With animals which have their sexes separated, the males necessarily differ from the females in their organs of reproduction; and these afford 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, in the male possessing certain organs of sense or locomotion, of which the female is quite destitute, or in having them more highly-developed, in order that he may readily find or reach her; or again, in the male having special organs of prehension so as to hold her securely. These latter organs of infinitely diversified kinds graduate into, and in some cases can hardly be distinguished from, those which are commonly ranked as primary, such as 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, as far as the organs of prehension are concerned, 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, as the mammary glands of mammals, and the abdominal sacks of the marsupials. The male, also, in some few cases differs from the female in possessing analogous organs, as the receptacles for the ova possessed by the males of certain fishes, and those temporarily developed in certain male frogs. Female bees have a special apparatus for collecting and carrying pollen, and their ovipositor is modified into a sting for the defence of their larvæ and the community. In the females of many insects the ovipositor is modified in the most complex manner for the safe placing of the eggs. Numerous similar cases could be given, but they do not here concern us. There are, however, other sexual differences quite disconnected with the primary organs with which 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 foregoing primary and secondary sexual differences, the male and female sometimes differ in structures connected with different habits of life, and not at all, or only indirectly, related to the reproductive functions. Thus the females of certain flies (Culicidæ and Tabanidæ) are blood-suckers, whilst the males live on flowers and have their mouths destitute of mandibles.336 The males alone 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 hermaphrodite form, and are destitute of a mouth and prehensile limbs. In these cases it is the male which has been modified and has lost certain important organs, which the other members of the same group possess. In other cases it is the female which has lost such parts; for instance, the female glow-worm is destitute of wings, as are many female moths, some of which never leave their cocoons. Many female parasitic crustaceans have lost their natatory legs. In some weevil-beetles (Curculionidæ) there is a great difference between the male and female in the length of the rostrum or snout;337 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. No doubt in most, but apparently not in all these cases, the differences are indirectly connected with the propagation of the species: thus a female which has to nourish a multitude of ova will require more food than the male, and consequently will require special means for procuring it. A male animal which lived for a very short time might without detriment lose through disuse its organs for procuring food; 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 that kind of selection, which I have called sexual selection. This depends on the advantage which certain individuals have over other individuals of the same sex and species, in exclusive relation to reproduction. When the two sexes differ in structure in relation to different habits of life, as in the cases above mentioned, they have no doubt been modified through natural selection, accompanied 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 this same head; for those individuals which generated or nourished their offspring best, would leave, cæteris 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 search for the female, he requires for this purpose 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 have their legs and antennæ modified in an extraordinary manner for the prehension of the female; hence we may suspect that owing to these animals being washed about by the waves of the open sea, they absolutely require these organs in order to propagate their kind, and if so their development will have been the result of ordinary or natural selection.

When the two sexes follow exactly the same habits of life, and the male has more highly developed sense or locomotive organs than the female, it may be that these in their perfected state are 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 the less well-endowed males, if time were allowed them, would succeed in pairing with the females; and they would in all other respects, judging from the structure of the female, be equally well adapted for their ordinary habits of life. In such cases sexual selection must have come into action, for 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. 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 males over their rivals. But in most cases it is scarcely possible to distinguish between the effects of natural and sexual selection. Whole chapters could easily 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 almost pass them over, 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 possessed by the males for fighting with and driving away their rivals—their courage and pugnacity—their ornaments of many kinds—their organs 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. That these characters are the result of sexual and not of ordinary selection is clear, as unarmed, unornamented, or unattractive males would succeed equally well in the battle for life and in leaving a numerous progeny, if better endowed males were not present. We may infer that this would be the case, for 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 they are in many respects interesting, but more especially as they depend 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 the strangest 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.

In the same manner as man can improve the breed 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. Through repeated deadly contests, a slight degree of variability, if it led to some advantage, however slight, would suffice for the work of sexual selection; and it is certain that secondary sexual characters are eminently variable. In the same manner as man can give beauty, according to his standard of taste, to his male poultry—can give to the Sebright bantam a new and elegant plumage, an erect and peculiar carriage—so it appears that in a state of nature female birds, by having long selected the more attractive males, have added to their beauty. No doubt this implies powers of discrimination and taste on the part of the female which will at first appear extremely improbable; but I hope hereafter to shew that this is not the case.

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 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 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 attractive 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 the 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 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 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 the 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 proportion 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, 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 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 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 more 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 respects, 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, and 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-birds, 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 be 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.”

