THE
DESCENT OF MAN,
AND
SELECTION IN RELATION TO SEX.
By CHARLES DARWIN, M.A., F.R.S., &c.
IN TWO VOLUMES.—Vol. I.
WITH ILLUSTRATIONS.
LONDON:
JOHN MURRAY, ALBEMARLE STREET.
1871.
[The right of Translation is reserved.]
BY THE SAME AUTHOR.
ON THE ORIGIN OF SPECIES BY MEANS OF NATURAL SELECTION; or, The Preservation of Favoured Races in the Struggle for Life; Fifth Edition (Tenth Thousand), with Additions and Corrections. 1869. Murray.
THE VARIATION OF ANIMALS AND PLANTS UNDER DOMESTICATION. In two vols. With Illustrations. 1868. Murray.
ON THE VARIOUS CONTRIVANCES by which BRITISH AND FOREIGN ORCHIDS ARE FERTILISED BY INSECTS; and on the Good Effects of Crossing. With numerous Woodcuts. Murray.
A NATURALIST‘S VOYAGE ROUND THE WORLD; or, A Journal of Researches into the Natural History and Geology of the Countries visited during the voyage of H.M.S. ‘Beagle,’ under the command of Captain FitzRoy, R.N. Eleventh Thousand. Murray.
ON THE STRUCTURE AND DISTRIBUTION OF CORAL REEFS. Smith, Elder, & Co.
GEOLOGICAL OBSERVATIONS ON VOLCANIC ISLANDS. Smith, Elder, & Co.
GEOLOGICAL OBSERVATIONS ON SOUTH AMERICA. Smith, Elder & Co.
A MONOGRAPH OF THE CIRRIPEDIA. With numerous Illustrations. 2 vols. 8vo. Hardwicke.
ON THE MOVEMENTS AND HABITS OF CLIMBING PLANTS. With Woodcuts. Williams & Norgate.
LONDON: PRINTED BY WILLIAM CLOWES AND SONS, STAMFORD STREET, AND CHARING CROSS.
CONTENTS.
Introduction Page 1-5
PART I.
ON THE DESCENT OF MAN
CHAPTER I. The Evidence of the Descent of man from some Lower form. Nature of the evidence bearing on the origin of man—Homologous structures in man and the lower animals—Miscellaneous points of correspondence—Development—Rudimentary structures, muscles, sense-organs, hair, bones, reproductive organs, &c.—The bearing of these three great classes of facts on the origin of man 9-33 CHAPTER II. Comparison of the Mental Powers of Man and the Lower Animals. The difference in mental power between the highest ape and the lowest savage, immense—Certain instincts in common—The emotions—Curiosity—Imitation—Attention—Memory—Imagination—Reason—Progressive improvement—Tools and weapons used by animals—Language—Self-consciousness—Sense of beauty—Belief in God, spiritual agencies, superstitions 34-69 CHAPTER III. Comparison of the Mental Powers of Man and the Lower Animals—continued. The difference in mental power between the highest ape and the lowest savage, immense—Certain instincts in common—The emotions—Curiosity—Imitation—Attention—Memory—Imagination—Reason—Progressive improvement—Tools and weapons used by animals—Language—Self-consciousness—Sense of beauty—Belief in God, spiritual agencies, superstitions 70-106 CHAPTER IV. On the Manner of Development of Man from some Lower Form. Variability of body and mind in man—Inheritance—Causes of variability—Laws of variation the same in man as in the lower animals—Direct action of the conditions of life—Effects of the increased use and disuse of parts—Arrested development—Reversion—Correlated variation—Rate of increase—Checks to increase—Natural selection—Man the most dominant animal in the world—Importance of his corporeal structure—The causes which have led to his becoming erect—Consequent changes of structure—Decrease in size of the canine teeth—Increased size and altered shape of the skull—Nakedness—Absence of a tail—Defenceless condition of man 107-157 CHAPTER V. On the Development of the Intellectual and Moral Faculties during Primeval and Civilised Times. The advancement of the intellectual powers through natural selection—Importance of imitation—Social and moral faculties—Their development within the limits of the same tribe—Natural selection as affecting civilised nations—Evidence that civilised nations were once barbarous 158-184 CHAPTER VI. On the Affinities and Genealogy of Man. Position of man in the animal series—The natural system genealogical—Adaptive characters of slight value—Various small points of resemblance between man and the Quadrumana—Rank of man in the natural system—Birthplace and antiquity of man—Absence of fossil connecting-links—Lower stages in the genealogy of man, as inferred, firstly from his affinities and secondly from his structure—Early androgynous condition of the Vertebrata—Conclusion 185-213 CHAPTER VII. On the Races of Man.PART II.
