have lost their natatory legs. In some weevil- beetles (Curculionidae) there is a great difference between the male and female in the length of the rostrum or snout (2. Kirby and Spence, 'Introduction to Entomology,' vol. iii. 1826, p. 309.); but the meaning of this and of many analogous differences, is not at all understood. Differences of structure between the two sexes in relation to different habits of life are generally confined to the lower animals; but with some few birds the beak of the male differs from that of the female. In the Huia of New Zealand the difference is wonderfully great, and we hear from Dr. Buller (3. 'Birds of New Zealand,' 1872, p. 66.) that the male uses his strong beak in chiselling the larvae of insects out of decayed wood, whilst the female probes the softer parts with her far longer, much curved and pliant beak: and thus they mutually aid each other. In most cases, differences of structure between the sexes are more or less directly connected with the propagation of the species: thus a female, which has to nourish a multitude of ova, requires more food than the male, and consequently requires special means for procuring it. A male animal, which lives for a very short time, might lose its organs for procuring food through disuse, without detriment; but he would retain his locomotive organs in a perfect state, so that he might reach the female. The female, on the other hand, might safely lose her organs for flying, swimming, or walking, if she gradually acquired habits which rendered such powers useless.
We are, however, here concerned only with sexual selection. This depends on the advantage which certain individuals have over others of the same sex and species solely in respect of reproduction. When, as in the cases above mentioned, the two sexes differ in structure in relation to different habits of life, they have no doubt been modified through natural selection, and by inheritance limited to one and the same sex. So again the primary sexual organs, and those for nourishing or protecting the young, come under the same influence; for those individuals which generated or nourished their offspring best, would leave, ceteris paribus, the greatest number to inherit their superiority; whilst those which generated or nourished their offspring badly, would leave but few to inherit their weaker powers. As the male has to find the female, he requires organs of sense and locomotion, but if these organs are necessary for the other purposes of life, as is generally the case, they will have been developed through natural selection. When the male has found the female, he sometimes absolutely requires prehensile organs to hold her; thus Dr. Wallace informs me that the males of certain moths cannot unite with the females if their tarsi or feet are broken. The males of many oceanic crustaceans, when adult, have their legs and antennae modified in an extraordinary manner for the prehension of the female; hence we may suspect that it is because these animals are washed about by the waves of the open sea, that they require these organs in order to propagate their kind, and if so, their development has been the result of ordinary or natural selection. Some animals extremely low in the scale have been modified for this same purpose; thus the males of certain parasitic worms, when fully grown, have the lower surface of the terminal part of their bodies roughened like a rasp, and with this they coil round and permanently hold the females. (4. M. Perrier advances this case ('Revue Scientifique,' Feb. 1, 1873, p. 865) as one fatal to the belief in sexual election, inasmuch as he supposes that I attribute all the differences between the sexes to sexual selection. This distinguished naturalist, therefore, like so many other Frenchmen, has not taken the trouble to understand even the first principles of sexual selection. An English naturalist insists that the claspers of certain male animals could not have been developed through the choice of the female! Had I not met with this remark, I should not have thought it possible for any one to have read this chapter and to have imagined that I maintain that the choice of the female had anything to do with the development of the prehensile organs in the male.)
When the two sexes follow exactly the same habits of life, and the male has the sensory or locomotive organs more highly developed than those of the female, it may be that the perfection of these is indispensable to the male for finding the female; but in the vast majority of cases, they serve only to give one male an advantage over another, for with sufficient time, the less well-endowed males would succeed in pairing with the females; and judging from the structure of the female, they would be in all other respects equally well adapted for their ordinary habits of life. Since in such cases the males have acquired their present structure, not from being better fitted to survive in the struggle for existence, but from having gained an advantage over other males, and from having transmitted this advantage to their male offspring alone, sexual selection must here have come into action. It was the importance of this distinction which led me to designate this form of selection as Sexual Selection. So again, if the chief service rendered to the male by his prehensile organs is to prevent the escape of the female before the arrival of other males, or when assaulted by them, these organs will have been perfected through sexual selection, that is by the advantage acquired by certain individuals over their rivals. But in most cases of this kind it is impossible to distinguish between the effects of natural and sexual selection. Whole chapters could be filled with details on the differences between the sexes in their sensory, locomotive, and prehensile organs. As, however, these structures are not more interesting than others adapted for the ordinary purposes of life I shall pass them over almost entirely, giving only a few instances under each class.
There are many other structures and instincts which must have been developed through sexual selection--such as the weapons of offence and the means of defence of the males for fighting with and driving away their rivals--their courage and pugnacity--their various ornaments--their contrivances for producing vocal or instrumental music--and their glands for emitting odours, most of these latter structures serving only to allure or excite the female. It is clear that these characters are the result of sexual and not of ordinary selection, since unarmed, unornamented, or unattractive males would succeed equally well in the battle for life and in leaving a numerous progeny, but for the presence of better endowed males. We may infer that this would be the case, because the females, which are unarmed and unornamented, are able to survive and procreate their kind. Secondary sexual characters of the kind just referred to, will be fully discussed in the following chapters, as being in many respects interesting, but especially as depending on the will, choice, and rivalry of the individuals of either sex. When we behold two males fighting for the possession of the female, or several male birds displaying their gorgeous plumage, and performing strange antics before an assembled body of females, we cannot doubt that, though led by instinct, they know what they are about, and consciously exert their mental and bodily powers.
