Extinction follows chiefly from the compet.i.tion of tribe with tribe, and race with race. Various checks are always in action, as specified in a former chapter, which serve to keep down the numbers of each savage tribe,-such as periodical famines, the wandering of the parents and the consequent deaths of infants, prolonged suckling, the stealing of women, wars, accidents, sickness, licentiousness, especially infanticide, and, perhaps, lessened fertility from less nutritious food, and many hardships. If from any cause any one of these checks is lessened, even in a slight degree, the tribe thus favoured will tend to increase; and when one of two adjoining tribes becomes more numerous and powerful than the other, the contest is soon settled by war, slaughter, cannibalism, slavery, and absorption. Even when a weaker tribe is not thus abruptly swept away, if it once begins to decrease, it generally goes on decreasing until it is extinct.[312]

When civilised nations come into contact with barbarians the struggle is short, except where a deadly climate gives its aid to the native race.

Of the causes which lead to the victory of civilised nations, some are plain and some very obscure. We can see that the cultivation of the land will be fatal in many ways to savages, for they cannot, or will not, change their habits. New diseases and vices are highly destructive; and it appears that in every nation a new disease causes much death, until those who are most susceptible to its destructive influence are gradually weeded out;[313] and so it may be with the evil effects from spirituous liquors, as well as with the unconquerably strong taste for them shewn by so many savages. It further appears, mysterious as is the fact, that the first meeting of distinct and separated people generates disease.[314] Mr. Sproat, who in Vancouver Island closely attended to the subject of extinction, believes that changed habits of life, which always follow from the advent of Europeans, induces much ill-health. He lays, also, great stress on so trifling a cause as that the natives become "bewildered and dull by the new life around them; they lose the motives for exertion, and get no new ones in their place."[315]

The grade of civilisation seems a most important element in the success of nations which come in compet.i.tion. A few centuries ago Europe feared the inroads of Eastern barbarians; now, any such fear would be ridiculous. It is a more curious fact, that savages did not formerly waste away, as Mr. Bagehot has remarked, before the cla.s.sical nations, as they now do before modern civilised nations; had they done so, the old moralists would have mused over the event; but there is no lament in any writer of that period over the perishing barbarians.[316]

Although the gradual decrease and final extinction of the races of man is an obscure problem, we can see that it depends on many causes, differing in different places and at different times. It is the same difficult problem as that presented by the extinction of one of the higher animals-of the fossil horse, for instance, which disappeared from South America, soon afterwards to be replaced, within the same districts, by countless troops of the Spanish horse. The New Zealander seems conscious of this parallelism, for he compares his future fate with that of the native rat almost exterminated by the European rat. The difficulty, though great to our imagination, and really great if we wish to ascertain the precise causes, ought not to be so to our reason, as long as we keep steadily in mind that the increase of each species and each race is constantly hindered by various checks; so that if any new check, or cause of destruction, even a slight one, be superadded, the race will surely decrease in number; and as it has everywhere been observed that savages are much opposed to any change of habits, by which means injurious checks could be counterbalanced, decreasing numbers will sooner or later lead to extinction; the end, in most cases, being promptly determined by the inroads of increasing and conquering tribes.

_On the Formation of the Races of Man._-It may be premised that when we find the same race, though broken up into distinct tribes, ranging over a great area, as over America, we may attribute their general resemblance to descent from a common stock. In some cases the crossing of races already, distinct has led to the formation of new races. The singular fact that Europeans and Hindoos, who belong to the same Aryan stock and speak a language fundamentally the same, differ widely in appearance, whilst Europeans differ but little from Jews, who belong to the Semitic stock and speak quite another language, has been accounted for by Broca[317] through the Aryan branches having been largely crossed during their wide diffusion by various indigenous tribes. When two races in close contact cross, the first result is a heterogeneous mixture: thus Mr. Hunter, in describing the Santali or hill-tribes of India, says that hundreds of imperceptible gradations may be traced "from the black, squat tribes of the mountains to the tall olive-coloured Brahman, with his intellectual brow, calm eyes, and high but narrow head;" so that it is necessary in courts of justice to ask the witnesses whether they are Santalis or Hindoos.[318] Whether a heterogeneous people, such as the inhabitants of some of the Polynesian islands, formed by the crossing of two distinct races, with few or no pure members left, would ever become h.o.m.ogeneous, is not known from direct evidence. But as with our domesticated animals, a crossed breed can certainly, in the course of a few generations, be fixed and made uniform by careful selection,[319] we may infer that the free and prolonged intercrossing during many generations of a heterogeneous mixture would supply the place of selection, and overcome any tendency to reversion, so that a crossed race would ultimately become h.o.m.ogeneous, though it might not partake in an equal degree of the characters of the two parent-races.

