The Descent of Man

Charles Darwin

Chapter 21 - General Summary and Conclusion


A BRIEF summary will be sufficient to recall to the reader's mind the more salient points in this work. Many of the views which have been advanced are highly speculative, and some no doubt will prove erroneous; but I have in every case given the reasons which have led me to one view rather than to another. It seemed worth while to try how far the principle of evolution would throw light on some of the more complex problems in the natural history of man. False facts are highly injurious to the progress of science, for they often endure long; but false views, if supported by some evidence, do little harm, for every one takes a salutary pleasure in proving their falseness: and when this is done, one path towards error is closed and the road to truth is often at the same time opened.

The main conclusion here arrived at, and now held by many naturalists who are well competent to form a sound judgment is that man is descended from some less highly organised form. The grounds upon which this conclusion rests will never be shaken, for the close similarity between man and the lower animals in embryonic development, as well as in innumerable points of structure and constitution, both of high and of the most trifling importance,- the rudiments which he retains, and the abnormal reversions to which he is occasionally liable,- are facts which cannot be disputed. They have long been known, but until recently they told us nothing with respect to the origin of man. Now when viewed by the light of our knowledge of the whole organic world, their meaning is unmistakable. The great principle of evolution stands up clear and firm, when these groups or facts are considered in connection with others, such as the mutual affinities of the members of the same group, their geographical distribution in past and present times, and their geological succession. It is incredible that all these facts should speak falsely. He who is not content to look, like a savage, at the phenomena of nature as disconnected, cannot any longer believe that man is the work of a separate act of creation. He will be forced to admit that the close resemblance of the embryo of man to that, for instance, of a dog- the construction of his skull, limbs and whole frame on the same plan with that of other mammals, independently of the uses to which the parts may be put- the occasional re-appearance of various structures, for instance of several muscles, which man does not normally possess, but which are common to the Quadrumana- and a crowd of analogous facts- all point in the plainest manner to the conclusion that man is the co-descendant with other mammals of a common progenitor.

We have seen that man incessantly presents individual differences in all parts of his body and in his mental faculties. These differences or variations seem to be induced by the same general causes, and to obey the same laws as with the lower animals. In both cases similar laws of inheritance prevail. Man tends to increase at a greater rate than his means of subsistence; consequently he is occasionally subjected to a severe struggle for existence, and natural selection will have effected whatever lies within its scope. A succession of strongly-marked variations of a similar nature is by no means requisite; slight fluctuating differences in the individual suffice for the work of natural selection; not that we have any reason to suppose that in the same species, all parts of the organisation tend to vary to the same degree. We may feel assured that the inherited effects of the long-continued use or disuse of parts will have done much in the same direction with natural selection. Modifications formerly of importance, though no longer of any special use, are long-inherited. When one part is modified, other parts change through the principle of correlation, of which we have instances in many curious cases of correlated monstrosities. Something may be attributed to the direct and definite action of the surrounding conditions of life, such as abundant food, heat or moisture; and lastly, many characters of slight physiological importance, some indeed of considerable importance, have been gained through sexual selection.

No doubt man, as well as every other animal, presents structures, which seem to our limited knowledge, not to be now of any service to him, nor to have been so formerly, either for the general conditions of life, or in the relations of one sex to the other. Such structures cannot be accounted for by any form of selection, or by the inherited effects of the use and disuse of parts. We know, however, that many strange and strongly-marked peculiarities of structure occasionally appear in our domesticated productions, and if their unknown causes were to act more uniformly, they would probably become common to all the individuals of the species. We may hope hereafter to understand something about the causes of such occasional modifications, especially through the study of monstrosities: hence the labours of experimentalists such as those of M. Camille Dareste, are full of promise for the future. In general we can only say that the cause of each slight variation and of each monstrosity lies much more in the constitution of the organism, than in the nature of the surrounding conditions; though new and changed conditions certainly play an important part in exciting organic changes of many kinds.

Through the means just specified, aided perhaps by others as yet undiscovered, man has been raised to his present state. But since he attained to the rank of manhood, he has diverged into distinct races, or as they may be more fitly called, sub-species. Some of these, such as the Negro and European, are so distinct that, if specimens had been brought to a naturalist without any further information, they would undoubtedly have been considered by him as good and true species. Nevertheless all the races agree in so many unimportant details of structure and in so many mental peculiarities that these can be accounted for only by inheritance from a common progenitor; and a progenitor thus characterised would probably deserve to rank as man.