It deserves notice that the instinct of pairing with a single female is easily lost under domestication. The wild-duck is strictly monogamous, the domestic-duck highly polygamous. The Rev. W. D. Fox informs me that with some half-tamed wild-ducks, kept on a large pond in his neighbourhood, so many mallards were shot by the gamekeeper that only one was left for every seven or eight females; yet unusually large broods were reared. The guinea-fowl is strictly monogamous; but Mr. Fox finds that his birds succeed best when he keeps one cock to two or three hens.344 Canary-birds pair in a state of nature, but the breeders in England successfully put one male to four or five females; nevertheless the first female, as Mr. Fox has been assured, is alone treated as the wife, she and her young ones being fed by him; the others are treated as concubines. I have noticed these cases, as it renders it in some degree probable that monogamous species, in a state of nature, might readily become either temporarily or permanently polygamous.

With respect to reptiles and fishes, too little is known of their habits to enable us to speak of their marriage arrangements. The stickle-back Gasterosteus, however, is said to be a polygamist;345 and the male during the breeding-season differs conspicuously from the female.

To sum up on the means through which, as far as we can judge, sexual selection has led to the development of secondary sexual characters. It has been shewn that the largest number of vigorous offspring will be reared from the pairing of the strongest and best-armed males, which have conquered other males, with the most vigorous and best-nourished females, which are the first to breed in the spring. Such females, if they select the more attractive, and at the same time vigorous, males, will rear a larger number of offspring than the retarded females, which must pair with the less vigorous and less attractive males. So it will be if the more vigorous males select the more attractive and at the same time healthy and vigorous females; and this will especially hold good if the male defends the female, and aids in providing food for the young. The advantage thus gained by the more vigorous pairs in rearing a larger number of offspring has apparently sufficed to render sexual selection efficient. But a large preponderance in number of the males over the females would be still more efficient; whether the preponderance was only occasional and local, or permanent; whether it occurred at birth, or subsequently from the greater destruction of the females; or whether it indirectly followed from the practice of polygamy.

The Male generally more modified than the Female.—Throughout the animal kingdom, when the sexes differ from each other in external appearance, it is the male which, with rare exceptions, has been chiefly modified; for the female still remains more like the young of her own species, and more like the other members of the same group. The cause of this seems to lie in the males of almost all animals having stronger passions than the females. Hence it is the males that fight together and sedulously display their charms before the females; and those which are victorious transmit their superiority to their male offspring. Why the males do not transmit their characters to both sexes will hereafter be considered. That the males of all mammals eagerly pursue the females is notorious to every one. So it is with birds; but many male birds do not so much pursue the female, as display their plumage, perform strange antics, and pour forth their song, in her presence. With the few fish which have been observed, the male seems much more eager than the female; and so it is with alligators, and apparently with Batrachians. Throughout the enormous class of insects, as Kirby remarks,346 “the law is, that the male shall seek the female.” With spiders and crustaceans, as I hear from two great authorities, Mr. Blackwall and Mr. C. Spence Bate, the males are more active and more erratic in their habits than the females. With insects and crustaceans, when the organs of sense or locomotion are present in the one sex and absent in the other, or when, as is frequently the case, they are more highly developed in the one than the other, it is almost invariably the male, as far as I can discover, which retains such organs, or has them most developed; and this shews that the male is the more active member in the courtship of the sexes.347

The female, on the other hand, with the rarest exception, is less eager than the male. As the illustrious Hunter348 long ago observed, she generally “requires to be courted;” she is coy, and may often be seen endeavouring for a long time to escape from the male. Every one who has attended to the habits of animals will be able to call to mind instances of this kind. Judging from various facts, hereafter to be given, and from the results which may fairly be attributed to sexual selection, the female, though comparatively passive, generally exerts some choice and accepts one male in preference to others. Or she may accept, as appearances would sometimes lead us to believe, not the male which is the most attractive to her, but the one which is the least distasteful. The exertion of some choice on the part of the female seems almost as general a law as the eagerness of the male.

