The Descent of Man, and Selection in Relation to Sex

by Charles Darwin

ON THE FORMATION OF THE RACES OF MAN.

In some cases the crossing of distinct races has led to the formation of a new race. The singular fact that the 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 (49. ‘On Anthropology,’ translation, ‘Anthropological Review,’ Jan. 1868, p. 38.), through certain Aryan branches having been largely crossed by indigenous tribes during their wide diffusion. 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. (50. ‘The Annals of Rural Bengal,’ 1868, p. 134.) 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 homogeneous, is not known from direct evidence. But as with our domesticated animals, a cross-breed can certainly be fixed and made uniform by careful selection (51. ‘The Variation of Animals and Plants under Domestication,’ vol. ii. p. 95.) in the course of a few generations, we may infer that the free intercrossing of a heterogeneous mixture during a long descent would supply the place of selection, and overcome any tendency to reversion; so that the crossed race would ultimately become homogeneous, 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. It was formerly thought that differences of this kind could be accounted for by long exposure to different climates; but Pallas first shewed that this is not tenable, and he has since been followed by almost all anthropologists. (52. Pallas, ‘Act. Acad. St. Petersburg,’ 1780, part ii. p. 69. He was followed by Rudolphi, in his ‘Beytrage zur Anthropologie,’ 1812. An excellent summary of the evidence is given by Godron, ‘De l’Espèce,’ 1859, vol. ii. p. 246, etc.) This 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. Some little weight may be given to such cases as that of the Dutch families, who, as we hear on excellent authority (53. Sir Andrew Smith, as quoted by Knox, ‘Races of Man,’ 1850, p. 473.), have not undergone the least change of colour after residing for three centuries in South Africa. An argument on the same side may likewise be drawn from the uniform appearance in various parts of the world of gipsies and Jews, though the uniformity of the latter has been somewhat exaggerated. (54. See De Quatrefages on this head, ‘Revue des Cours Scientifiques,’ Oct. 17, 1868, p. 731.) 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. (55. Livingstone’s ‘Travels and Researches in S. Africa,’ 1857, pp. 338, 339. D’Orbigny, as quoted by Godron, ‘De l’Espece,’ vol. ii. p. 266.)

Various facts, which I have given elsewhere, 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 from the deadly influence of the miasma of their native countries, during a long series of generations.

I afterwards found that this same idea had long ago occurred to Dr. Wells. (56. See a paper read before the Royal Soc. in 1813, and published in his Essays in 1818. I have given an account of Dr. Wells’ views in the Historical Sketch (p. xvi.) to my ‘Origin of Species.’ Various cases of colour correlated with constitutional peculiarities are given in my ‘Variation of Animals and Plants under Domestication,’ vol. ii. pp. 227, 335.) It has long been known that negroes, and even mulattoes, are almost completely exempt from the yellow-fever, so destructive in tropical America. (57. See, for instance, Nott and Gliddon, ‘Types of Mankind,’ p. 68.) They likewise escape to a large extent the fatal intermittent fevers, that prevail along at least 2600 miles of the shores of Africa, and which annually cause one-fifth of the white settlers to die, and another fifth to return home invalided. (58. Major Tulloch, in a paper read before the Statistical Society, April 20, 1840, and given in the ‘Athenaeum,’ 1840, p. 353.) This immunity in the negro seems to be partly inherent, depending on some unknown peculiarity of constitution, and partly the result of acclimatisation. Pouchet (59. ‘The Plurality of the Human Race’ (translat.), 1864, p. 60.) states that the negro regiments recruited near the Soudan, and borrowed from the Viceroy of Egypt for the Mexican war, escaped the yellow-fever almost equally 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 have become somewhat liable to tropical fevers, after having resided for some time in a colder climate. (60. Quatrefages, ‘Unité de l’Espèce Humaine,’ 1861, p. 205. Waitz, ‘Introduction to Anthropology,’ translat., vol. i. 1863, p. 124. Livingstone gives analogous cases in his ‘Travels.’) 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 latitude 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. H.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 (61. In the spring of 1862 I obtained permission from the Director-General of the Medical department of the Army, to transmit to the surgeons of the various regiments on foreign service a blank table, with the following appended remarks, but I have received no returns. “As several well-marked cases have been recorded with our domestic animals of a relation between the colour of the dermal appendages and the constitution; and it being notorious that there is some limited degree of relation between the colour of the races of man and the climate inhabited by them; the following investigation seems worth consideration. Namely, whether there is any relation in Europeans between the colour of their hair, and their liability to the diseases of tropical countries. If the surgeons of the several regiments, when stationed in unhealthy tropical districts, would be so good as first to count, as a standard of comparison, how many men, in the force whence the sick are drawn, have dark and light-coloured hair, and hair of intermediate or doubtful tints; and if a similar account were kept by the same medical gentlemen, of all the men who suffered from malarious and yellow fevers, or from dysentery, it would soon be apparent, after some thousand cases had been tabulated, whether there exists any relation between the colour of the hair and constitutional liability to tropical diseases. Perhaps no such relation would be discovered, but the investigation is well worth making. In case any positive result were obtained, it might be of some practical use in selecting men for any particular service. Theoretically the result would be of high interest, as indicating one means by which a race of men inhabiting from a remote period an unhealthy tropical climate, might have become dark-coloured by the better preservation of dark-haired or dark-complexioned individuals during a long succession of generations.”), to ascertain how far it holds 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, writes to me that dark-coloured Europeans escape the yellow fever more than those that are light-coloured. Mr. J.M. Harris altogether denies 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. (62. ‘Anthropological Review,’ Jan. 1866, p. xxi. Dr. Sharpe also says, with respect to India (‘Man a Special Creation,’ 1873, p. 118), “that it has been noticed by some medical officers that Europeans with light hair and florid complexions suffer less from diseases of tropical countries than persons with dark hair and sallow complexions; and, so far as I know, there appear to be good grounds for this remark.” On the other hand, Mr. Heddle, of Sierra Leone, “who has had more clerks killed under him than any other man,” by the climate of the West African Coast (W. Reade, ‘African Sketch Book,’ vol. ii. p. 522), holds a directly opposite view, as does Capt. Burton.) As far, therefore, as these slight indications go, there seems no foundation for the hypothesis, that blackness has resulted from the darker and darker individuals having survived better during long exposure to fever-generating miasma.