It must not be supposed that the divergence of each race from the other races, and of all from a common stock, can be traced back to any one pair of progenitors. On the contrary, at every stage in the process of modification, all the individuals which were in any way better fitted for their conditions of life, though in different degrees, would have survived in greater numbers than the less well-fitted. The process would have been like that followed by man, when he does not intentionally select particular individuals, but breeds from all the superior individuals, and neglects the inferior. He thus slowly but surely modifies his stock, and unconsciously forms a new strain. So with respect to modifications acquired independently of selection, and due to variations arising from the nature of the organism and the action of the surrounding conditions, or from changed habits of life, no single pair will have been modified much more than the other pairs inhabiting the same country, for all will have been continually blended through free intercrossing.

By considering the embryological structure of man,- the homologies which he presents with the lower animals,- the rudiments which he retains,- and the reversions to which he is liable, we can partly recall in imagination the former condition of our early progenitors; and can approximately place them in their proper place in the zoological series. We thus learn that man is descended from a hairy, tailed quadruped, probably arboreal in its habits, and an inhabitant of the Old World. This creature, if its whole structure had been examined by a naturalist, would have been classed amongst the Quadrumana, as surely as the still more ancient progenitor of the Old and New World monkeys. The Quadrumana and all the higher mammals are probably derived from an ancient marsupial animal, and this through a long series of diversified forms, from some amphibian-like creature, and this again from some fish-like animal. In the dim obscurity of the past we can see that the early progenitor of all the Vertebrata must have been an aquatic animal provided with branchiae, with the two sexes united in the same individual, and with the most important organs of the body (such as the brain and heart) imperfectly or not at all developed. This animal seems to have been more like the larvae of the existing marine ascidians than any other known form.

The high standard of our intellectual powers and moral disposition is the greatest difficulty which presents itself, after we have been driven to this conclusion on the origin of man. But every one who admits the principle of evolution, must see that the mental powers of the higher animals, which are the same in kind with those of man, though so different in degree, are capable of advancement. Thus the interval between the mental powers of one of the higher apes and of a fish, or between those of an ant and scale-insect, is immense; yet their development does not offer any special difficulty; for with our domesticated animals, the mental faculties are certainly variable, and the variations are inherited. No one doubts that they are of the utmost importance to animals in a state of nature. Therefore the conditions are favourable for their development through natural selection. The same conclusion may be extended to man; the intellect must have been all-important to him, even at a very remote period, as enabling him to invent and use language, to make weapons, tools, traps, &c., whereby with the aid of his social habits, he long ago became the most dominant of all living creatures.

A great stride in the development of the intellect will have followed, as soon as the half-art and half-instinct of language came into use; for the continued use of language will have reacted on the brain and produced an inherited effect; and this again will have reacted on the improvement of language. As Mr. Chauncey Wright* has well remarked, the largeness of the brain in man relatively to his body, compared with the lower animals, may be attributed in chief part to the early use of some simple form of language,- that wonderful engine which affixes signs to all sorts of objects and qualities, and excites trains of thought which would never arise from the mere impression of the senses, or if they did arise could not be followed out. The higher intellectual powers of man, such as those of ratiocination, abstraction, self-consciousness, &c., probably follow from the continued improvement and exercise of the other mental faculties.

* "On the Limits of Natural Selection," in the North American Review, Oct., 1870, p. 295.

The development of the moral qualities is a more interesting problem. The foundation lies in the social instincts, including under this term the family ties. These instincts are highly complex, and in the case of the lower animals give special tendencies towards certain definite actions; but the more important elements are love, and the distinct emotion of sympathy. Animals endowed with the social instincts take pleasure in one another's company, warn one another of danger, defend and aid one another in many ways. These instincts do not extend to all the individuals of the species, but only to those of the same community. As they are highly beneficial to the species, they have in all probability been acquired through natural selection.