We are naturally led to enquire why the male in so many and such widely distinct classes has been rendered 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 both sexes were mutually to search for each other; but why should the male almost always be the seeker? With plants, the ovules after fertilisation have to be nourished for a time; hence the pollen is necessarily brought to the female organs—being placed on the stigma, through the agency of insects or of the wind, or by the spontaneous movements of the stamens; and with the Algæ, &c., by the locomotive power of the antherozooids. With lowly-organised animals permanently affixed to the same spot and having their sexes separate, the male element is invariably brought to the female; and we can see the reason; for the ova, even if detached before being fertilised and not requiring subsequent nourishment or protection, would be, from their larger relative size, less easily transported than the male element. Hence plants349 and many of the lower animals are, in this respect, analogous. In the case of animals not affixed to the same spot, but enclosed within a shell with no power of protruding any part of their bodies, and in the case of animals having little power of locomotion, the males must trust the fertilising element to the risk of at least a short transit through the waters of the sea. It would, therefore, be a great advantage to such animals, as their organisation became perfected, if the males when ready to emit the fertilising element, were to acquire the habit of approaching the female as closely as possible. The males of various lowly-organised animals having thus aboriginally acquired the habit of approaching and seeking the females, the same habit would naturally be transmitted to their more highly developed male descendants; and in order that they should become efficient seekers, they would have to be endowed with strong passions. The acquirement of such passions would naturally follow from the more eager males leaving a larger number of offspring than the less eager.

The great eagerness of the male has thus indirectly led to the much more frequent development of secondary sexual characters in the male than in the female. But the development of such characters will have been much aided, if the conclusion at which I arrived after studying domesticated animals, can be trusted, namely, that the male is more liable to vary than the female. I am aware how difficult it is to verify a conclusion of this kind. Some slight evidence, however, can be gained by comparing the two sexes in mankind, as man has been more carefully observed than any other animal. During the Novara Expedition350 a vast number of measurements of various parts of the body in different races were made, and the men were found in almost every case to present a greater range of variation than the women; but I shall have to recur to this subject in a future chapter. Mr. J. Wood,351 who has carefully attended to the variation of the muscles in man, puts in italics the conclusion that “the greatest number of abnormalities in each subject is found in the males.” He had previously remarked that “altogether in 102 subjects the varieties of redundancy were found to be half as many again as in females, contrasting widely with the greater frequency of deficiency in females before described.” Professor Macalister like wise remarks352 that variations in the muscles “are probably more common in males than females.” Certain muscles which are not normally present in mankind are also more frequently developed in the male than in the female sex, although exceptions to this rule are said to occur. Dr. Burt Wilder353 has tabulated the cases of 152 individuals with supernumerary digits, of which 86 were males, and 39, or less than half, females; the remaining 27 being of unknown sex. It should not, however, be overlooked that women would more frequently endeavour to conceal a deformity of this kind than men. Whether the large proportional number of deaths of the male offspring of man and apparently of sheep, compared with the female offspring, before, during, and shortly after birth (see supplement), has any relation to a stronger tendency in the organs of the male to vary and thus to become abnormal in structure or function, I will not pretend to conjecture.

In various classes of animals a few exceptional cases occur, in which the female instead of the male has acquired well pronounced secondary sexual characters, such as brighter colours, greater size, strength, or pugnacity. With birds, as we shall hereafter see, there has sometimes been a complete transposition of the ordinary characters proper to each sex; the females having become the more eager in courtship, the males remaining comparatively passive, but apparently selecting, as we may infer from the results, the more attractive females. Certain female birds have thus been rendered more highly coloured or otherwise ornamented, as well as more powerful and pugnacious than the males, these characters being transmitted to the female offspring alone.

It may be suggested that in some cases a double process of selection has been carried on; the males having selected the more attractive females, and the latter the more attractive males. This process however, though it might lead to the modification of both sexes, would not make the one sex different from the other, unless indeed their taste for the beautiful differed; but this is a supposition too improbable in the case of any animal, excepting man, to be worth considering. There are, however, many animals, in which the sexes resemble each other, both being furnished with the same ornaments, which analogy would lead us to attribute to the agency of sexual selection. In such cases it may be suggested with more plausibility, that there has been a double or mutual process of sexual selection; the more vigorous and precocious females having selected the more attractive and vigorous males, the latter having rejected all except the more attractive females. But from what we know of the habits of animals, this view is hardly probable, the male being generally eager to pair with any female. It is more probable that the ornaments common to both sexes were acquired by one sex, generally the male, and then transmitted to the offspring of both sexes. If, indeed, during a lengthened period the males of any species were greatly to exceed the females in number, and then during another lengthened period under different conditions the reverse were to occur, a double, but not simultaneous, process of sexual selection might easily be carried on, by which the two sexes might be rendered widely different.