SEXUAL SELECTION.
CHAPTER VIII. Principles of Sexual Selection. Secondary sexual characters—Sexual selection—Manner of action—Excess of males—Polygamy—The male alone generally modified through sexual selection—Eagerness of the male—Variability of the male—Choice exerted by the female—Sexual compared with natural selection—Inheritance at corresponding periods of life, at corresponding seasons of the year, and as limited by sex—Relations between the several forms of inheritance—Causes why one sex and the young are not modified through sexual selection—Supplement on the proportional numbers of the two sexes throughout the animal kingdom—On the limitation of the numbers of the two sexes through natural selection 253-320 CHAPTER IX. Secondary Sexual Characters in the Lower Classes of the Animal Kingdom. These characters absent in the lowest classes—Brilliant colours—Mollusca—Annelids—Crustacea, secondary sexual characters strongly developed; dimorphism; colour; characters not acquired before maturity—Spiders, sexual colours of; stridulation by the males—Myriapoda 321-340 CHAPTER X. Secondary Sexual Characters of Insects. Diversified structures possessed by the males for seizing the females—Differences between the sexes, of which the meaning is not understood—Difference in size between the sexes—Thysanura—Diptera—Hemiptera—Homoptera, musical powers possessed by the males alone—Orthoptera, musical instruments of the males, much diversified in structure; pugnacity; colours—Neuroptera, sexual differences in colour—Hymenoptera, pugnacity and colours—Coleoptera, colours; furnished with great horns, apparently as an ornament; battles; stridulating organs generally common to both sexes 341-385 CHAPTER XI. Insects, continued.—Order Lepidoptera. Courtship of butterflies—Battles—Ticking noise—Colours common to both sexes, or more brilliant in the males—Examples—Not due to the direct action of the conditions of life—Colours adapted for protection—Colours of moths—Display—Perceptive powers of the Lepidoptera—Variability—Causes of the difference in colour between the males and females—Mimickry, female butterflies more brilliantly coloured than the males—Bright colours of caterpillars—Summary and concluding remarks on the secondary sexual characters of insects—Birds and insects compared 386-423 THE DESCENT OF MAN;
AND ON
SELECTION IN RELATION TO SEX.
INTRODUCTION.
The nature of the following work will be best understood by a brief account of how it came to be written. During many years I collected notes on the origin or descent of man, without any intention of publishing on the subject, but rather with the determination not to publish, as I thought that I should thus only add to the prejudices against my views. It seemed to me sufficient to indicate, in the first edition of my ‘Origin of Species,’ that by this work “light would be thrown on the origin of man and his history;” and this implies that man must be included with other organic beings in any general conclusion respecting his manner of appearance on this earth. Now the case wears a wholly different aspect. When a naturalist like Carl Vogt ventures to say in his address as President of the National Institution of Geneva (1869), “personne, en Europe au moins, n’ose plus soutenir la création indépendante et de toutes pièces, des espèces,” it is manifest that at least a large number of naturalists must admit that species are the modified descendants of other species; and this especially holds good with the younger and rising naturalists. The greater number accept the agency of natural selection; though some urge, whether with justice the future must decide, that I have greatly overrated its importance. Of the older and honoured chiefs in natural science, many unfortunately are still opposed to evolution in every form.