A moral being is one who is capable of reflecting on his past actions and their motives- of approving of some and disapproving of others; and the fact that man is the one being who certainly deserves this designation, is the greatest of all distinctions between him and the lower animals. But in the fourth chapter I have endeavoured to shew that the moral sense follows, firstly, from the enduring and ever-present nature of the social instincts; secondly, from man's appreciation of the approbation and disapprobation of his fellows; and thirdly, from the high activity of his mental faculties, with past impressions extremely vivid; and in these latter respects he differs from the lower animals. Owing to this condition of mind, man cannot avoid looking both backwards and forwards, and comparing past impressions. Hence after some temporary desire or passion has mastered his social instincts, he reflects and compares the now weakened impression of such past impulses with the ever-present social instincts; and he then feels that sense of dissatisfaction which all unsatisfied instincts leave behind them, he therefore resolves to act differently for the future,- and this is conscience. Any instinct, permanently stronger or more enduring than another, gives rise to a feeling which we express by saying that it ought to be obeyed. A pointer dog, if able to reflect on his past conduct, would say to himself, I ought (as indeed we say of him) to have pointed at that hare and not have yielded to the passing temptation of hunting it.

Social animals are impelled partly by a wish to aid the members of their community in a general manner, but more commonly to perform certain definite actions. Man is impelled by the same general wish to aid his fellows; but has few or no special instincts. He differs also from the lower animals in the power of expressing his desires by words, which thus become a guide to the aid required and bestowed. The motive to give aid is likewise much modified in man: it no longer consists solely of a blind instinctive impulse, but is much influenced by the praise or blame of his fellows. The appreciation and the bestowal of praise and blame both rest on sympathy; and this emotion, as we have seen, is one of the most important elements of the social instincts. Sympathy, though gained as an instinct, is also much strengthened by exercise or habit. As all men desire their own happiness, praise or blame is bestowed on actions and motives, according as they lead to this end; and as happiness is an essential part of the general good, the greatest-happinesss principle indirectly serves as a nearly safe standard of right and wrong. As the reasoning powers advance and experience is gained, the remoter effects of certain lines of conduct on the character of the individual, and on the general good, are perceived; and then the self-regarding virtues come within the scope of public opinion, and receive praise, and their opposites blame. But with the less civilised nations reason often errs, and many bad customs and base superstitions come within the same scope, and are then esteemed as high virtues, and their breach as heavy crimes.

The moral faculties are generally and justly esteemed as of higher value than the intellectual powers. But we should bear in mind that the activity of the mind in vividly recalling past impressions is one of the fundamental though secondary bases of conscience. This affords the strongest argument for educating and stimulating in all possible ways the intellectual faculties of every human being. No doubt a man with a torpid mind, if his social affections and sympathies are well developed, will be led to good actions, and may have a fairly sensitive conscience. But whatever renders the imagination more vivid and strengthens the habit of recalling and comparing past impressions, will make the conscience more sensitive, and may even somewhat compensate for weak social affections and sympathies.

The moral nature of man has reached its present standard, partly through the advancement of his reasoning powers and consequently of a just public opinion, but especially from his sympathies having been rendered more tender and widely diffused through the effects of habit, example, instruction, and reflection. It is not improbable that after long practice virtuous tendencies may be inherited. With the more civilised races, the conviction of the existence of an all-seeing Deity has had a potent influence on the advance of morality. Ultimately man does not accept the praise or blame of his fellows as his sole guide, though few escape this influence, but his habitual convictions, controlled by reason, afford him the safest rule. His conscience then becomes the supreme judge and monitor. Nevertheless the first foundation or origin of the moral sense lies in the social instincts, including sympathy; and these instincts no doubt were primarily gained, as in the case of the lower animals, through natural selection.

The belief in God has often been advanced as not only the greatest, but the most complete of all the distinctions between man and the lower animals. It is however impossible, as we have seen, to maintain that this belief is innate or instinctive in man. On the other hand a belief in all-pervading spiritual agencies seems to be universal; and apparently follows from a considerable advance in man's reason, and from a still greater advance in his faculties of imagination, curiosity and wonder. I am aware that the assumed instinctive belief in God has been used by many persons as an argument for His existence. But this is a rash argument, as we should thus be compelled to believe in the existence of many cruel and malignant spirits, only a little more powerful than man; for the belief in them is far more general than in a beneficent Deity. The idea of a universal and beneficent Creator does not seem to arise in the mind of man, until he has been elevated by long-continued culture.