Dr. Sharpe remarks (63. ‘Man a Special Creation,’ 1873, p. 119.), that a tropical sun, which burns and blisters a white skin, does not injure a black one at all; and, as he adds, this is not due to habit in the individual, for children only six or eight months old are often carried about naked, and are not affected. I have been assured by a medical man, that some years ago during each summer, but not during the winter, his hands became marked with light brown patches, like, although larger than freckles, and that these patches were never affected by sun-burning, whilst the white parts of his skin have on several occasions been much inflamed and blistered. With the lower animals there is, also, a constitutional difference in liability to the action of the sun between those parts of the skin clothed with white hair and other parts. (64. ‘Variation of Animals and Plants under Domestication,’ vol. ii. pp. 336, 337.) Whether the saving of the skin from being thus burnt is of sufficient importance to account for a dark tint having been gradually acquired by man through natural selection, I am unable to judge. If it be so, we should have to assume that the natives of tropical America have lived there for a much shorter time than the Negroes in Africa, or the Papuans in the southern parts of the Malay archipelago, just as the lighter-coloured Hindoos have resided in India for a shorter time than the darker aborigines of the central and southern parts of the peninsula.

Although with our present knowledge we cannot account for the differences of colour in the races of man, through any advantage thus gained, or from 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. (65. See, for instance, Quatrefages (‘Revue des Cours Scientifiques,’ Oct. 10, 1868, p. 724) on the effects of residence in Abyssinia and Arabia, and other analogous cases. Dr. Rolle (‘Der Mensch, seine Abstammung,’ etc., 1865, s. 99) states, on the authority of Khanikof, that the greater number of German families settled in Georgia, have acquired in the course of two generations dark hair and eyes. Mr. D. Forbes informs me that the Quichuas in the Andes vary greatly in colour, according to the position of the valleys inhabited by them.)

We have seen in the second chapter that the conditions of life affect the development of the bodily frame in a direct manner, and that the effects are transmitted. Thus, as is generally admitted, the European settlers in the United States undergo a slight but extraordinary rapid change of appearance. Their bodies and limbs become elongated; and I hear from Col. Bernys that during the late war in the United States, good evidence was afforded of this fact by the ridiculous appearance presented by the German regiments, when dressed in ready-made clothes manufactured for the American market, and which were much too long for the men in every way. 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. (66. Harlan, ‘Medical Researches,’ p. 532. Quatrefages (‘Unité de l’Espèce Humaine,’ 1861, p. 128) has collected much evidence on this head.)

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 shores; 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 shores 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 by the inherited effects of the increased or decreased use of parts, except to a quite insignificant degree. Men who habitually live in canoes, may have their legs somewhat stunted; those who inhabit lofty regions may have their chests enlarged; and those who constantly use certain sense-organs may 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 when compared with savages. (67. See Prof. Schaaffhausen, translat., in ‘Anthropological Review,’ Oct. 1868, p. 429.) Increased bodily stature, without any corresponding increase in the size of the brain, may (judging from the previously adduced case of rabbits), have given to some races an elongated skull of the dolichocephalic type.

Lastly, the little-understood principle of correlated development has sometimes come into action, as in the case of great muscular development and strongly projecting supra-orbital ridges. The colour of the skin and hair are plainly correlated, as is the texture of the hair with its colour in the Mandans of North America. (68. Mr. Catlin states (‘N. American Indians,’ 3rd ed., 1842, vol. i. p. 49) that in the whole tribe of the Mandans, about one in ten or twelve of the members, of all ages and both sexes, have bright silvery grey hair, which is hereditary. Now this hair is as coarse and harsh as that of a horse’s mane, whilst the hair of other colours is fine and soft.) The colour also 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 space and the number of excretory pores are related. (69. On the odour of the skin, Godron, ‘Sur l’Espèce,’ tom. ii. p. 217. On the pores in the skin, Dr. Wilckens, ‘Die Aufgaben der Landwirth. Zootechnik,’ 1869, s. 7.) If we may judge from the analogy of our domesticated animals, many modifications of structure in man probably come under this principle of correlated development.

We have now seen that the external 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 enquire 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 err on this head, none of the 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. The great variability of all the external differences between the races of man, likewise indicates that they cannot be of much importance; for if 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 such variations being of an indifferent nature, and to their having thus 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 Sexual Selection, which appears to have acted powerfully on man, as on many other animals. I do not intend to assert that sexual selection will account for all the differences between the races. An unexplained residuum is left, about which we can only say, in our ignorance, 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 variations come under the provisional class, alluded to in our second chapter, which for want of a better term are often called spontaneous. Nor do I pretend that the effects of sexual 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 appears to have acted powerfully on innumerable animals. It can further be shewn that the differences between the races of man, as in colour, hairiness, form of features, etc., are of a kind which might have been expected to come under the influence of sexual selection. But in order to treat this subject properly, I have found it necessary to pass 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 sexual selection, will give a brief summary of the chapters in this First Part.

NOTE ON THE RESEMBLANCES AND DIFFERENCES IN THE STRUCTURE AND THE DEVELOPMENT OF THE BRAIN IN MAN AND APES BY PROFESSOR HUXLEY, F.R.S.

The controversy respecting the nature and the extent of the differences in the structure of the brain in man and the apes, which arose some fifteen years ago, has not yet come to an end, though the subject matter of the dispute is, at present, totally different from what it was formerly. It was originally asserted and re-asserted, with singular pertinacity, that the brain of all the apes, even the highest, differs from that of man, in the absence of such conspicuous structures as the posterior lobes of the cerebral hemispheres, with the posterior cornu of the lateral ventricle and the hippocampus minor, contained in those lobes, which are so obvious in man.