Just as man can improve the breeds of his game-cocks by the selection of those birds which are victorious in the cockpit, so it appears that the strongest and most vigorous males, or those provided with the best weapons, have prevailed under nature, and have led to the improvement of the natural breed or species. A slight degree of variability leading to some advantage, however slight, in reiterated deadly contests would suffice for the work of sexual selection; and it is certain that secondary sexual characters are eminently variable. Just as man can give beauty, according to his standard of taste, to his male poultry, or more strictly can modify the beauty originally acquired by the parent species, can give to the Sebright bantam a new and elegant plumage, an erect and peculiar carriage-- so it appears that female birds in a state of nature, have by a long selection of the more attractive males, added to their beauty or other attractive qualities. No doubt this implies powers of discrimination and taste on the part of the female which will at first appear extremely improbable; but by the facts to be adduced hereafter, I hope to be able to shew that the females actually have these powers. When, however, it is said that the lower animals have a sense of beauty, it must not be supposed that such sense is comparable with that of a cultivated man, with his multiform and complex associated ideas. A more just comparison would be between the taste for the beautiful in animals, and that in the lowest savages, who admire and deck themselves with any brilliant, glittering, or curious object.
From our ignorance on several points, the precise manner in which sexual selection acts is somewhat 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 amongst 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 have the opportunity of selecting one out of several males, on the supposition that their mental capacity suffices for the exertion of a choice. In many cases special circumstances tend to make the struggle between the males particularly severe. Thus the males of our migratory birds generally arrive at their places of breeding before the females, so that many males are ready to contend for each female. I am informed by Mr. Jenner Weir, that the bird-catchers assert that this is invariably the case with the nightingale and blackcap, and with respect to the latter he can himself confirm the statement.
Mr. Swaysland of Brighton has been in the habit, during the last forty years, of catching our migratory birds on their first arrival, and he has 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 the dissection of those snipes which arrive the first in this country, that the males come before the females. And the like holds good with most of the migratory birds of the United States. (5. J.A. Allen, on the 'Mammals and Winter Birds of Florida,' Bulletin of Comparative Zoology, Harvard College, p. 268.) The majority of the male salmon in our rivers, on coming up from the sea, are ready to breed before the females. So it appears to be with frogs and toads. Throughout the great class of insects the males almost always are the first to emerge from the pupal state, so that they generally abound for a time before any females can be seen. (6. Even with those plants in which the sexes are separate, the male flowers are generally mature before the female. As first shewn by C.K. Sprengel, many hermaphrodite plants are dichogamous; that is, their male and female organs are not ready at the same time, so that they cannot be self-fertilised. Now in such flowers, the pollen is in general matured before the stigma, though there are exceptional cases in which the female organs are beforehand.) 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. It must be borne in mind that it would have been impossible to change very materially the time of sexual maturity in the females, without at the same time interfering with the period of the production of the young--a period which must be determined by the seasons of the year. 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 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. (7. Here is excellent evidence on the character of the offspring from an experienced ornithologist. Mr. J.A. Allen, in speaking ('Mammals and Winter Birds of E. Florida,' p. 229) of the later broods, after the accidental destruction of the first, says, that these "are found to be smaller and paler-coloured than those hatched earlier in the season. In cases where several broods are reared each year, as a general rule the birds of the earlier broods seem in all respects the most perfect and vigorous.") 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. (8. Hermann Muller has come to this same conclusion with respect to those female bees which are the first to emerge from the pupa each year. See his remarkable essay, 'Anwendung der Darwin'schen Lehre auf Bienen,' 'Verh. d. V. Jahrg.' xxix. p. 45.) 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.
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. (9. With respect to poultry, I have received information, hereafter to be given, to this effect. Even birds, such as pigeons, which pair for life, the female, as I hear from Mr. Jenner Weir, will desert her mate if he is injured or grows weak.) 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.
NUMERICAL PROPORTION OF THE TWO SEXES.I have remarked that sexual selection would be a simple affair if the males were considerably more numerous than 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 means 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 statistics, from which it appears that with most of our domestic animals the sexes are nearly equal at birth. Thus 25,560 births of race- horses have been recorded during twenty-one years, and the male births were to the female births as 99.7 to 100. In greyhounds the inequality is greater than with any other animal, for out of 6878 births during twelve years, the male births were to the female as 110.1 to 100. It is, however, in some degree doubtful whether it is safe to infer that the proportion would be the same under natural conditions as under domestication; for slight and unknown differences in the conditions affect 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 Livonia as 120, to 100 female births. But I shall recur to this curious point of the excess of male births in the supplement to this chapter. At the Cape of Good Hope, however, male children of European extraction have been born during several years in the proportion of between 90 and 99 to 100 female children.