He who believes in the advancement of man from some low organised form, will naturally ask how does this bear on the belief in the immortality of the soul. The barbarous races of man, as Sir J. Lubbock has shewn, possess no clear belief of this kind; but arguments derived from the primeval beliefs of savages are, as we have just seen, of little or no avail. Few persons feel any anxiety from the impossibility of determining at what precise period in the development of the individual, from the first trace of a minute germinal vesicle, man becomes an immortal being; and there is no greater cause for anxiety because the period cannot possibly be determined in the gradually ascending organic scale.*

* The Rev. J. A. Picton gives a discussion to this effect in his New Theories and the Old Faith, 1870.

I am aware that the conclusions arrived at in this work will be denounced by some as highly irreligious; but he who denounces them is bound to shew why it is more irreligious to explain the origin of man as a distinct species by descent from some lower form, through the laws of variation and natural selection, than to explain the birth of the individual through the laws of ordinary reproduction. The birth both of the species and of the individual are equally parts of that grand sequence of events, which our minds refuse to accept as the result of blind chance. The understanding revolts at such a conclusion, whether or not we are able to believe that every slight variation of structure,- the union of each pair in marriage, the dissemination of each seed,- and other such events, have all been ordained for some special purpose.

Sexual selection has been treated at great length in this work; for, as I have attempted to shew, it has played an important part in the history of the organic world. I am aware that much remains doubtful, but I have endeavoured to give a fair view of the whole case. In the lower divisions of the animal kingdom, sexual selection seems to have done nothing: such animals are often affixed for life to the same spot, or have the sexes combined in the same individual, or what is still more important, their perceptive and intellectual faculties are not sufficiently advanced to allow of the feelings of love and jealousy, or of the exertion of choice. When, however, we come to the Arthropoda and Vertebrata, even to the lowest classes in these two great sub-kingdoms, sexual selection has effected much.

In the several great classes of the animal kingdom,- in mammals, birds, reptiles, fishes, insects, and even crustaceans,- the differences between the sexes follow nearly the same rules. The males are almost always the wooers; and they alone are armed with special weapons for fighting with their rivals. They are generally stronger and larger than the females, and are endowed with the requisite qualities of courage and pugnacity. They are provided, either exclusively or in a much higher degree than the females, with organs for vocal or instrumental music, and with odoriferous glands. They are ornamental with infinitely diversified appendages, and with the most brilliant or conspicuous colours, often arranged in elegant patterns, whilst the females are unadorned. When the sexes differ in more important structures, it is the male which is provided with special sense-organs for discovering the female, with locomotive organs for reaching her, and often with prehensile organs for holding her. These various structures for charming or securing the female are often developed in the male during only part of the year, namely the breeding-season. They have in many cases been more or less transferred to the females; and in the latter case they often appear in her as mere rudiments. They are lost or never gained by the males after emasculation. Generally they are not developed in the male during early youth, but appear a short time before the age for reproduction. Hence in most cases the young of both sexes resemble each other; and the female somewhat resembles her young offspring throughout life. In almost every great class a few anomalous cases occur, where there has been an almost complete transposition of the characters proper to the two sexes; the females assuming characters which properly belong to the males. This surprising uniformity in the laws regulating the differences between the sexes in so many and such widely separated classes, is intelligible if we admit the action of one common cause, namely sexual selection.

Sexual selection depends on the success of certain individuals over others of the same sex, in relation to the propagation of the species; whilst natural selection depends on the success of both sexes, at all ages, in relation to the general conditions of life. The sexual struggle is of two kinds; in the one it is between individuals of the same sex, generally the males, in order to drive away or kill their rivals, the females remaining passive; whilst in the other, the struggle is likewise between the individuals of the same sex, in order to excite or charm those of the opposite sex, generally the females, which no longer remain passive, but select the more agreeable partners. This latter kind of selection is closely analogous to that which man unintentionally, yet effectually, brings to bear on his domesticated productions, when he preserves during a long period the most pleasing or useful individuals, without any wish to modify the breed.

The laws of inheritance determine whether characters gained through sexual selection by either sex shall be transmitted to the same sex, or to both; as well as the age at which they shall be developed. It appears that variations arising late in life are commonly transmitted to one and the same sex. Variability is the necessary basis for the action of selection, and is wholly independent of it. It follows from this, that variations of the same general nature have often been taken advantage of and accumulated through sexual selection in relation to the propagation of the species, as well as through natural selection in relation to the general purposes of life. Hence secondary sexual characters, when equally transmitted to both sexes can be distinguished from ordinary specific characters only by the light of analogy. The modifications acquired through sexual selection are often so strongly pronounced that the two sexes have frequently been ranked as distinct species, or even as distinct genera. Such strongly-marked differences must be in some manner highly important; and we know that they have been acquired in some instances at the cost not only of inconvenience, but of exposure to actual danger.