In consequence of the views now adopted by most naturalists, and which will ultimately, as in every other case, be followed by other men, I have been led to put together my notes, so as to see how far the general conclusions arrived at in my former works were applicable to man. This seemed all the more desirable as I had never deliberately applied these views to a species taken singly. When we confine our attention to any one form, we are deprived of the weighty arguments derived from the nature of the affinities which connect together whole groups of organisms—their geographical distribution in past and present times, and their geological succession. The homological structure, embryological development, and rudimentary organs of a species, whether it be man or any other animal, to which our attention may be directed, remain to be considered; but these great classes of facts afford, as it appears to me, ample and conclusive evidence in favour of the principle of gradual evolution. The strong support derived from the other arguments should, however, always be kept before the mind.
The sole object of this work is to consider, firstly, whether man, like every other species, is descended from some pre-existing form; secondly, the manner of his development; and thirdly, the value of the differences between the so-called races of man. As I shall confine myself to these points, it will not be necessary to describe in detail the
This work contains hardly any original facts in regard to man; but as the conclusions at which I arrived, after drawing up a rough draft, appeared to me interesting, I thought that they might interest others. It has often and confidently been asserted, that man’s origin can never be known: but ignorance more frequently begets confidence than does knowledge: it is those who know little, and not those who know much, who so positively assert that this or that problem will never be solved by science. The conclusion that man is the co-descendant with other species of some ancient, lower, and extinct form, is not in any degree new. Lamarck long ago came to this conclusion, which has lately been maintained by several eminent naturalists and philosophers; for instance by Wallace, Huxley, Lyell, Vogt, Lubbock, Büchner, Rolle, &c.,1 and especially by Häckel. This last naturalist, besides his great work, 'Generelle Morphologie ‘(1866), has recently (1868, with a second edit. in 1870), published his ‘Natürliche Schöpfungsgeschichte, ‘in which he fully discusses the genealogy of man. If this work had appeared before my essay had been written, I should probably never have completed it. Almost all the conclusions at which I have arrived I find confirmed by this naturalist, whose knowledge on many points is much fuller than mine. Wherever I have added any fact or view from Prof. Häckel’s writings, I give his authority in the text, other statements I leave as they originally stood in my manuscript, occasionally giving in the foot-notes references to his works, as a confirmation of the more doubtful or interesting points.
During many years it has seemed to me highly probable that sexual selection has played an important part in differentiating the races of man; but in my ‘Origin of Species’ (first edition, p. 199) I contented myself by merely alluding to this belief. When I came to apply this view to man, I found it indispensable to treat the whole subject in full detail.2 Consequently the second part of the present work, treating of sexual selection, has extended to an inordinate length, compared with the first part; but this could not be avoided.
I had intended adding to the present volumes an essay on the expression of the various emotions by man and the lower animals. My attention was called to this subject many years ago by Sir Charles Bell’s admirable work. This illustrious anatomist maintains that man is endowed with certain muscles solely for the sake of expressing his emotions. As this view is obviously opposed to the belief that man is descended from some other and lower form, it was necessary for me to consider it. I likewise wished to ascertain how far the emotions are expressed in the same manner by the different races of man. But owing to the length of the present work, I have thought it better to reserve my essay, which is partially completed, for separate publication.
Part I.
THE DESCENT OR ORIGIN OF MAN.
Part I.—THE DESCENT OF MAN.
CHAPTER I.
The Evidence of the Descent of Man from some Lower Form.
Nature of the evidence bearing on the origin of man—Homologous structures in man and the lower animals—Miscellaneous points of correspondence—Development—Rudimentary structures, muscles, sense-organs, hair, bones, reproductive organs, &c.—The bearing of these three great classes of facts on the origin of man.
He who wishes to decide whether man is the modified descendant of some pre-existing form, would probably first enquire whether man varies, however slightly, in bodily structure and in mental faculties; and if so, whether the variations are transmitted to his offspring in accordance with the laws which prevail with the lower animals; such as that of the transmission of characters to the same age or sex. Again, are the variations the result, as far as our ignorance permits us to judge, of the same general causes, and are they governed by the same general laws, as in the case of other organisms; for instance by correlation, the inherited effects of use and disuse, &c.? Is man subject to similar malconformations, the result of arrested development, of reduplication of parts, &c., and does he display in any of his anomalies reversion to some former and ancient type of structure? It might also naturally be enquired whether man, like so many other animals, has given rise to varieties and sub-races, differing but slightly from each other, or to races differing so much that they must be classed as doubtful species? How are such races distributed over the world; and how, when crossed, do they react on each other, both in the first and succeeding generations? And so with many other points.