But the truth that the three structures in question are as well developed in apes’ as in human brains, or even better; and that it is characteristic of all the Primates (if we exclude the Lemurs) to have these parts well developed, stands at present on as secure a basis as any proposition in comparative anatomy. Moreover, it is admitted by every one of the long series of anatomists who, of late years, have paid special attention to the arrangement of the complicated sulci and gyri which appear upon the surface of the cerebral hemispheres in man and the higher apes, that they are disposed after the very same pattern in him, as in them. Every principal gyrus and sulcus of a chimpanzee’s brain is clearly represented in that of a man, so that the terminology which applies to the one answers for the other. On this point there is no difference of opinion. Some years since, Professor Bischoff published a memoir (70. ‘Die Grosshirn-Windungen des Menschen;’ ‘Abhandlungen der K. Bayerischen Akademie,’ B. x. 1868.) on the cerebral convolutions of man and apes; and as the purpose of my learned colleague was certainly not to diminish the value of the differences between apes and men in this respect, I am glad to make a citation from him.

“That the apes, and especially the orang, chimpanzee and gorilla, come very close to man in their organisation, much nearer than to any other animal, is a well known fact, disputed by nobody. Looking at the matter from the point of view of organisation alone, no one probably would ever have disputed the view of Linnaeus, that man should be placed, merely as a peculiar species, at the head of the mammalia and of those apes. Both shew, in all their organs, so close an affinity, that the most exact anatomical investigation is needed in order to demonstrate those differences which really exist. So it is with the brains. The brains of man, the orang, the chimpanzee, the gorilla, in spite of all the important differences which they present, come very close to one another” (loc. cit. p. 101).

There remains, then, no dispute as to the resemblance in fundamental characters, between the ape’s brain and man’s: nor any as to the wonderfully close similarity between the chimpanzee, orang and man, in even the details of the arrangement of the gyri and sulci of the cerebral hemispheres. Nor, turning to the differences between the brains of the highest apes and that of man, is there any serious question as to the nature and extent of these differences. It is admitted that the man’s cerebral hemispheres are absolutely and relatively larger than those of the orang and chimpanzee; that his frontal lobes are less excavated by the upward protrusion of the roof of the orbits; that his gyri and sulci are, as a rule, less symmetrically disposed, and present a greater number of secondary plications. And it is admitted that, as a rule, in man, the temporo-occipital or “external perpendicular” fissure, which is usually so strongly marked a feature of the ape’s brain is but faintly marked. But it is also clear, that none of these differences constitutes a sharp demarcation between the man’s and the ape’s brain. In respect to the external perpendicular fissure of Gratiolet, in the human brain for instance, Professor Turner remarks: (71. ‘Convolutions of the Human Cerebrum Topographically Considered,’ 1866, p. 12.)

“In some brains it appears simply as an indentation of the margin of the hemisphere, but, in others, it extends for some distance more or less transversely outwards. I saw it in the right hemisphere of a female brain pass more than two inches outwards; and on another specimen, also the right hemisphere, it proceeded for four-tenths of an inch outwards, and then extended downwards, as far as the lower margin of the outer surface of the hemisphere. The imperfect definition of this fissure in the majority of human brains, as compared with its remarkable distinctness in the brain of most Quadrumana, is owing to the presence, in the former, of certain superficial, well marked, secondary convolutions which bridge it over and connect the parietal with the occipital lobe. The closer the first of these bridging gyri lies to the longitudinal fissure, the shorter is the external parieto-occipital fissure” (loc. cit. p. 12).

The obliteration of the external perpendicular fissure of Gratiolet, therefore, is not a constant character of the human brain. On the other hand, its full development is not a constant character of the higher ape’s brain. For, in the chimpanzee, the more or less extensive obliteration of the external perpendicular sulcus by “bridging convolutions,” on one side or the other, has been noted over and over again by Prof. Rolleston, Mr. Marshall, M. Broca and Professor Turner. At the conclusion of a special paper on this subject the latter writes: (72. Notes more especially on the bridging convolutions in the Brain of the Chimpanzee, ‘Proceedings of the Royal Society of Edinburgh,’ 1865-6.)

“The three specimens of the brain of a chimpanzee, just described, prove, that the generalisation which Gratiolet has attempted to draw of the complete absence of the first connecting convolution and the concealment of the second, as essentially characteristic features in the brain of this animal, is by no means universally applicable. In only one specimen did the brain, in these particulars, follow the law which Gratiolet has expressed. As regards the presence of the superior bridging convolution, I am inclined to think that it has existed in one hemisphere, at least, in a majority of the brains of this animal which have, up to this time, been figured or described. The superficial position of the second bridging convolution is evidently less frequent, and has as yet, I believe, only been seen in the brain (A) recorded in this communication. The asymmetrical arrangement in the convolutions of the two hemispheres, which previous observers have referred to in their descriptions, is also well illustrated in these specimens” (pp. 8, 9).

Even were the presence of the temporo-occipital, or external perpendicular, sulcus, a mark of distinction between the higher apes and man, the value of such a distinctive character would be rendered very doubtful by the structure of the brain in the Platyrrhine apes. In fact, while the temporo-occipital is one of the most constant of sulci in the Catarrhine, or Old World, apes, it is never very strongly developed in the New World apes; it is absent in the smaller Platyrrhini; rudimentary in Pithecia (73. Flower, ‘On the Anatomy of Pithecia Monachus,’ ‘Proceedings of the Zoological Society,’ 1862.); and more or less obliterated by bridging convolutions in Ateles.

A character which is thus variable within the limits of a single group can have no great taxonomic value.

It is further established, that the degree of asymmetry of the convolution of the two sides in the human brain is subject to much individual variation; and that, in those individuals of the Bushman race who have been examined, the gyri and sulci of the two hemispheres are considerably less complicated and more symmetrical than in the European brain, while, in some individuals of the chimpanzee, their complexity and asymmetry become notable. This is particularly the case in the brain of a young male chimpanzee figured by M. Broca. (‘L’ordre des Primates,’ p. 165, fig. 11.)