The belief in the power of sexual selection rests chiefly on the following considerations. Certain characters are confined to one sex; and this alone renders it probable that in most cases they are connected with the act of reproduction. In innumerable instances these characters are fully developed only at maturity, and often during only a part of the year, which is always the breeding-season. The males (passing over a few exceptional cases) are the more active in courtship; they are the better armed, and are rendered the more attractive in various ways. It is to be especially observed that the males display their attractions with elaborate care in the presence of the females; and that they rarely or never display them excepting during the season of love. It is incredible that all this should be purposeless. Lastly we have distinct evidence with some quadrupeds and birds, that the individuals of one sex are capable of feeling a strong antipathy or preference for certain individuals of the other sex.

Bearing in mind these facts, and the marked results of man's unconscious selection, when applied to domesticated animals and cultivated plants, it seems to me almost certain that if the individuals of one sex were during a long series of generations to prefer pairing with certain individuals of the other sex, characterised in some peculiar manner, the offspring would slowly but surely become modified in this same manner. I have not attempted to conceal that, excepting when the males are more numerous than the females, or when polygamy prevails, it is doubtful how the more attractive males succeed in leaving a large number of offspring to inherit their superiority in ornaments or other charms than the less attractive males; but I have shewn that this would probably follow from the females,- especially the more vigorous ones, which would be the first to breed,- preferring not only the more attractive but at the same time the more vigorous and victorious males.

Although we have some positive evidence that birds appreciate bright and beautiful objects, as with the bower-birds of Australia, and although they certainly appreciate the power of song, yet I fully admit that it is astonishing that the females of many birds and some mammals should be endowed with sufficient taste to appreciate ornaments, which we have reason to attribute to sexual selection; and this is even more astonishing in the case of reptiles, fish, and insects. But we really know little about the minds of the lower animals. It cannot be supposed, for instance, that male birds of paradise or peacocks should take such pains in erecting, spreading, and vibrating their beautiful plumes before the females for no purpose. We should remember the fact given on excellent authority in a former chapter, that several peahens, when debarred from an admired male, remained widows during a whole season rather than pair with another bird.

Nevertheless I know of no fact in natural history more wonderful than that the female Argus pheasant should appreciate the exquisite shading of the ball-and-socket ornaments and the elegant patterns on the wing-feather of the male. He who thinks that the male was created as he now exists must admit that the great plumes, which prevent the wings from being used for flight, and which are displayed during courtship and at no other time in a manner quite peculiar to this one species, were given to him as an ornament. If so, he must likewise admit that the female was created and endowed with the capacity of appreciating such ornaments. I differ only in the conviction that the male Argus pheasant acquired his beauty gradually, through the preference of the females during many generations for the more highly ornamented males; the aesthetic capacity of the females having been advanced through exercise or habit, just as our own taste is gradually improved. In the male through the fortunate chance of a few feathers being left unchanged, we can distinctly trace how simple spots with a little fulvous shading on one side may have been developed by small steps into the wonderful ball-and-socket ornaments; and it is probable that they were actually thus developed.

Everyone who admits the principle of evolution, and yet feels great difficulty in admitting that female mammals, birds, reptiles, and fish, could have acquired the high taste implied by the beauty of the males, and which generally coincides with our own standard, should reflect that the nerve-cells of the brain in the highest as well as in the lowest members of the vertebrate series, are derived from those of the common progenitor of this great kingdom. For we can thus see how it has come to pass that certain mental faculties, in various and widely distinct groups of animals, have been developed in nearly the same manner and to nearly the same degree.

The reader who has taken the trouble to go through the several chapters devoted to sexual selection, will be able to judge how far the conclusions at which I have arrived are supported by sufficient evidence. If he accepts these conclusions he may, I think, safely extend them to mankind; but it would be superfluous here to repeat what I have so lately said on the manner in which sexual selection apparently has acted on man, both on the male and female side, causing the two sexes to differ in body and mind, and the several races to differ from each other in various characters, as well as from their ancient and lowly-organised progenitors.