We shall hereafter see that many animals exist, of which neither sex is brilliantly coloured or provided with special ornaments, and yet the members of both sexes or of one alone have probably been modified through sexual selection. The absence of bright tints or other ornaments may be the result of variations of the right kind never having occurred, or of the animals themselves preferring simple colours, such as plain black or white. Obscure colours have often been acquired through natural selection for the sake of protection, and the acquirement through sexual selection of conspicuous colours, may have been checked from the danger thus incurred. But in other cases the males have probably struggled together during long ages, through brute force, or by the display of their charms, or by both means combined, and yet no effect will have been produced unless a larger number of offspring were left by the more successful males to inherit their superiority, than by the less successful males; and this, as previously shewn, depends on various complex contingencies.

Sexual selection acts in a less rigorous manner than natural selection. The latter produces its effects by the life or death at all ages of the more or less successful individuals. Death, indeed, not rarely ensues from the conflicts of rival males. But generally the less successful male merely fails to obtain a female, or obtains later in the season a retarded and less vigorous female, or, if polygamous, obtains fewer females; so that they leave fewer, or less vigorous, or no offspring. In regard to structures acquired through ordinary or natural selection, there is in most cases, as long as the conditions of life remain the same, a limit to the amount of advantageous modification in relation to certain special ends; but in regard to structures adapted to make one male victorious over another, either in fighting or in charming the female, there is no definite limit to the amount of advantageous modification; so that as long as the proper variations arise the work of sexual selection will go on. This circumstance may partly account for the frequent and extraordinary amount of variability presented by secondary sexual characters. Nevertheless, natural selection will determine that characters of this kind shall not be acquired by the victorious males, which would be injurious to them in any high degree, either by expending too much of their vital powers, or by exposing them to any great danger. The development, however, of certain structures—of the horns, for instance, in certain stags—has been carried to a wonderful extreme; and in some instances to an extreme which, as far as the general conditions of life are concerned, must be slightly injurious to the male. From this fact we learn that the advantages which favoured males have derived from conquering other males in battle or courtship, and thus leaving a numerous progeny, have been in the long run greater than those derived from rather more perfect adaptation to the external conditions of life. We shall further see, and this could never have been anticipated, that the power to charm the female has been in some few instances more important than the power to conquer other males in battle.

LAWS OF INHERITANCE.

In order to understand how sexual selection has acted, and in the course of ages has produced conspicuous results with many animals of many classes, it is necessary to bear in mind the laws of inheritance, as far as they are known. Two distinct elements are included under the term “inheritance,” namely the transmission and the development of characters; but as these generally go together, the distinction is often overlooked. We see this distinction in those characters which are transmitted through the early years of life, but are developed only at maturity or during old age. We see the same distinction more clearly with secondary sexual characters, for these are transmitted through both sexes, though developed in one alone. That they are present in both sexes, is manifest when two species, having strongly-marked sexual characters, are crossed, for each transmits the characters proper to its own male and female sex to the hybrid offspring of both sexes. The same fact is likewise manifest, when characters proper to the male are occasionally developed in the female when she grows old or becomes diseased; and so conversely with the male. Again, characters occasionally appear, as if transferred from the male to the female, as when, in certain breeds of the fowl, spurs regularly appear in the young and healthy females; but in truth they are simply developed in the female; for in every breed each detail in the structure of the spur is transmitted through the female to her male offspring. In all cases of reversion, characters are transmitted through two, three, or many generations, and are then under certain unknown favourable conditions developed. This important distinction between transmission and development will be easiest kept in mind by the aid of the hypothesis of pangenesis, whether or not it be accepted as true. According to this hypothesis, every unit or cell of the body throws off gemmules or undeveloped atoms, which are transmitted to the offspring of both sexes, and are multiplied by self-division. They may remain undeveloped during the early years of life or during successive generations; their development into units or cells, like those from which they were derived, depending on their affinity for, and union with, other units or cells previously developed in the due order of growth.