Again, as respects the question of absolute size, it is established that the difference between the largest and the smallest healthy human brain is greater than the difference between the smallest healthy human brain and the largest chimpanzee’s or orang’s brain.

Moreover, there is one circumstance in which the orang’s and chimpanzee’s brains resemble man’s, but in which they differ from the lower apes, and that is the presence of two corpora candicantia—the Cynomorpha having but one.

In view of these facts I do not hesitate in this year 1874, to repeat and insist upon the proposition which I enunciated in 1863: (74. ‘Man’s Place in Nature,’ p. 102.)

“So far as cerebral structure goes, therefore, it is clear that man differs less from the chimpanzee or the orang, than these do even from the monkeys, and that the difference between the brain of the chimpanzee and of man is almost insignificant when compared with that between the chimpanzee brain and that of a Lemur.”

In the paper to which I have referred, Professor Bischoff does not deny the second part of this statement, but he first makes the irrelevant remark that it is not wonderful if the brains of an orang and a Lemur are very different; and secondly, goes on to assert that, “If we successively compare the brain of a man with that of an orang; the brain of this with that of a chimpanzee; of this with that of a gorilla, and so on of a Hylobates, Semnopithecus, Cynocephalus, Cercopithecus, Macacus, Cebus, Callithrix, Lemur, Stenops, Hapale, we shall not meet with a greater, or even as great a, break in the degree of development of the convolutions, as we find between the brain of a man and that of an orang or chimpanzee.”

To which I reply, firstly, that whether this assertion be true or false, it has nothing whatever to do with the proposition enunciated in ‘Man’s Place in Nature,’ which refers not to the development of the convolutions alone, but to the structure of the whole brain. If Professor Bischoff had taken the trouble to refer to p. 96 of the work he criticises, in fact, he would have found the following passage: “And it is a remarkable circumstance that though, so far as our present knowledge extends, there IS one true structural break in the series of forms of Simian brains, this hiatus does not lie between man and the manlike apes, but between the lower and the lowest Simians, or in other words, between the Old and New World apes and monkeys and the Lemurs. Every Lemur which has yet been examined, in fact, has its cerebellum partially visible from above; and its posterior lobe, with the contained posterior cornu and hippocampus minor, more or less rudimentary. Every marmoset, American monkey, Old World monkey, baboon or manlike ape, on the contrary, has its cerebellum entirely hidden, posteriorly, by the cerebral lobes, and possesses a large posterior cornu with a well-developed hippocampus minor.”

This statement was a strictly accurate account of what was known when it was made; and it does not appear to me to be more than apparently weakened by the subsequent discovery of the relatively small development of the posterior lobes in the Siamang and in the Howling monkey. Notwithstanding the exceptional brevity of the posterior lobes in these two species, no one will pretend that their brains, in the slightest degree, approach those of the Lemurs. And if, instead of putting Hapale out of its natural place, as Professor Bischoff most unaccountably does, we write the series of animals he has chosen to mention as follows: Homo, Pithecus, Troglodytes, Hylobates, Semnopithecus, Cynocephalus, Cercopithecus, Macacus, Cebus, Callithrix, Hapale, Lemur, Stenops, I venture to reaffirm that the great break in this series lies between Hapale and Lemur, and that this break is considerably greater than that between any other two terms of that series. Professor Bischoff ignores the fact that long before he wrote, Gratiolet had suggested the separation of the Lemurs from the other Primates on the very ground of the difference in their cerebral characters; and that Professor Flower had made the following observations in the course of his description of the brain of the Javan Loris: (75. ‘Transactions of the Zoological Society,’ vol. v. 1862.)

“And it is especially remarkable that, in the development of the posterior lobes, there is no approximation to the Lemurine, short hemisphered brain, in those monkeys which are commonly supposed to approach this family in other respects, viz. the lower members of the Platyrrhine group.”

So far as the structure of the adult brain is concerned, then, the very considerable additions to our knowledge, which have been made by the researches of so many investigators, during the past ten years, fully justify the statement which I made in 1863. But it has been said, that, admitting the similarity between the adult brains of man and apes, they are nevertheless, in reality, widely different, because they exhibit fundamental differences in the mode of their development. No one would be more ready than I to admit the force of this argument, if such fundamental differences of development really exist. But I deny that they do exist. On the contrary, there is a fundamental agreement in the development of the brain in men and apes.

Gratiolet originated the statement that there is a fundamental difference in the development of the brains of apes and that of man—consisting in this; that, in the apes, the sulci which first make their appearance are situated on the posterior region of the cerebral hemispheres, while, in the human foetus, the sulci first become visible on the frontal lobes. (76. Chez tous les singes, les plis postérieurs se developpent les premiers; les plis antérieurs se developpent plus tard, aussi la vertèbre occipitale et la parietale sont-elles relativement tres-grandes chez le foetus. L’Homme présente une exception remarquable quant a l’époque de l’apparition des plis frontaux, qui sont les premiers indiqués; mais le développement general du lobe frontal, envisagé seulement par rapport a son volume, suit les mêmes lois que dans les singes: Gratiolet, ‘Mémoire sur les plis cérèbres de l’Homme et des Primateaux,’ p. 39, Tab. iv, fig. 3.)