He who admits the principle of sexual selection will be led to the remarkable conclusion that the nervous system not only regulates most of the existing functions of the body, but has indirectly influenced the progressive development of various bodily structures and of certain mental qualities. Courage, pugnacity, perseverance, strength and size of body, weapons of all kinds, musical organs, both vocal and instrumental, bright colours and ornamental appendages, have all been indirectly gained by the one sex or the other, through the exertion of choice, the influence of love and jealousy, and the appreciation of the beautiful in sound, colour or form; and these powers of the mind manifestly depend on the development of the brain.

Man scans with scrupulous care the character and pedigree of his horses, cattle, and dogs before he matches them; but when he comes to his own marriage he rarely, or never, takes any such care. He is impelled by nearly the same motives as the lower animals, when they are left to their own free choice, though he is in so far superior to them that he highly values mental charms and virtues. On the other hand he is strongly attracted by mere wealth or rank. Yet he might by selection do something not only for the bodily constitution and frame of his offspring, but for their intellectual and moral qualities. Both sexes ought to refrain from marriage if they are in any marked degree inferior in body or mind; but such hopes are Utopian and will never be even partially realised until the laws of inheritance are thoroughly known. Everyone does good service, who aids towards this end. When the principles of breeding and inheritance are better understood, we shall not hear ignorant members of our legislature rejecting with scorn a plan for ascertaining whether or not consanguineous marriages are injurious to man.

The advancement of the welfare of mankind is a most intricate problem: all ought to refrain from marriage who cannot avoid abject poverty for their children; for poverty is not only a great evil, but tends to its own increase by leading to recklessness in marriage. On the other hand, as Mr. Galton has remarked, if the prudent avoid marriage, whilst the reckless marry, the inferior members tend to supplant the better members of society. Man, like every other animal, has no doubt advanced to his present high condition through a struggle for existence consequent on his rapid multiplication; and if he is to advance still higher, it is to be feared that he must remain subject to a severe struggle. Otherwise he would sink into indolence, and the more gifted men would not be more successful in the battle of life than the less gifted. Hence our natural rate of increase, though leading to many and obvious evils, must not be greatly diminished by any means. There should be open competition for all men; and the most able should not be prevented by laws or customs from succeeding best and rearing the largest number of offspring. Important as the struggle for existence has been and even still is, yet as far as the highest part of man's nature is concerned there are other agencies more important. For the moral qualities are advanced, either directly or indirectly, much more through the effects of habit, the reasoning powers, instruction, religion, &c., than through natural selection; though to this latter agency may be safely attributed the social instincts, which afforded the basis for the development of the moral sense.

The main conclusion arrived at in this work, namely, that man is descended from some lowly organised form, will, I regret to think, be highly distasteful to many. But there can hardly be a doubt that we are descended from barbarians. The astonishment which I felt on first seeing a party of Fuegians on a wild and broken shore will never be forgotten by me, for the reflection at once rushed into my mind- such were our ancestors. These men were absolutely naked and bedaubed with paint, their long hair was tangled, their mouths frothed with excitement, and their expression was wild, startled, and distrustful. They possessed hardly any arts, and like wild animals lived on what they could catch; they had no government, and were merciless to every one not of their own small tribe. He who has seen a savage in his native land will not feel much shame, if forced to acknowledge that the blood of some more humble creature flows in his veins. For my own part I would as soon be descended from that heroic little monkey, who braved his dreaded enemy in order to save the life of his keeper, or from that old baboon, who descending from the mountains, carried away in triumph his young comrade from a crowd of astonished dogs- as from a savage who delights to torture his enemies, offers up bloody sacrifices, practices infanticide without remorse, treats his wives like slaves, knows no decency, and is haunted by the grossest superstitions.

Man may be excused for feeling some pride at having risen, though not through his own exertions, to the very summit of the organic scale; and the fact of his having thus risen, instead of having been aboriginally placed there, may give him hope for a still higher destiny in the distant future. But we are not here concerned with hopes or fears, only with the truth as far as our reason permits us to discover it; and I have given the evidence to the best of my ability. We must, however, acknowledge, as it seems to me, that man with all his noble qualities, with sympathy which feels for the most debased, with benevolence which extends not only to other men but to the humblest living creature, with his god-like intellect which has penetrated into the movements and constitution of the solar system- with all these exalted powers- Man still bears in his bodily frame the indelible stamp of his lowly origin.