Inheritance at Corresponding Periods of Life.—This tendency is well established. If a new character appears in an animal whilst young, whether it endures throughout life or lasts only for a time, it will reappear, as a general rule, at the same age and in the same manner in the offspring. If, on the other hand, a new character appears at maturity, or even during old age, it tends to reappear in the offspring at the same advanced age. When deviations from this rule occur, the transmitted characters much oftener appear before than after the corresponding age. As I have discussed this subject at sufficient length in another work,354 I will here merely give two or three instances, for the sake of recalling the subject to the reader’s mind. In several breeds of the Fowl, the chickens whilst covered with down, in their first true plumage, and in their adult plumage, differ greatly from each other, as well as from their common parent-form, the Gallus bankiva; and these characters are faithfully transmitted by each breed to their offspring at the corresponding period of life. For instance, the chickens of spangled Hamburghs, whilst covered with down, have a few dark spots on the head and rump, but are not longitudinally striped, as in many other breeds; in their first true plumage, “they are beautifully pencilled,” that is each feather is transversely marked by numerous dark bars; but in their second plumage the feathers all become spangled or tipped with a dark round spot.355 Hence in this breed variations have occurred and have been transmitted at three distinct periods of life. The Pigeon offers a more remarkable case, because the aboriginal parent-species does not undergo with advancing age any change of plumage, excepting that at maturity the breast becomes more iridescent; yet there are breeds which do not acquire their characteristic colours until they have moulted two, three, or four times; and these modifications of plumage are regularly transmitted.

Inheritance at Corresponding Seasons of the Year.—With animals in a state of nature innumerable instances occur of characters periodically appearing at different seasons. We see this with the horns of the stag, and with the fur of arctic animals which becomes thick and white during the winter. Numerous birds acquire bright colours and other decorations during the breeding-season alone. I can throw but little light on this form of inheritance from facts observed under domestication. Pallas states,356 that in Siberia domestic cattle and horses periodically become lighter-coloured during the winter; and I have observed a similar marked change of colour in certain ponies in England. Although I do not know that this tendency to assume a differently coloured coat during different seasons of the year is transmitted, yet it probably is so, as all shades of colour are strongly inherited by the horse. Nor is this form of inheritance, as limited by season, more remarkable than inheritance as limited by age or sex.

Inheritance as Limited by Sex.—The equal transmission of characters to both, sexes is the commonest form of inheritance, at least with those animals which do not present strongly-marked sexual differences, and indeed with many of these. But characters are not rarely transferred exclusively to that sex, in which they first appeared. Ample evidence on this head has been advanced in my work on Variation under Domestication; but a few instances may here be given. There are breeds of the sheep and goat, in which the horns of the male differ greatly in shape from those of the female; and these differences, acquired under domestication, are regularly transmitted to the same sex. With tortoise-shell cats the females alone, as a general rule, are thus coloured, the males being rusty-red. With most breeds of the fowl, the characters proper to each sex are transmitted to the same sex alone. So general is this form of transmission that it is an anomaly when we see in certain breeds variations transmitted equally to both sexes. There are also certain sub-breeds of the fowl in which the males can hardly be distinguished from each other, whilst the females differ considerably in colour. With the pigeon the sexes of the parent-species do not differ in any external character; nevertheless in certain domesticated breeds the male is differently coloured from the female.357 The wattle in the English Carrier pigeon and the crop in the Pouter are more highly developed in the male than in the female; and although these characters have been gained through long-continued selection by man, the difference between the two sexes is wholly due to the form of inheritance which has prevailed; for it has arisen, not from, but rather in opposition to, the wishes of the breeder.

Most of our domestic races have been formed by the accumulation of many slight variations; and as some of the successive steps have been transmitted to one sex alone, and some to both sexes, we find in the different breeds of the same species all gradations between great sexual dissimilarity and complete similarity. Instances have already been given with the breeds of the fowl and pigeon; and under nature analogous cases are of frequent occurrence. With animals under domestication, but whether under nature I will not venture to say, one sex may lose characters proper to it, and may thus come to resemble to a certain extent the opposite sex; for instance, the males of some breeds of the fowl have lost their masculine plumes and hackles. On the other hand the differences between the sexes may be increased under domestication, as with merino sheep, in which the ewes have lost their horns. Again, characters proper to one sex may suddenly appear in the other sex; as with those sub-breeds of the fowl in which the hens whilst young acquire spurs; or, as in certain Polish sub-breeds, in which the females, as there is reason to believe, originally acquired a crest, and subsequently transferred it to the males. All these cases are intelligible on the hypothesis of pangenesis; for they depend on the gemmules of certain units of the body, although present in both sexes, becoming through the influence of domestication dormant in the one sex; or if naturally dormant, becoming developed.