This general statement is based upon two observations, the one of a Gibbon almost ready to be born, in which the posterior gyri were “well developed,” while those of the frontal lobes were “hardly indicated” (77. Gratiolet’s words are (loc. cit. p. 39): “Dans le foetus dont il s’agit les plis cérébraux posterieurs sont bien developpés, tandis que les plis du lobe frontal sont a peine indiqués.” The figure, however (Pl. iv, fig. 3), shews the fissure of Rolando, and one of the frontal sulci plainly enough. Nevertheless, M. Alix, in his ‘Notice sur les travaux anthropologiques de Gratiolet’ (‘Mem. de la Societé d’Anthropologie de Paris,’ 1868, page 32), writes thus: “Gratiolet a eu entre les mains le cerveau d’un foetus de Gibbon, singe eminemment supérieur, et tellement rapproché de l’orang, que des naturalistes tres-compétents l’ont rangé parmi les anthropoides. M. Huxley, par exemple, n’hesite pas sur ce point. Eh bien, c’est sur le cerveau d’un foetus de Gibbon que Gratiolet a vu LES CIRCONVOLUTIONS DU LOBE TEMPORO-SPHENOIDAL DÉJÀ DEVELOPPÉES LORSQU’IL N’EXISTENT PAS ENCORE DE PLIS SUR LE LOBE FRONTAL. Il etait donc bien autorisé a dire que, chez l’homme les circonvolutions apparaissent d’a en w, tandis que chez les singes elles se developpent d’w en a.”), and the other of a human foetus at the 22nd or 23rd week of uterogestation, in which Gratiolet notes that the insula was uncovered, but that nevertheless “des incisures sement de lobe anterieur, une scissure peu profonde indique la separation du lobe occipital, tres-reduit, d’ailleurs dès cette époque. Le reste de la surface cérébrale est encore absolument lisse.”

Three views of this brain are given in Plate II, figs. 1, 2, 3, of the work cited, shewing the upper, lateral and inferior views of the hemispheres, but not the inner view. It is worthy of note that the figure by no means bears out Gratiolet’s description, inasmuch as the fissure (antero-temporal) on the posterior half of the face of the hemisphere is more marked than any of those vaguely indicated in the anterior half. If the figure is correct, it in no way justifies Gratiolet’s conclusion: “Il y a donc entre ces cerveaux [those of a Callithrix and of a Gibbon] et celui du foetus humain une différence fondamental. Chez celui-ci, longtemps avant que les plis temporaux apparaissent, les plis frontaux, ESSAYENT d’exister.”

Since Gratiolet’s time, however, the development of the gyri and sulci of the brain has been made the subject of renewed investigation by Schmidt, Bischoff, Pansch (78. ‘Ueber die typische Anordnung der Furchen und Windungen auf den Grosshirn-Hemisphären des Menschen und der Affen,’ ‘Archiv für Anthropologie,’ iii. 1868.), and more particularly by Ecker (79. ‘Zur Entwicklungs Geschichte der Furchen und Windungen der Grosshirn-Hemisphären im Foetus des Menschen,’ ‘Archiv für Anthropologie,’ iii. 1868.), whose work is not only the latest, but by far the most complete, memoir on the subject.

The final results of their inquiries may be summed up as follows:— 1. In the human foetus, the sylvian fissure is formed in the course of the third month of uterogestation. In this, and in the fourth month, the cerebral hemispheres are smooth and rounded (with the exception of the sylvian depression), and they project backwards far beyond the cerebellum.

2. The sulci, properly so called, begin to appear in the interval between the end of the fourth and the beginning of the sixth month of foetal life, but Ecker is careful to point out that, not only the time, but the order, of their appearance is subject to considerable individual variation. In no case, however, are either the frontal or the temporal sulci the earliest.

The first which appears, in fact, lies on the inner face of the hemisphere (whence doubtless Gratiolet, who does not seem to have examined that face in his foetus, overlooked it), and is either the internal perpendicular (occipito-parietal), or the calcarine sulcus, these two being close together and eventually running into one another. As a rule the occipito-parietal is the earlier of the two.

3. At the latter part of this period, another sulcus, the “posterio-parietal,” or “Fissure of Rolando” is developed, and it is followed, in the course of the sixth month, by the other principal sulci of the frontal, parietal, temporal and occipital lobes. There is, however, no clear evidence that one of these constantly appears before the other; and it is remarkable that, in the brain at the period described and figured by Ecker (loc. cit. pp. 212-213, Taf. II, figs. 1, 2, 3, 4), the antero-temporal sulcus (scissure parallele) so characteristic of the ape’s brain, is as well, if not better developed than the fissure of Rolando, and is much more marked than the proper frontal sulci.

Taking the facts as they now stand, it appears to me that the order of the appearance of the sulci and gyri in the foetal human brain is in perfect harmony with the general doctrine of evolution, and with the view that man has been evolved from some ape-like form; though there can be no doubt that form was, in many respects, different from any member of the Primates now living.

Von Baer taught us, half a century ago, that, in the course of their development, allied animals put on at first, the characters of the greater groups to which they belong, and, by degrees, assume those which restrict them within the limits of their family, genus, and species; and he proved, at the same time, that no developmental stage of a higher animal is precisely similar to the adult condition of any lower animal. It is quite correct to say that a frog passes through the condition of a fish, inasmuch as at one period of its life the tadpole has all the characters of a fish, and if it went no further, would have to be grouped among fishes. But it is equally true that a tadpole is very different from any known fish.

In like manner, the brain of a human foetus, at the fifth month, may correctly be said to be, not only the brain of an ape, but that of an Arctopithecine or marmoset-like ape; for its hemispheres, with their great posterior lobster, and with no sulci but the sylvian and the calcarine, present the characteristics found only in the group of the Arctopithecine Primates. But it is equally true, as Gratiolet remarks, that, in its widely open sylvian fissure, it differs from the brain of any actual marmoset. No doubt it would be much more similar to the brain of an advanced foetus of a marmoset. But we know nothing whatever of the development of the brain in the marmosets. In the Platyrrhini proper, the only observation with which I am acquainted is due to Pansch, who found in the brain of a foetal Cebus Apella, in addition to the sylvian fissure and the deep calcarine fissure, only a very shallow antero-temporal fissure (scissure parallele of Gratiolet).