SUPPLEMENTAL NOTE.

ON SEXUAL SELECTION IN RELATION TO MONKEYS.

Reprinted from NATURE, November 2, 1876, p. 18.

IN the discussion on Sexual Selection in my Descent of Man, no case interested and perplexed me so much as the brightly-coloured hinder ends and adjoining parts of certain monkeys. As these parts are more brightly coloured in one sex than the other, and as they become more brilliant during the season of love, I concluded that the colours had been gained as a sexual attraction. I was well aware that I thus laid myself open to ridicule; though in fact it is not more surprising that a monkey should display his bright-red hinder end than that a peacock should display his magnificent tail. I had, however, at that time no evidence of monkeys exhibiting this part of their bodies during their courtship; and such display in the case of birds affords the best evidence that the ornaments of the males are of service to them by attracting or exciting the females. I have lately read an article by Joh. von Fischer, of Gotha, published in Der Zoologische Garten, April, 1876, on the expression of monkeys under various emotions, which is well worthy of study by any one interested in the subject, and which shews that the author is a careful and acute observer. In this article there is an account of the behaviour of a young male mandrill when he first beheld himself in a looking-glass, and it is added, that after a time he turned round and presented his red hinder end to the glass. Accordingly I wrote to Herr J. von Fischer to ask what he supposed was the meaning of this strange action, and he has sent me two long letters full of new and curious details, which will, I hope, be hereafter published. He says that he was himself at first perplexed by the above action, and was thus led carefully to observe several individuals of various other species of monkeys, which he has long kept in his house. He finds that not only the mandrill (Cynocephalus mormon) but the drill (C. leucophaeus) and three other kinds of baboons (C. hamadryas, sphinx, and babouin), also Cynopithecus niger, and Macacus rhesus and nemestrinus, turn this part of their bodies, which in all these species is more or less brightly coloured, to him when they are pleased, and to other persons as a sort of greeting. He took pains to cure a Macacus rhesus, which he had kept for five years, of this indecorous habit, and at last succeeded. These monkeys are particularly apt to act in this manner, grinning at the same time, when first introduced to a new monkey, but often also to their old monkey friends; and after this mutual display they begin to play together. The young mandrill ceased spontaneously after a time to act in this manner towards his master, von Fischer, but continued to do so towards persons who were strangers and to new monkeys. A young Cynopithecus niger never acted, excepting on one occasion, in this way towards his master, but frequently towards strangers, and continues to do so up to the present time. From these facts von Fischer concludes that the monkeys which behaved in this manner before a looking-glass (viz., the mandrill, drill, Cynopithecus niger, Macacus rhesus and nemestrinus) acted as if their reflection were a new acquaintance. The mandrill and drill, which have their hinder ends especially ornamented, display it even whilst quite young, more frequently and more ostentatiously than do the other kinds. Next in order comes Cynocephalus hamadryas, whilst the other species act in this manner seldomer. The individuals, however, of the same species vary in this respect, and some which were very shy never displayed their hinder ends. It deserves especial attention that von Fischer has never seen any species purposely exhibit the hinder part of its body, if not at all coloured. This remark applies to many individuals of Macacus cynomolgus and Cercocebus radiatus (which is closely allied to M. rhesus), to three species of Cercopithecus and several American monkeys. The habit of turning the hinder ends as a greeting to an old friend or new acquaintance, which seems to us so odd, is not really more so than the habits of many savages, for instance that of rubbing their bellies with their hands, or rubbing noses together. The habit with the mandrill and drill seems to be instinctive or inherited, as it was followed by very young animals; but it is modified or guided, like so many other instincts, by observation, for von Fischer says that they take pains to make their display fully; and if made before two observers, they turn to him who seems to pay the most attention.

