The Descent of Man, and Selection in Relation to Sex

We here see that in the interval of forty years, between 1832 and 1872, the population has decreased no less than sixty-eight per cent.! This has been attributed by most writers to the profligacy of the women, to former bloody wars, and to the severe labour imposed on conquered tribes and to newly introduced diseases, which have been on several occasions extremely destructive. No doubt these and other such causes have been highly efficient, and may account for the extraordinary rate of decrease between the years 1832 and 1836; but the most potent of all the causes seems to be lessened fertility. According to Dr. Ruschenberger of the U.S. Navy, who visited these islands between 1835 and 1837, in one district of Hawaii, only twenty-five men out of 1134, and in another district only ten out of 637, had a family with as many as three children. Of eighty married women, only thirty-nine had ever borne children; and “the official report gives an average of half a child to each married couple in the whole island.” This is almost exactly the same average as with the Tasmanians at Oyster Cove. Jarves, who published his History in 1843, says that “families who have three children are freed from all taxes; those having more, are rewarded by gifts of land and other encouragements.” This unparalleled enactment by the government well shews how infertile the race had become. The Rev. A. Bishop stated in the Hawaiian ‘Spectator’ in 1839, that a large proportion of the children die at early ages, and Bishop Staley informs me that this is still the case, just as in New Zealand. This has been attributed to the neglect of the children by the women, but it is probably in large part due to innate weakness of constitution in the children, in relation to the lessened fertility of their parents. There is, moreover, a further resemblance to the case of New Zealand, in the fact that there is a large excess of male over female births: the census of 1872 gives 31,650 males to 25,247 females of all ages, that is 125.36 males for every 100 females; whereas in all civilised countries the females exceed the males. No doubt the profligacy of the women may in part account for their small fertility; but their changed habits of life is a much more probable cause, and which will at the same time account for the increased mortality, especially of the children. The islands were visited by Cook in 1779, Vancouver in 1794, and often subsequently by whalers. In 1819 missionaries arrived, and found that idolatry had been already abolished, and other changes effected by the king. After this period there was a rapid change in almost all the habits of life of the natives, and they soon became “the most civilised of the Pacific Islanders.” One of my informants, Mr. Coan, who was born on the islands, remarks that the natives have undergone a greater change in their habits of life in the course of fifty years than Englishmen during a thousand years. From information received from Bishop Staley, it does not appear that the poorer classes have ever much changed their diet, although many new kinds of fruit have been introduced, and the sugar-cane is in universal use. Owing, however, to their passion for imitating Europeans, they altered their manner of dressing at an early period, and the use of alcoholic drinks became very general. Although these changes appear inconsiderable, I can well believe, from what is known with respect to animals, that they might suffice to lessen the fertility of the natives. (43. The foregoing statements are taken chiefly from the following works: Jarves’ ‘History of the Hawaiian Islands,’ 1843, pp. 400-407. Cheever, ‘Life in the Sandwich Islands,’ 1851, p. 277. Ruschenberger is quoted by Bonwick, ‘Last of the Tasmanians,’ 1870, p. 378. Bishop is quoted by Sir E. Belcher, ‘Voyage Round the World,’ 1843, vol. i. p. 272. I owe the census of the several years to the kindness of Mr. Coan, at the request of Dr. Youmans of New York; and in most cases I have compared the Youmans figures with those given in several of the above-named works. I have omitted the census for 1850, as I have seen two widely different numbers given.)

Lastly, Mr. Macnamara states (44. ‘The Indian Medical Gazette,’ Nov. 1, 1871, p. 240.) that the low and degraded inhabitants of the Andaman Islands, on the eastern side of the Gulf of Bengal, are “eminently susceptible to any change of climate: in fact, take them away from their island homes, and they are almost certain to die, and that independently of diet or extraneous influences.” He further states that the inhabitants of the Valley of Nepal, which is extremely hot in summer, and also the various hill-tribes of India, suffer from dysentery and fever when on the plains; and they die if they attempt to pass the whole year there.

We thus see that many of the wilder races of man are apt to suffer much in health when subjected to changed conditions or habits of life, and not exclusively from being transported to a new climate. Mere alterations in habits, which do not appear injurious in themselves, seem to have this same effect; and in several cases the children are particularly liable to suffer. It has often been said, as Mr. Macnamara remarks, that man can resist with impunity the greatest diversities of climate and other changes; but this is true only of the civilised races. Man in his wild condition seems to be in this respect almost as susceptible as his nearest allies, the anthropoid apes, which have never yet survived long, when removed from their native country.

Lessened fertility from changed conditions, as in the case of the Tasmanians, Maories, Sandwich Islanders, and apparently the Australians, is still more interesting than their liability to ill-health and death; for even a slight degree of infertility, combined with those other causes which tend to check the increase of every population, would sooner or later lead to extinction. The diminution of fertility may be explained in some cases by the profligacy of the women (as until lately with the Tahitians), but Mr. Fenton has shewn that this explanation by no means suffices with the New Zealanders, nor does it with the Tasmanians.

In the paper above quoted, Mr. Macnamara gives reasons for believing that the inhabitants of districts subject to malaria are apt to be sterile; but this cannot apply in several of the above cases. Some writers have suggested that the aborigines of islands have suffered in fertility and health from long continued inter-breeding; but in the above cases infertility has coincided too closely with the arrival of Europeans for us to admit this explanation. Nor have we at present any reason to believe that man is highly sensitive to the evil effects of inter-breeding, especially in areas so large as New Zealand, and the Sandwich archipelago with its diversified stations. On the contrary, it is known that the present inhabitants of Norfolk Island are nearly all cousins or near relations, as are the Todas in India, and the inhabitants of some of the Western Islands of Scotland; and yet they seem not to have suffered in fertility. (45. On the close relationship of the Norfolk Islanders, Sir W. Denison, ‘Varieties of Vice-Regal Life,’ vol. i. 1870, p. 410. For the Todas, see Col. Marshall’s work 1873, p. 110. For the Western Islands of Scotland, Dr. Mitchell, ‘Edinburgh Medical Journal,’ March to June, 1865.)