There is one difficult question which it will be convenient to defer to a future chapter; namely, whether a character at first developed in both sexes, can be rendered through selection limited in its development to one sex alone. If, for instance, a breeder observed that some of his pigeons (in which species characters are usually transferred in an equal degree to both sexes) varied into pale blue; could he by long-continued selection make a breed, in which the males alone should be of this tint, whilst the females remained unchanged? I will here only say, that this, though perhaps not impossible, would be extremely difficult; for the natural result of breeding from the pale-blue males would be to change his whole stock, including both sexes, into this tint. If, however, variations of the desired tint appeared, which were from the first limited in their development to the male sex, there would not be the least difficulty in making a breed characterised by the two sexes being of a different colour, as indeed has been effected with a Belgian breed, in which the males alone are streaked with black. In a similar manner, if any variation appeared in a female pigeon, which was from the first sexually limited in its development, it would be easy to make a breed with the females alone thus characterised; but if the variation was not thus originally limited, the process would be extremely difficult, perhaps impossible.

On the Relation between the period of Development of a Character and its transmission to one sex or to both sexes.—Why certain characters should be inherited by both sexes, and other characters by one sex alone, namely by that sex in which the character first appeared, is in most cases quite unknown. We cannot even conjecture why with certain sub-breeds of the pigeon, black striæ, though transmitted through the female, should be developed in the male alone, whilst every other character is equally transferred to both sexes. Why, again, with cats, the tortoise-shell colour should, with rare exceptions, be developed in the female alone. The very same characters, such as deficient or supernumerary digits, colour-blindness, &c., may with mankind be inherited by the males alone of one family, and in another family by the females alone, though in both cases transmitted through the opposite as well as the same sex.358 Although we are thus ignorant, two rules often hold good, namely that variations which, first appear in either sex at a late period of life, tend to be developed in the same sex alone; whilst variations which first appear early in life in either sex tend to be developed in both sexes. I am, however, far from supposing that this is the sole determining cause. As I have not elsewhere discussed this subject, and as it has an important bearing on sexual selection, I must here enter into lengthy and somewhat intricate details.

It is in itself probable that any character appearing at an early age would tend to be inherited equally by both sexes, for the sexes do not differ much in constitution, before the power of reproduction is gained. On the other hand, after this power has been gained and the sexes have come to differ in constitution, the gemmules (if I may again use the language of pangenesis) which are cast off from each varying part in the one sex would be much more likely to possess the proper affinities for uniting with the tissues of the same sex, and thus becoming developed, than with those of the opposite sex.

I was first led to infer that a relation of this kind exists, from the fact that whenever and in whatever manner the adult male has come to differ from the adult female, he differs in the same manner from the young of both sexes. The generality of this fact is quite remarkable: it holds good with almost all mammals, birds, amphibians, and fishes; also with many crustaceans, spiders and some few insects, namely certain orthoptera and libellulæ. In all these cases the variations, through the accumulation of which the male acquired his proper masculine characters, must have occurred at a somewhat late period of life; otherwise the young males would have been similarly characterised; and conformably with our rule, they are transmitted to and developed in the adult males alone. When, on the other hand, the adult male closely resembles the young of both sexes (these, with rare exceptions, being alike), he generally resembles the adult female; and in most of these cases the variations through which the young and old acquired their present characters, probably occurred in conformity with our rule during youth. But there is here room for doubt, as characters are sometimes transferred to the offspring at an earlier age than that at which they first appeared in the parents, so that the parents may have varied when adult, and have transferred their characters to their offspring whilst young. There are, moreover, many animals, in which the two sexes closely resemble each other, and yet both differ from their young; and here the characters of the adults must have been acquired late in life; nevertheless, these characters in apparent contradiction to our rule, are transferred to both sexes. We must not, however, overlook the possibility or even probability of successive variations of the same nature sometimes occurring, under exposure to similar conditions, simultaneously in both sexes at a rather late period of life; and in this case the variations would be transferred to the offspring of both sexes at a corresponding late age; and there would be no real contradiction to our rule of the variations which occur late in life being transferred exclusively to the sex in which they first appeared. This latter rule seems to hold true more generally than the second rule, namely, that variations which occur in either sex early in life tend to be transferred to both sexes. As it was obviously impossible even to estimate in how large a number of cases throughout the animal kingdom these two propositions hold good, it occurred to me to investigate some striking or crucial instances, and to rely on the result.