Of all the differences between the races of man, the colour of the skin is the most conspicuous and one of the best marked. Differences of this kind, it was formerly thought, could be accounted for by long exposure under different climates; but Pallas first shewed that this view is not tenable, and he has been followed by almost all anthropologists.[320]

The view has been rejected chiefly because the distribution of the variously coloured races, most of whom must have long inhabited their present homes, does not coincide with corresponding differences of climate. Weight must also be given to such cases as that of the Dutch families, who, as we hear on excellent authority,[321] have not undergone the least change of colour, after residing for three centuries in South Africa. The uniform appearance in various parts of the world of gypsies and Jews, though the uniformity of the latter has been somewhat exaggerated,[322] is likewise an argument on the same side. A very damp or a very dry atmosphere has been supposed to be more influential in modifying the colour of the skin than mere heat; but as D'Orbigny in South America, and Livingstone in Africa, arrived at diametrically opposite conclusions with respect to dampness and dryness, any conclusion on this head must be considered as very doubtful.[323]

Various facts, which I have elsewhere given, prove that the colour of the skin and hair is sometimes correlated in a surprising manner with a complete immunity from the action of certain vegetable poisons and from the attacks of certain parasites. Hence it occurred to me, that negroes and other dark races might have acquired their dark tints by the darker individuals escaping during a long series of generations from the deadly influence of the miasmas of their native countries.

I afterwards found that the same idea had long ago occurred to Dr.

Wells.[324] That negroes, and even mulattoes, are almost completely exempt from the yellow-fever, which is so destructive in tropical America, has long been known.[325] They likewise escape to a large extent the fatal intermittent fevers that prevail along, at least, 2600 miles of the sh.o.r.es of Africa, and which annually cause one-fifth of the white settlers to die, and another fifth to return home invalided.[326]

This immunity in the negro seems to be partly inherent, depending on some unknown peculiarity of const.i.tution, and partly the result of acclimatisation. Pouchet[327] states that the negro regiments, borrowed from the Viceroy of Egypt for the Mexican war, which had been recruited near the Soudan, escaped the yellow-fever almost equally well with the negroes originally brought from various parts of Africa, and accustomed to the climate of the West Indies. That acclimatisation plays a part is shewn by the many cases in which negroes, after having resided for some time in a colder climate, have become to a certain extent liable to tropical fevers.[328] The nature of the climate under which the white races have long resided, likewise has some influence on them; for during the fearful epidemic of yellow-fever in Demerara during 1837, Dr. Blair found that the death-rate of the immigrants was proportional to the lat.i.tude of the country whence they had come. With the negro the immunity, as far as it is the result of acclimatisation, implies exposure during a prodigious length of time; for the aborigines of tropical America, who have resided there from time immemorial, are not exempt from yellow-fever; and the Rev. B. Tristram states, that there are districts in Northern Africa which the native inhabitants are compelled annually to leave, though the negroes can remain with safety.

That the immunity of the negro is in any degree correlated with the colour of his skin is a mere conjecture: it may be correlated with some difference in his blood, nervous system, or other tissues. Nevertheless, from the facts above alluded to, and from some connection apparently existing between complexion and a tendency to consumption, the conjecture seemed to me not improbable. Consequently I endeavoured, with but little success,[329] to ascertain how far it held good. The late Dr. Daniell, who had long lived on the West Coast of Africa, told me that he did not believe in any such relation. He was himself unusually fair, and had withstood the climate in a wonderful manner. When he first arrived as a boy on the coast, an old and experienced negro chief predicted from his appearance that this would prove the case. Dr.

Nicholson, of Antigua, after having attended to this subject, wrote to me that he did not think that dark-coloured Europeans escaped the yellow-fever better than those that were light-coloured. Mr. J. M.

Harris altogether denies[330] that Europeans with dark hair withstand a hot climate better than other men; on the contrary, experience has taught him in making a selection of men for service on the coast of Africa, to choose those with red hair. As far, therefore, as these slight indications serve, there seems no foundation for the hypothesis, which has been accepted by several writers, that the colour of the black races may have resulted from darker and darker individuals having survived in greater numbers, during their exposure to the fever-generating miasmas of their native countries.