With respect to the origin of the habit, von Fischer remarks that his monkeys like to have their naked hinder ends patted or stroked, and that they then grunt with pleasure. They often also turn this part of their bodies to other monkeys to have bits of dirt picked off, and so no doubt it would be with respect to thorns. But the habit with adult animals is connected to a certain extent with sexual feelings, for von Fischer watched through a glass door a female Cynopithecus niger, and she during several days, "umdrehte und dem Mannchen mit gurgelnden Tonen die stark gerothete Sitzflache zeigte, was ich fruher nie an diesem Thier bemerkt hatte. Beim Anblick dieses Gegenstandes erregte sich das Mannchen sichtlich, denn es polterte heftig an den Staben, ebenfalls gurgelnde Laute ausstossend." As all the monkeys which have the hinder parts of their bodies more or less brightly coloured live, according to von Fischer, in open rocky places, he thinks that these colours serve to render one sex conspicuous at a distance to the other; but, as monkeys are such gregarious animals, I should have thought that there was no need for the sexes to recognise each other at a distance. It seems to me more probable that the bright colours, whether on the face or hinder end, or, as in the mandrill, on both, serve as a sexual ornament and attraction. Anyhow, as we now know that monkeys have the habit of turning their hinder ends towards other monkeys, it ceases to be at all surprising that it should have been this part of their bodies which has been more or less decorated. The fact that it is only the monkeys thus characterised which, as far as at present known, act in this manner as a greeting towards other monkeys renders it doubtful whether the habit was first acquired from some independent cause, and that afterwards the parts in question were coloured as a sexual ornament; or whether the colouring and the habit of turning round were first acquired through variation and sexual selection, and that afterwards the habit was retained as a sign of pleasure or as a greeting, through the principle of inherited association. This principle apparently comes into play on many occasions: thus it is generally admitted that the songs of birds serve mainly as an attraction during the season of love, and that the leks, or great congregations of the black-grouse, are connected with their courtship; but the habit of singing has been retained by some birds when they feel happy, for instance by the common robin, and the habit of congregating has been retained by the black-grouse during other seasons of the year.

I beg leave to refer to one other point in relation to sexual selection. It has been objected that this form of selection, as far as the ornaments of the males are concerned, implies that all females within the same district must possess and exercise exactly the same taste. It should, however, be observed, in the first place, that although the range of variation of a species may be very large, it is by no means indefinite. I have elsewhere given a good instance of this fact in the pigeon, of which there are at least a hundred varieties differing widely in their colours, and at least a score of varieties of the fowl differing in the same kind of way; but the range of colour in these two species is extremely distinct. Therefore the females of natural species cannot have an unlimited scope for their taste. In the second place, I presume that no supporter of the principle of sexual selection believes that the females select particular points of beauty in the males; they are merely excited or attracted in a greater degree by one male than by another, and this seems often to depend, especially with birds, on brilliant colouring. Even man, excepting perhaps an artist, does not analyse the slight differences in the features of the woman whom he may admire, on which her beauty depends. The male mandrill has not only the hinder end of his body, but his face gorgeously coloured and marked with oblique ridges, a yellow beard, and other ornaments. We may infer from what we see of the variation of animals under domestication, that the above several ornaments of the mandrill were gradually acquired by one individual varying a little in one way, and another individual in another way. The males which were the handsomest or the most attractive in any manner to the females would pair oftenest, and would leave rather more offspring than other males. The offspring of the former, although variously intercrossed, would either inherit the peculiarities of their fathers or transmit an increased tendency to vary in the same manner. Consequently the whole body of males inhabiting the same country would tend from the effects of constant intercrossing to become modified almost uniformly, but sometimes a little more in one character and sometimes in another, though at an extremely slow rate; all ultimately being thus rendered more attractive to the females. The process is like that which I have called unconscious selection by man, and of which I have given several instances. In one country the inhabitants value a fleet or light dog or horse, and in another country a heavier and more powerful one; in neither country is there any selection of individual animals with lighter or stronger bodies and limbs; nevertheless after a considerable lapse of time the individuals are found to have been modified in the desired manner almost uniformly, though differently in each country. In two absolutely distinct countries inhabited by the same species, the individuals of which can never during long ages have intermigrated and intercrossed, and where, moreover, the variations will probably not have been identically the same, sexual selection might cause the males to differ. Nor does the belief appear to me altogether fanciful that two sets of females, surrounded by a very different environment, would be apt to acquire somewhat different tastes with respect to form, sound, or colour. However this may be, I have given in my Descent of Man instances of closely-allied birds inhabiting distinct countries, of which the young and the females cannot be distinguished, whilst the adult males differ considerably, and this may be attributed with much probability to the action of sexual selection.

THE END .