Now this fact, taken together with the circumstance that the antero-temporal sulcus is present in such Platyrrhini as the Saimiri, which present mere traces of sulci on the anterior half of the exterior of the cerebral hemispheres, or none at all, undoubtedly, so far as it goes, affords fair evidence in favour of Gratiolet’s hypothesis, that the posterior sulci appear before the anterior, in the brains of the Platyrrhini. But, it by no means follows, that the rule which may hold good for the Platyrrhini extends to the Catarrhini. We have no information whatever respecting the development of the brain in the Cynomorpha; and, as regards the Anthropomorpha, nothing but the account of the brain of the Gibbon, near birth, already referred to. At the present moment there is not a shadow of evidence to shew that the sulci of a chimpanzee’s, or orang’s, brain do not appear in the same order as a man’s.

Gratiolet opens his preface with the aphorism: “Il est dangereux dans les sciences de conclure trop vite.” I fear he must have forgotten this sound maxim by the time he had reached the discussion of the differences between men and apes, in the body of his work. No doubt, the excellent author of one of the most remarkable contributions to the just understanding of the mammalian brain which has ever been made, would have been the first to admit the insufficiency of his data had he lived to profit by the advance of inquiry. The misfortune is that his conclusions have been employed by persons incompetent to appreciate their foundation, as arguments in favour of obscurantism. (80. For example, M. l’Abbe Lecomte in his terrible pamphlet, ‘Le Darwinisme et l’origine de l’Homme,’ 1873.)

But it is important to remark that, whether Gratiolet was right or wrong in his hypothesis respecting the relative order of appearance of the temporal and frontal sulci, the fact remains; that before either temporal or frontal sulci, appear, the foetal brain of man presents characters which are found only in the lowest group of the Primates (leaving out the Lemurs); and that this is exactly what we should expect to be the case, if man has resulted from the gradual modification of the same form as that from which the other Primates have sprung.

PART II.
SEXUAL SELECTION.

CHAPTER VIII.
PRINCIPLES OF SEXUAL SELECTION.

Secondary sexual characters—Sexual selection—Manner of action—Excess of males—Polygamy—The male alone generally modified through sexual selection—Eagerness of the male—Variability of the male—Choice exerted by the female—Sexual compared with natural selection—Inheritance, at corresponding periods of life, at corresponding seasons of the year, and as limited by sex—Relations between the several forms of inheritance—Causes why one sex and the young are not modified through sexual selection—Supplement on the proportional numbers of the two sexes throughout the animal kingdom—The proportion of the sexes in relation to natural selection.

With animals which have their sexes separated, the males necessarily differ from the females in their organs of reproduction; and these are the primary sexual characters. But the sexes often differ in what Hunter has called secondary sexual characters, which are not directly connected with the act of reproduction; for instance, the male possesses certain organs of sense or locomotion, of which the female is quite destitute, or has them more highly-developed, in order that he may readily find or reach her; or again the male has special organs of prehension for holding her securely. These latter organs, of infinitely diversified kinds, graduate into those which are commonly ranked as primary, and in some cases can hardly be distinguished from them; we see instances of this in the complex appendages at the apex of the abdomen in male insects. Unless indeed we confine the term “primary” to the reproductive glands, it is scarcely possible to decide which ought to be called primary and which secondary.

The female often differs from the male in having organs for the nourishment or protection of her young, such as the mammary glands of mammals, and the abdominal sacks of the marsupials. In some few cases also the male possesses similar organs, which are wanting in the female, such as the receptacles for the ova in certain male fishes, and those temporarily developed in certain male frogs. The females of most bees are provided with a special apparatus for collecting and carrying pollen, and their ovipositor is modified into a sting for the defence of the larvae and the community. Many similar cases could be given, but they do not here concern us. There are, however, other sexual differences quite unconnected with the primary reproductive organs, and it is with these that we are more especially concerned—such as the greater size, strength, and pugnacity of the male, his weapons of offence or means of defence against rivals, his gaudy colouring and various ornaments, his power of song, and other such characters.

Besides the primary and secondary sexual differences, such as the foregoing, the males and females of some animals differ in structures related to different habits of life, and not at all, or only indirectly, to the reproductive functions. Thus the females of certain flies (Culicidae and Tabanidae) are blood-suckers, whilst the males, living on flowers, have mouths destitute of mandibles. (1. Westwood, ‘Modern Classification of Insects,’ vol. ii. 1840, p. 541. For the statement about Tanais, mentioned below, I am indebted to Fritz Muller.) The males of certain moths and of some crustaceans (e.g. Tanais) have imperfect, closed mouths, and cannot feed. The complemental males of certain Cirripedes live like epiphytic plants either on the female or the hermaphrodite form, and are destitute of a mouth and of prehensile limbs. In these cases it is the male which has been modified, and has lost certain important organs, which the females possess. In other cases it is the female which has lost such parts; for instance, the female glow-worm is destitute of wings, as also are many female moths, some of which never leave their cocoons. Many female parasitic crustaceans have lost their natatory legs. In some weevil-beetles (Curculionidae) there is a great difference between the male and female in the length of the rostrum or snout (2. Kirby and Spence, ‘Introduction to Entomology,’ vol. iii. 1826, p. 309.); but the meaning of this and of many analogous differences, is not at all understood. Differences of structure between the two sexes in relation to different habits of life are generally confined to the lower animals; but with some few birds the beak of the male differs from that of the female. In the Huia of New Zealand the difference is wonderfully great, and we hear from Dr. Buller (3. ‘Birds of New Zealand,’ 1872, p. 66.) that the male uses his strong beak in chiselling the larvae of insects out of decayed wood, whilst the female probes the softer parts with her far longer, much curved and pliant beak: and thus they mutually aid each other. In most cases, differences of structure between the sexes are more or less directly connected with the propagation of the species: thus a female, which has to nourish a multitude of ova, requires more food than the male, and consequently requires special means for procuring it. A male animal, which lives for a very short time, might lose its organs for procuring food through disuse, without detriment; but he would retain his locomotive organs in a perfect state, so that he might reach the female. The female, on the other hand, might safely lose her organs for flying, swimming, or walking, if she gradually acquired habits which rendered such powers useless.