Although with our present knowledge we cannot account for the strongly-marked differences in colour between the races of man, either through correlation with const.i.tutional peculiarities, or through the direct action of climate; yet we must not quite ignore the latter agency, for there is good reason to believe that some inherited effect is thus produced.[331]

We have seen in our third chapter that the conditions of life, such as abundant food and general comfort, affect in a direct manner the development of the bodily frame, the effects being transmitted. Through the combined influences of climate and changed habits of life, European settlers, in the United States undergo, as is generally admitted, a slight but extraordinarily rapid change of appearance. There is, also, a considerable body of evidence shewing that in the Southern States the house-slaves of the third generation present a markedly different appearance from the field-slaves.[332]

If, however, we look to the races of man, as distributed over the world, we must infer that their characteristic differences cannot be accounted for by the direct action of different conditions of life, even after exposure to them for an enormous period of time. The Esquimaux live exclusively on animal food; they are clothed in thick fur, and are exposed to intense cold and to prolonged darkness; yet they do not differ in any extreme degree from the inhabitants of Southern China, who live entirely on vegetable food and are exposed almost naked to a hot, glaring climate. The unclothed Fuegians live on the marine productions of their inhospitable sh.o.r.es; the Botocudos of Brazil wander about the hot forests of the interior and live chiefly on vegetable productions; yet these tribes resemble each other so closely that the Fuegians on board the "Beagle" were mistaken by some Brazilians for Botocudos. The Botocudos again, as well as the other inhabitants of tropical America, are wholly different from the Negroes who inhabit the opposite sh.o.r.es of the Atlantic, are exposed to a nearly similar climate, and follow nearly the same habits of life.

Nor can the differences between the races of man be accounted for, except to a quite insignificant degree, by the inherited effects of the increased or decreased use of parts. Men who habitually live in canoes, may have their legs somewhat stunted; those who inhabit lofty regions have their chests enlarged; and those who constantly use certain sense-organs have the cavities in which they are lodged somewhat increased in size, and their features consequently a little modified.

With civilised nations, the reduced size of the jaws from lessened use, the habitual play of different muscles serving to express different emotions, and the increased size of the brain from greater intellectual activity, have together produced a considerable effect on their general appearance in comparison with savages.[333] It is also possible that increased bodily stature, with no corresponding increase in the size of the brain, may have given to some races (judging from the previously adduced cases of the rabbits) an elongated skull of the dolichocephalic type.

Lastly, the little-understood principle of correlation will almost certainly have come into action, as in the case of great muscular development and strongly projecting supra-orbital ridges. It is not improbable that the texture of the hair, which differs much in the different races, may stand in some kind of correlation with the structure of the skin; for the colour of the hair and skin are certainly correlated, as is its colour and texture with the Mandans.[334] The colour of the skin and the odour emitted by it are likewise in some manner connected. With the breeds of sheep the number of hairs within a given s.p.a.ce and the number of the excretory pores stand in some relation to each other.[335] If we may judge from the a.n.a.logy of our domesticated animals, many modifications of structure in man probably come under this principle of correlated growth.

We have now seen that the characteristic differences between the races of man cannot be accounted for in a satisfactory manner by the direct action of the conditions of life, nor by the effects of the continued use of parts, nor through the principle of correlation. We are therefore led to inquire whether slight individual differences, to which man is eminently liable, may not have been preserved and augmented during a long series of generations through natural selection. But here we are at once met by the objection that beneficial variations alone can be thus preserved; and as far as we are enabled to judge (although always liable to error on this head) not one of the external differences between the races of man are of any direct or special service to him. The intellectual and moral or social faculties must of course be excepted from this remark; but differences in these faculties can have had little or no influence on external characters. The variability of all the characteristic differences between the races, before referred to, likewise indicates that these differences cannot be of much importance; for, had they been important, they would long ago have been either fixed and preserved, or eliminated. In this respect man resembles those forms, called by naturalists protean or polymorphic, which have remained extremely variable, owing, as it seems, to their variations being of an indifferent nature, and consequently to their having escaped the action of natural selection.

We have thus far been baffled in all our attempts to account for the differences between the races of man; but there remains one important agency, namely s.e.xual Selection, which appears to have acted as powerfully on man, as on many other animals. I do not intend to a.s.sert that s.e.xual selection will account for all the differences between the races. An unexplained residuum is left, about which we can in our ignorance only say, that as individuals are continually born with, for instance, heads a little rounder or narrower, and with noses a little longer or shorter, such slight differences might become fixed and uniform, if the unknown agencies which induced them were to act in a more constant manner, aided by long-continued intercrossing. Such modifications come under the provisional cla.s.s, alluded to in our fourth chapter, which for the want of a better term have been called spontaneous variations. Nor do I pretend that the effects of s.e.xual selection can be indicated with scientific precision; but it can be shewn that it would be an inexplicable fact if man had not been modified by this agency, which has acted so powerfully on innumerable animals, both high and low in the scale. It can further be shewn that the differences between the races of man, as in colour, hairyness, form of features, &c., are of the nature which it might have been expected would have been acted on by s.e.xual selection. But in order to treat this subject in a fitting manner, I have found it necessary to pa.s.s the whole animal kingdom in review; I have therefore devoted to it the Second Part of this work. At the close I shall return to man, and, after attempting to shew how far he has been modified through s.e.xual selection, will give a brief summary of the chapters in this First Part.