An excellent case for investigation is afforded by the Deer Family. In all the species, excepting one, the horns are developed in the male alone, though certainly transmitted through the female, and capable of occasional abnormal development in her. In the reindeer, on the other hand, the female is provided with horns; so that in this species, the horns ought, according to our rule, to appear early in life, long before the two sexes had arrived at maturity and had come to differ much in constitution. In all the other species of deer the horns ought to appear later in life, leading to their development in that sex alone, in which they first appeared in the progenitor of the whole Family. Now in seven species, belonging to distinct sections of the family and inhabiting different regions, in which the stags alone bear horns, I find that the horns first appear at periods varying from nine months after birth in the roebuck to ten or twelve or even more months in the stags of the six other larger species.359 But with the reindeer the case is widely different, for as I hear from Prof. Nilsson, who kindly made special enquiries for me in Lapland, the horns appear in the young animals within four or five weeks after birth, and at the same time in both sexes. So that here we have a structure, developed at a most unusually early age in one species of the family, and common to both sexes in this one species.

In several kinds of antelopes the males alone are provided with horns, whilst in the greater number both sexes have horns. With respect to the period of development, Mr. Blyth informs me that there lived at one time in the Zoological Gardens a young koodoo (Ant. strepsiceros), in which species the males alone are horned, and the young of a closely-allied species, viz. the eland (Ant. oreas), in which both sexes are horned. Now in strict conformity with our rule, in the young male koodoo, although arrived at the age of ten months, the horns were remarkably small considering the size ultimately attained by them: whilst in the young male eland, although only three months old, the horns were already very much larger than in the koodoo. It is also worth notice that in the prong-horned antelope,360 in which species the horns, though present in both sexes, are almost rudimentary in the female, they do not appear until about five or six months after birth. With sheep, goats, and cattle, in which the horns are well developed in both sexes, though not quite equal in size, they can be felt, or even seen, at birth or soon afterwards.361 Our rule, however, fails in regard to some breeds of sheep, for instance merinos, in which the rams alone are horned; for I cannot find on enquiry,362 that the horns are developed later in life in this breed than in ordinary sheep in which both sexes are horned. But with domesticated sheep the presence or absence of horns is not a firmly fixed character; a certain proportion of the merino ewes bearing small horns, and some of the rams being hornless; whilst with ordinary sheep hornless ewes are occasionally produced.

In most of the species of the splendid family of the Pheasants, the males differ conspicuously from the females, and they acquire their ornaments at a rather late period of life. The eared pheasant (Crossoptilon auritum), however, offers a remarkable exception, for both sexes possess the fine caudal plumes, the large ear-tufts and the crimson velvet about the head; and I find on enquiry in the Zoological Gardens that all these characters, in accordance with our rule, appear very early in life. The adult male can, however, be distinguished from the adult female by one character, namely by the presence of spurs; and conformably with our rule, these do not begin to be developed, as I am assured by Mr. Bartlett, before the age of six months, and even at this age, can hardly be distinguished in the two sexes.363 The male and female Peacock differ conspicuously from each other in almost every part of their plumage, except in the elegant head-crest, which is common to both sexes; and this is developed very early in life, long before the other ornaments which are confined to the male. The wild-duck offers an analogous case, for the beautiful green speculum on the wings is common to both sexes, though duller and somewhat smaller in the female, and it is developed early in life, whilst the curled tail-feathers and other ornaments peculiar to the male are developed later.364 Between such extreme cases of close sexual resemblance and wide dissimilarity, as those of the Crossoptilon and peacock, many intermediate ones could be given, in which the characters follow in their order of development our two rules.

As most insects emerge from their pupal state in a mature condition, it is doubtful whether the period of development determines the transference of their characters to one or both sexes. But we do not know that the coloured scales, for instance, in two species of butterflies, in one of which the sexes differ in colour, whilst in the other they are alike, are developed at the same relative age in the cocoon. Nor do we know whether all the scales are simultaneously developed on the wings of the same species of butterfly, in which certain coloured marks are confined to one sex, whilst other marks are common to both sexes. A difference of this kind in the period of development is not so improbable as it may at first appear; for with the Orthoptera, which assume their adult state, not by a single metamorphosis, but by a succession of moults, the young males of some species at first resemble the females, and acquire their distinctive masculine characters only during a later moult. Strictly analogous cases occur during the successive moults of certain male crustaceans.