We are, however, here concerned only with sexual selection. This depends on the advantage which certain individuals have over others of the same sex and species solely in respect of reproduction. When, as in the cases above mentioned, the two sexes differ in structure in relation to different habits of life, they have no doubt been modified through natural selection, and by inheritance limited to one and the same sex. So again the primary sexual organs, and those for nourishing or protecting the young, come under the same influence; for those individuals which generated or nourished their offspring best, would leave, ceteris paribus, the greatest number to inherit their superiority; whilst those which generated or nourished their offspring badly, would leave but few to inherit their weaker powers. As the male has to find the female, he requires organs of sense and locomotion, but if these organs are necessary for the other purposes of life, as is generally the case, they will have been developed through natural selection. When the male has found the female, he sometimes absolutely requires prehensile organs to hold her; thus Dr. Wallace informs me that the males of certain moths cannot unite with the females if their tarsi or feet are broken. The males of many oceanic crustaceans, when adult, have their legs and antennae modified in an extraordinary manner for the prehension of the female; hence we may suspect that it is because these animals are washed about by the waves of the open sea, that they require these organs in order to propagate their kind, and if so, their development has been the result of ordinary or natural selection. Some animals extremely low in the scale have been modified for this same purpose; thus the males of certain parasitic worms, when fully grown, have the lower surface of the terminal part of their bodies roughened like a rasp, and with this they coil round and permanently hold the females. (4. M. Perrier advances this case (‘Revue Scientifique,’ Feb. 1, 1873, p. 865) as one fatal to the belief in sexual election, inasmuch as he supposes that I attribute all the differences between the sexes to sexual selection. This distinguished naturalist, therefore, like so many other Frenchmen, has not taken the trouble to understand even the first principles of sexual selection. An English naturalist insists that the claspers of certain male animals could not have been developed through the choice of the female! Had I not met with this remark, I should not have thought it possible for any one to have read this chapter and to have imagined that I maintain that the choice of the female had anything to do with the development of the prehensile organs in the male.)

When the two sexes follow exactly the same habits of life, and the male has the sensory or locomotive organs more highly developed than those of the female, it may be that the perfection of these is indispensable to the male for finding the female; but in the vast majority of cases, they serve only to give one male an advantage over another, for with sufficient time, the less well-endowed males would succeed in pairing with the females; and judging from the structure of the female, they would be in all other respects equally well adapted for their ordinary habits of life. Since in such cases the males have acquired their present structure, not from being better fitted to survive in the struggle for existence, but from having gained an advantage over other males, and from having transmitted this advantage to their male offspring alone, sexual selection must here have come into action. It was the importance of this distinction which led me to designate this form of selection as Sexual Selection. So again, if the chief service rendered to the male by his prehensile organs is to prevent the escape of the female before the arrival of other males, or when assaulted by them, these organs will have been perfected through sexual selection, that is by the advantage acquired by certain individuals over their rivals. But in most cases of this kind it is impossible to distinguish between the effects of natural and sexual selection. Whole chapters could be filled with details on the differences between the sexes in their sensory, locomotive, and prehensile organs. As, however, these structures are not more interesting than others adapted for the ordinary purposes of life I shall pass them over almost entirely, giving only a few instances under each class.

There are many other structures and instincts which must have been developed through sexual selection—such as the weapons of offence and the means of defence of the males for fighting with and driving away their rivals—their courage and pugnacity—their various ornaments—their contrivances for producing vocal or instrumental music—and their glands for emitting odours, most of these latter structures serving only to allure or excite the female. It is clear that these characters are the result of sexual and not of ordinary selection, since unarmed, unornamented, or unattractive males would succeed equally well in the battle for life and in leaving a numerous progeny, but for the presence of better endowed males. We may infer that this would be the case, because the females, which are unarmed and unornamented, are able to survive and procreate their kind. Secondary sexual characters of the kind just referred to, will be fully discussed in the following chapters, as being in many respects interesting, but especially as depending on the will, choice, and rivalry of the individuals of either sex. When we behold two males fighting for the possession of the female, or several male birds displaying their gorgeous plumage, and performing strange antics before an assembled body of females, we cannot doubt that, though led by instinct, they know what they are about, and consciously exert their mental and bodily powers.

Just as man can improve the breeds of his game-cocks by the selection of those birds which are victorious in the cockpit, so it appears that the strongest and most vigorous males, or those provided with the best weapons, have prevailed under nature, and have led to the improvement of the natural breed or species. A slight degree of variability leading to some advantage, however slight, in reiterated deadly contests would suffice for the work of sexual selection; and it is certain that secondary sexual characters are eminently variable. Just as man can give beauty, according to his standard of taste, to his male poultry, or more strictly can modify the beauty originally acquired by the parent species, can give to the Sebright bantam a new and elegant plumage, an erect and peculiar carriage—so it appears that female birds in a state of nature, have by a long selection of the more attractive males, added to their beauty or other attractive qualities. No doubt this implies powers of discrimination and taste on the part of the female which will at first appear extremely improbable; but by the facts to be adduced hereafter, I hope to be able to shew that the females actually have these powers. When, however, it is said that the lower animals have a sense of beauty, it must not be supposed that such sense is comparable with that of a cultivated man, with his multiform and complex associated ideas. A more just comparison would be between the taste for the beautiful in animals, and that in the lowest savages, who admire and deck themselves with any brilliant, glittering, or curious object.

From our ignorance on several points, the precise manner in which sexual selection acts is somewhat uncertain. Nevertheless if those naturalists who already believe in the mutability of species, will read the following chapters, they will, I think, agree with me, that sexual selection has played an important part in the history of the organic world. It is certain that amongst almost all animals there is a struggle between the males for the possession of the female. This fact is so notorious that it would be superfluous to give instances. Hence the females have the opportunity of selecting one out of several males, on the supposition that their mental capacity suffices for the exertion of a choice. In many cases special circumstances tend to make the struggle between the males particularly severe. Thus the males of our migratory birds generally arrive at their places of breeding before the females, so that many males are ready to contend for each female. I am informed by Mr. Jenner Weir, that the bird-catchers assert that this is invariably the case with the nightingale and blackcap, and with respect to the latter he can himself confirm the statement.