A much more probable view is suggested by the analogy of the lower animals. The reproductive system can be shewn to be susceptible to an extraordinary degree (though why we know not) to changed conditions of life; and this susceptibility leads both to beneficial and to evil results. A large collection of facts on this subject is given in chap. xviii. of vol. ii. of my ‘Variation of Animals and Plants under Domestication.’ I can here give only the briefest abstract; and every one interested in the subject may consult the above work. Very slight changes increase the health, vigour, and fertility of most or all organic beings, whilst other changes are known to render a large number of animals sterile. One of the most familiar cases, is that of tamed elephants not breeding in India; though they often breed in Ava, where the females are allowed to roam about the forests to some extent, and are thus placed under more natural conditions. The case of various American monkeys, both sexes of which have been kept for many years together in their own countries, and yet have very rarely or never bred, is a more apposite instance, because of their relationship to man. It is remarkable how slight a change in the conditions often induces sterility in a wild animal when captured; and this is the more strange as all our domesticated animals have become more fertile than they were in a state of nature; and some of them can resist the most unnatural conditions with undiminished fertility. (46. For the evidence on this head, see ‘Variation of Animals,’ etc., vol. ii. p. 111.) Certain groups of animals are much more liable than others to be affected by captivity; and generally all the species of the same group are affected in the same manner. But sometimes a single species in a group is rendered sterile, whilst the others are not so; on the other hand, a single species may retain its fertility whilst most of the others fail to breed. The males and females of some species when confined, or when allowed to live almost, but not quite free, in their native country, never unite; others thus circumstanced frequently unite but never produce offspring; others again produce some offspring, but fewer than in a state of nature; and as bearing on the above cases of man, it is important to remark that the young are apt to be weak and sickly, or malformed, and to perish at an early age.

Seeing how general is this law of the susceptibility of the reproductive system to changed conditions of life, and that it holds good with our nearest allies, the Quadrumana, I can hardly doubt that it applies to man in his primeval state. Hence if savages of any race are induced suddenly to change their habits of life, they become more or less sterile, and their young offspring suffer in health, in the same manner and from the same cause, as do the elephant and hunting-leopard in India, many monkeys in America, and a host of animals of all kinds, on removal from their natural conditions.

We can see why it is that aborigines, who have long inhabited islands, and who must have been long exposed to nearly uniform conditions, should be specially affected by any change in their habits, as seems to be the case. Civilised races can certainly resist changes of all kinds far better than savages; and in this respect they resemble domesticated animals, for though the latter sometimes suffer in health (for instance European dogs in India), yet they are rarely rendered sterile, though a few such instances have been recorded. (47. ‘Variation of Animals,’ etc., vol. ii. p. 16.) The immunity of civilised races and domesticated animals is probably due to their having been subjected to a greater extent, and therefore having grown somewhat more accustomed, to diversified or varying conditions, than the majority of wild animals; and to their having formerly immigrated or been carried from country to country, and to different families or sub-races having inter-crossed. It appears that a cross with civilised races at once gives to an aboriginal race an immunity from the evil consequences of changed conditions. Thus the crossed offspring from the Tahitians and English, when settled in Pitcairn Island, increased so rapidly that the island was soon overstocked; and in June 1856 they were removed to Norfolk Island. They then consisted of 60 married persons and 134 children, making a total of 194. Here they likewise increased so rapidly, that although sixteen of them returned to Pitcairn Island in 1859, they numbered in January 1868, 300 souls; the males and females being in exactly equal numbers. What a contrast does this case present with that of the Tasmanians; the Norfolk Islanders INCREASED in only twelve and a half years from 194 to 300; whereas the Tasmanians DECREASED during fifteen years from 120 to 46, of which latter number only ten were children. (48. These details are taken from ‘The Mutineers of the “Bounty,”’ by Lady Belcher, 1870; and from ‘Pitcairn Island,’ ordered to be printed by the House of Commons, May 29, 1863. The following statements about the Sandwich Islanders are from the ‘Honolulu Gazette,’ and from Mr. Coan.)

So again in the interval between the census of 1866 and 1872 the natives of full blood in the Sandwich Islands decreased by 8081, whilst the half-castes, who are believed to be healthier, increased by 847; but I do not know whether the latter number includes the offspring from the half-castes, or only the half-castes of the first generation.

The cases which I have here given all relate to aborigines, who have been subjected to new conditions as the result of the immigration of civilised men. But sterility and ill-health would probably follow, if savages were compelled by any cause, such as the inroad of a conquering tribe, to desert their homes and to change their habits. It is an interesting circumstance that the chief check to wild animals becoming domesticated, which implies the power of their breeding freely when first captured, and one chief check to wild men, when brought into contact with civilisation, surviving to form a civilised race, is the same, namely, sterility from changed conditions of life.

Finally, although the gradual decrease and ultimate extinction of the races of man is a highly complex problem, depending on many causes which differ in different places and at different times; it is the same 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 troups of the Spanish horse. The New Zealander seems conscious of this parallelism, for he compares his future fate with that of the native rat now almost exterminated by the European rat. Though the difficulty is great to our imagination, and really great, if we wish to ascertain the precise causes and their manner of action, it 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 checked in various ways; so that if any new check, even a slight one, be superadded, the race will surely decrease in number; and decreasing numbers will sooner or later lead to extinction; the end, in most cases, being promptly determined by the inroads of conquering tribes.

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.