PART II.-s.e.xUAL SELECTION.

CHAPTER VIII.

PRINCIPLES OF s.e.xUAL SELECTION.

Secondary s.e.xual characters-s.e.xual selection-Manner of action-Excess of males-Polygamy-The male alone generally modified through s.e.xual selection-Eagerness of the male-Variability of the male-Choice exerted by the female-s.e.xual compared with natural selection-Inheritance, at corresponding periods of life, at corresponding seasons of the year, and as limited by s.e.x-Relations between the several forms of inheritance-Causes why one s.e.x and the young are not modified through s.e.xual selection-Supplement on the proportional numbers of the two s.e.xes throughout the animal kingdom-On the limitation of the numbers of the two s.e.xes through natural selection.

With animals which have their s.e.xes separated, the males necessarily differ from the females in their organs of reproduction; and these afford the primary s.e.xual characters. But the s.e.xes often differ in what Hunter has called secondary s.e.xual 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 dest.i.tute, 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 a.n.a.logous 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 larvae 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 s.e.xual 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 s.e.xual 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 (Culicidae and Tabanidae) are blood-suckers, whilst the males live on flowers and have their mouths dest.i.tute 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 dest.i.tute 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 dest.i.tute of wings, as are many female moths, some of which never leave their coc.o.o.ns. 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;[337] but the meaning of this and of many a.n.a.logous differences, is not at all understood. Differences of structure between the two s.e.xes 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 mult.i.tude 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 s.e.xual selection. This depends on the advantage which certain individuals have over other individuals of the same s.e.x and species, in exclusive relation to reproduction. When the two s.e.xes 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 s.e.x.

So again the primary s.e.xual 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, _caeteris 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 antennae 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 s.e.xes 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 s.e.xual 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 s.e.xual 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 a.s.saulted by them, these organs will have been perfected through s.e.xual 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 s.e.xual selection. Whole chapters could easily be filled with details on the differences between the s.e.xes 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 pa.s.s them over, giving only a few instances under each cla.s.s.

There are many other structures and instincts which must have been developed through s.e.xual 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 s.e.xual 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 s.e.xual 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 s.e.x. 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 a.s.sembled 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-c.o.c.ks by the selection of those birds which are victorious in the c.o.c.kpit, 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 s.e.xual selection; and it is certain that secondary s.e.xual 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 s.e.xual 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 s.e.xual 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 a.s.sert 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 cla.s.s of insects the males almost always emerge from the pupal state before the other s.e.x, 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 const.i.tutions. On the whole there can be no doubt that with almost all animals, in which the s.e.xes are separate, there is a constantly recurrent struggle between the males for the possession of the females.

Our difficulty in regard to s.e.xual 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 s.e.xual selection. When the s.e.xes 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 s.e.xual 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 s.e.xes, I do not believe that any great inequality in number commonly exists. In most cases s.e.xual 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 r.e.t.a.r.ded females, which would be compelled, supposing the s.e.xes 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 mult.i.tude 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 s.e.xes mutually preferred and selected certain individuals of the opposite s.e.x; supposing that they selected not only the more attractive, but likewise the more vigorous individuals.

_Numerical Proportion of the Two s.e.xes._-I have remarked that s.e.xual 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 s.e.xes 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 s.e.xes 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 997 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 1101 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 s.e.xes. Thus with mankind, the male births in England are as 1045, in Russia as 1089, and with the Jews of Livornia as 120 _to_ 100 females. The proportion is also mysteriously affected by the circ.u.mstance of the births being legitimate or illegitimate.

For our present purpose we are concerned with the proportion of the s.e.xes, 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 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 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 s.e.xes 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 s.e.xes fluctuates slightly during successive years: thus with race-horses, for every 100 females born, the males 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 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 s.e.xual 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 s.e.xes 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 s.e.xes; for if each male secures two or more females, many males will not be able to pair; and the latter a.s.suredly will be the weaker or less attractive individuals. Many mammals and some few birds are polygamous, but with animals belonging to the lower cla.s.ses 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 s.e.xual characters, appears nearly certain; and this supports the view that a numerical preponderance of males would be eminently favourable to the action of s.e.xual selection. Nevertheless many animals, especially birds, which are strictly monogamous, display strongly-marked secondary s.e.xual characters; whilst some few animals, which are polygamous, are not thus characterised.

We will first briefly run through the cla.s.s 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 s.e.xual 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 s.e.xual 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 Pallas[341] 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 s.e.xual 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, pa.s.ses much of his time in solitude; but when a.s.sociating 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 s.e.xes 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 s.e.xual 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.