Mr. Swaysland of Brighton has been in the habit, during the last forty years, of catching our migratory birds on their first arrival, and he has never known the females of any species to arrive before their males. During one spring he shot thirty-nine males of Ray’s wagtail (Budytes Raii) before he saw a single female. Mr. Gould has ascertained by the dissection of those snipes which arrive the first in this country, that the males come before the females. And the like holds good with most of the migratory birds of the United States. (5. J.A. Allen, on the ‘Mammals and Winter Birds of Florida,’ Bulletin of Comparative Zoology, Harvard College, p. 268.) The majority of the male salmon in our rivers, on coming up from the sea, are ready to breed before the females. So it appears to be with frogs and toads. Throughout the great class of insects the males almost always are the first to emerge from the pupal state, so that they generally abound for a time before any females can be seen. (6. Even with those plants in which the sexes are separate, the male flowers are generally mature before the female. As first shewn by C.K. Sprengel, many hermaphrodite plants are dichogamous; that is, their male and female organs are not ready at the same time, so that they cannot be self-fertilised. Now in such flowers, the pollen is in general matured before the stigma, though there are exceptional cases in which the female organs are beforehand.) The cause of this difference between the males and females in their periods of arrival and maturity is sufficiently obvious. Those males which annually first migrated into any country, or which in the spring were first ready to breed, or were the most eager, would leave the largest number of offspring; and these would tend to inherit similar instincts and constitutions. It must be borne in mind that it would have been impossible to change very materially the time of sexual maturity in the females, without at the same time interfering with the period of the production of the young—a period which must be determined by the seasons of the year. On the whole there can be no doubt that with almost all animals, in which the sexes are separate, there is a constantly recurrent struggle between the males for the possession of the females.

Our difficulty in regard to sexual selection lies in understanding how it is that the males which conquer other males, or those which prove the most attractive to the females, leave a greater number of offspring to inherit their superiority than their beaten and less attractive rivals. Unless this result does follow, the characters which give to certain males an advantage over others, could not be perfected and augmented through sexual selection. When the sexes exist in exactly equal numbers, the worst-endowed males will (except where polygamy prevails), ultimately find females, and leave as many offspring, as well fitted for their general habits of life, as the best-endowed males. From various facts and considerations, I formerly inferred that with most animals, in which secondary sexual characters are well developed, the males considerably exceeded the females in number; but this is not by any means always true. If the males were to the females as two to one, or as three to two, or even in a somewhat lower ratio, the whole affair would be simple; for the better-armed or more attractive males would leave the largest number of offspring. But after investigating, as far as possible, the numerical proportion of the sexes, I do not believe that any great inequality in number commonly exists. In most cases sexual selection appears to have been effective in the following manner.

Let us take any species, a bird for instance, and divide the females inhabiting a district into two equal bodies, the one consisting of the more vigorous and better-nourished individuals, and the other of the less vigorous and healthy. The former, there can be little doubt, would be ready to breed in the spring before the others; and this is the opinion of Mr. Jenner Weir, who has carefully attended to the habits of birds during many years. There can also be no doubt that the most vigorous, best-nourished and earliest breeders would on an average succeed in rearing the largest number of fine offspring. (7. Here is excellent evidence on the character of the offspring from an experienced ornithologist. Mr. J.A. Allen, in speaking (‘Mammals and Winter Birds of E. Florida,’ p. 229) of the later broods, after the accidental destruction of the first, says, that these “are found to be smaller and paler-coloured than those hatched earlier in the season. In cases where several broods are reared each year, as a general rule the birds of the earlier broods seem in all respects the most perfect and vigorous.”) The males, as we have seen, are generally ready to breed before the females; the strongest, and with some species the best armed of the males, drive away the weaker; and the former would then unite with the more vigorous and better-nourished females, because they are the first to breed. (8. Hermann Müller has come to this same conclusion with respect to those female bees which are the first to emerge from the pupa each year. See his remarkable essay, ‘Anwendung der Darwin’schen Lehre auf Bienen,’ ‘Verh. d. V. Jahrg.’ xxix. p. 45.) Such vigorous pairs would surely rear a larger number of offspring than the retarded females, which would be compelled to unite with the conquered and less powerful males, supposing the sexes to be numerically equal; and this is all that is wanted to add, in the course of successive generations, to the size, strength and courage of the males, or to improve their weapons.

But in very many cases the males which conquer their rivals, do not obtain possession of the females, independently of the choice of the latter. The courtship of animals is by no means so simple and short an affair as might be thought. The females are most excited by, or prefer pairing with, the more ornamented males, or those which are the best songsters, or play the best antics; but it is obviously probable that they would at the same time prefer the more vigorous and lively males, and this has in some cases been confirmed by actual observation. (9. With respect to poultry, I have received information, hereafter to be given, to this effect. Even with birds, such as pigeons, which pair for life, the female, as I hear from Mr. Jenner Weir, will desert her mate if he is injured or grows weak.) Thus the more vigorous females, which are the first to breed, will have the choice of many males; and though they may not always select the strongest or best armed, they will select those which are vigorous and well armed, and in other respects the most attractive. Both sexes, therefore, of such early pairs would as above explained, have an advantage over others in rearing offspring; and this apparently has sufficed during a long course of generations to add not only to the strength and fighting powers of the males, but likewise to their various ornaments or other attractions.

In the converse and much rarer case of the males selecting particular females, it is plain that those which were the most vigorous and had conquered others, would have the freest choice; and it is almost certain that they would select vigorous as well as attractive females. Such pairs would have an advantage in rearing offspring, more especially if the male had the power to defend the female during the pairing-season as occurs with some of the higher animals, or aided her in providing for the young. The same principles would apply if each sex preferred and selected certain individuals of the opposite sex; supposing that they selected not only the more attractive, but likewise the more vigorous individuals.