For our present purpose we are concerned with the proportions of the sexes, not only at birth, but also at maturity, and this adds another element of doubt; for it is a wellascertained fact that with man the number of males dying before or during birth, and during the first two years of infancy, is considerably larger than that of females. So it almost certainly is with male lambs, and probably with some other animals. The males of some species kill one another by fighting; or they drive one another about until they become greatly emaciated. They must also be often exposed to various dangers, whilst wandering about in eager search for the females. In 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. The females of some birds appear to die earlier than the males; they are also liable to be destroyed on their nests, or whilst in charge of their young. With insects the female larvae 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, and could not escape so well from danger. Hence, with animals in a state of nature, we must rely on mere estimation, in order to judge of the proportions of the sexes at maturity; and this is but little trustworthy, except when the inequality is strongly marked. 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, are considerably more numerous than the females.
The proportion between the sexes fluctuates slightly during successive years: thus with race-horses, for every 100 mares born the stallions varied from 107.1 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 an area more extensive than England, these fluctuations would probably have disappeared; and such as they are, would hardly suffice to lead to effective sexual selection in a state of nature. Nevertheless, in the cases of some few wild animals, as shewn in the supplement, the proportions seem to fluctuate either during different seasons or in different localities in a sufficient degree to lead to such selection. For it should be observed that any advantage, gained during certain years or in certain localities by those males which were able to conquer their rivals, 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 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 weaker or less attractive.
POLYGAMY.The practice of polygamy leads to the 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 cannot 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, which are strictly monogamous, especially birds, display strongly-marked secondary sexual characters; whilst some few animals, which are polygamous, do not have such characters.
We will first briefly run through the mammals, and then turn to birds. The gorilla seems to be polygamous, 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. (10. On the Gorilla, Savage and Wyman, 'Boston Journal of Natural History,' vol. v. 1845-47, p. 423. On Cynocephalus, Brehm, 'Thierleben,' B. i. 1864, s. 77. On Mycetes, Rengger, 'Naturgeschichte der Saugethiere von Paraguay,' 1830, ss. 14, 20. On Cebus, Brehm, ibid. s. 108.) 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 present sexual differences more frequently than almost any other group of mammals; this holds good, especially in their weapons, but also in other characters. Most deer, cattle, and sheep are polygamous; as are most antelopes, though some 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 Pallas (11. Pallas, 'Spicilegia Zoolog., Fasc.' xii. 1777, p. 29. Sir Andrew Smith, 'Illustrations of the Zoology of S. Africa,' 1849, pl. 29, on the Kobus. Owen, in his 'Anatomy of Vertebrates' (vol. iii. 1868, p. 633) gives a table shewing incidentally which species of antelopes are gregarious.) states that the male drives away all rivals, and collects a herd of about a hundred females and kids together; the female is hornless and has softer hair, but does not otherwise differ much from the male. The wild horse of the Falkland Islands and of the Western States of N. America is polygamous, but, except in his greater size and in the proportions of his body, differs but little from the mare. The wild boar presents well-marked sexual characters, in his great tusks and some other points. In Europe and in India he leads a solitary life, except during the breeding-season; but as is believed by Sir W. Elliot, who has had many opportunities in India of observing this animal, he consorts at this season with several females. Whether this holds good in Europe is doubtful, but it is supported by some evidence. The adult male Indian elephant, like the boar, passes much of his time in solitude; but as Dr. Campbell states, when with others, "It is rare to find more than one male with a whole herd of females"; the larger males expelling or killing the smaller and weaker ones. The male differs from the female in his immense tusks, greater size, strength, and endurance; so great is the difference in these respects that the males when caught are valued at one-fifth more than the females. (12. Dr. Campbell, in 'Proc. Zoolog. Soc.' 1869, p. 138. See also an interesting paper by Lieut. Johnstone, in 'Proceedings, Asiatic Society of Bengal,' May 1868.) The sexes of other pachydermatous animals differ very little or not at all, and, as far as known, they are not polygamists. Nor have I heard of any species in the Orders of Cheiroptera, Edentata, Insectivora and Rodents being polygamous, excepting that amongst the Rodents, the common rat, according to some ratcatchers, lives with several females. Nevertheless the two sexes of some sloths (Edentata) differ in the character and colour of certain patches of hair on their shoulders. (13. Dr. Gray, in 'Annals and Magazine of Natural History,' 1871, p. 302.) And many kinds of bats (Cheiroptera) present well-marked sexual differences, chiefly in the males possessing odoriferous glands and pouches, and by their being of a lighter colour. (14. See Dr. Dobson's excellent paper in 'Proceedings of the Zoological Society,' 1873, p. 241.) In the great order of Rodents, as far as I can learn, the sexes rarely differ, and when they do so, it is but slightly in the tint of the fur.
As I hear from Sir Andrew Smith, the lion in South Africa sometimes lives with a single female, but generally with more, and, in one case, was found with as many as five females; so that he is polygamous. As far as I can discover, he is the only polygamist amongst all the terrestrial Carnivora, and he alone presents well-marked sexual characters. If