We have as yet only considered the transference of characters, relatively to their period of development, with species in a natural state; we will now turn to domesticated animals; first touching on monstrosities and diseases. The presence of supernumerary digits, and the absence of certain phalanges, must be determined at an early embryonic period—the tendency to profuse bleeding is at least congenital, as is probably colour-blindness—yet these peculiarities, and other similar ones, are often limited in their transmission to one sex; so that the rule that characters which are developed at an early period tend to be transmitted to both sexes, here wholly fails. But this rule, as before remarked, does not appear to be nearly so generally true as the converse proposition, namely, that characters which appear late in life in one sex are transmitted exclusively to the same sex. From the fact of the above abnormal peculiarities becoming attached to one sex. long before the sexual functions are active, we may infer that there must be a difference of some kind between the sexes at an extremely early age. With respect to sexually-limited diseases, we know too little of the period at which they originate, to draw any fair conclusion. Gout, however, seems to fall under our rule; for it is generally caused by intemperance after early youth, and is transmitted from the father to his sons in a much more marked manner than to his daughters.

In the various domestic breeds of sheep, goats, and cattle, the males differ from their respective females in the shape or development of their horns, forehead, mane, dewlap, tail, and hump on the shoulders; and these peculiarities, in accordance with our rule, are not fully developed until rather late in life. With dogs, the sexes do not differ, except that in certain breeds, especially in the Scotch deer-hound, the male is much larger and heavier than the female; and as we shall see in a future chapter, the male goes on increasing in size to an unusually late period of life, which will account, according to our rule, for his increased size being transmitted to his male offspring alone. On the other hand, the tortoise-shell colour of the hair, which is confined to female cats, is quite distinct at birth, and this case violates our rule. There is a breed of pigeons in which the males alone are streaked with black, and the streaks can be detected even in the nestlings; but they become more conspicuous at each successive moult, so that this case partly opposes and partly supports the rule. With the English Carrier and Pouter pigeon the full development of the wattle and the crop occurs rather late in life, and these characters, conformably with our rule, are transmitted in full perfection to the males alone. The following cases perhaps come within the class previously alluded to, in which the two sexes have varied in the same manner at a rather late period of life, and have consequently transferred their new characters to both sexes at a corresponding late period; and if so, such cases are not opposed to our rule. Thus there are sub-breeds of the pigeon, described by Neumeister,365 both sexes of which change colour after moulting twice or thrice, as does likewise the Almond Tumbler; nevertheless these changes, though occurring rather late in life, are common to both sexes. One variety of the Canary-bird, namely the London Prize, offers a nearly analogous case.

With the breeds of the Fowl the inheritance of various characters by one sex or by both sexes, seems generally determined by the period at which such characters are developed. Thus in all the many breeds in which the adult male differs greatly in colour from the female and from the adult male parent-species, he differs from the young male, so that the newly acquired characters must have appeared at a rather late period of life. On the other hand with most of the breeds in which the two sexes resemble each other, the young are coloured in nearly the same manner as their parents, and this renders it probable that their colours first appeared early in life. We have instances of this fact in all black and white breeds, in which the young and old of both sexes are alike; nor can it be maintained that there is something peculiar in a black or white plumage, leading to its transference to both sexes; for the males alone of many natural species are either black or white, the females being very differently coloured. With the so-called Cuckoo sub-breeds of the fowl, in which the feathers are transversely pencilled with dark stripes, both sexes and the chickens are coloured in nearly the same manner. The laced plumage of the Sebright bantam is the same in both sexes, and in the chickens the feathers are tipped with black, which makes a near approach to lacing. Spangled Hamburghs, however, offer a partial exception, for the two sexes, though not quite alike, resemble each other more closely than do the sexes of the aboriginal parent-species, yet they acquire their characteristic plumage late in life, for the chickens are distinctly pencilled. Turning to other characters besides colour: the males alone of the wild parent-species and of most domestic breeds possess a fairly well developed comb, but in the young of the Spanish fowl it is largely developed at a very early age, and apparently in consequence of this it is of unusual size in the adult females. In the Game breeds pugnacity is developed at a wonderfully early age, of which curious proofs could be given; and this character is transmitted to both sexes, so that the hens, from their extreme pugnacity, are now generally exhibited in separate pens. With the Polish breeds the bony protuberance of the skull which supports the crest is partially developed even before the chickens are hatched, and the crest itself soon begins to grow, though at first feebly;366 and in this breed a great bony protuberance and an immense crest characterise the adults of both sexes.