The enquirer would next come to the important point, whether man tends to increase at so rapid a rate, as to lead to occasional severe struggles for existence, and consequently to beneficial variations, whether in body or mind, being preserved, and injurious ones eliminated. Do the races or species of men, whichever term may be applied, encroach on and replace each other, so that some finally become extinct? We shall see that all these questions, as indeed is obvious in respect to most of them, must be answered in the affirmative, in the same manner as with the lower animals. But the several considerations just referred to may be conveniently deferred for a time; and we will first see how far the bodily structure of man shows traces, more or less plain, of his descent from some lower form. In the two succeeding chapters the mental powers of man, in comparison with those of the lower animals, will be considered.
The Bodily Structure of Man.—It is notorious that man is constructed on the same general type or model with other mammals. All the bones in his skeleton can be compared with corresponding bones in a monkey, bat, or seal. So it is with his muscles, nerves, blood-vessels and internal viscera. The brain, the most important of all the organs, follows the same law, as shewn by Huxley and other anatomists. Bischoff,3 who is a hostile witness, admits that every chief fissure and fold in the brain
It may, however, be worth while to specify a few points, not directly or obviously connected with structure, by which this correspondence or relationship is well shewn.
Man is liable to receive from the lower animals, and to communicate to them, certain diseases as hydrophobia, variola, the glanders, &c.; and this fact proves the close similarity of their tissues and blood, both in minute structure and composition, far more plainly than does their comparison under the best microscope, or by the aid of the best chemical analysis. Monkeys are liable to many of the same non-contagious diseases as we are; thus Rengger,5 who carefully observed for a long time the Cebus Azaræ in its native land, found it liable to catarrh, with the usual symptoms, and which when often recurrent led to consumption. These monkeys suffered also from apoplexy, inflammation of the bowels, and cataract in the eye. The younger ones when shedding their milk-teeth often died from fever. Medicines produced the same effect on them as on us. Many kinds of monkeys have a strong taste for tea, coffee, and spirituous liquors: they will also, as I have myself seen, smoke tobacco with pleasure. Brehm asserts that the natives of north-eastern Africa catch the wild baboons by exposing vessels with strong beer, by which they are made drunk. He has seen some of these animals, which he kept in confinement, in this state; and he gives a laughable account of their behaviour and strange grimaces. On the following morning they were very cross and dismal; they held their aching heads with both hands and wore a most pitiable expression: when beer or wine was offered them, they turned away with disgust, but relished the juice of lemons.6 An American monkey, an Ateles, after getting drunk on brandy, would never touch it again, and thus was wiser than many men. These trifling facts prove how similar the nerves of taste must be in monkeys and man, and how similarly their whole nervous system is affected.
Man is infested with internal parasites, sometimes causing fatal effects, and is plagued by external parasites, all of which belong to the same genera or families with those infesting other mammals. Man is subject like other mammals, birds, and even insects, to that mysterious law, which causes certain normal processes, such as gestation, as well as the maturation and duration of various diseases, to follow lunar periods.7 His wounds are repaired by the same process of healing; and the stumps left after the amputation of his limbs occasionally possess, especially during an early embryonic period, some power of regeneration, as in the lowest animals.8
The whole process of that most important function, the reproduction of the species, is strikingly the same in all mammals, from the first act of courtship by the male9 to the birth and nurturing of the young. Monkeys are born in almost as helpless a condition as our own infants; and in certain genera the young differ fully as much in appearance from the adults, as do our children from their full-grown parents.10 It has been urged by some writers as an important distinction, that with man the young arrive at maturity at a much later age than with any other animal; but if we look to the races of mankind which inhabit tropical countries the difference is not great, for the orang is believed not to be adult till the age of from ten to fifteen years.11 Man differs from woman in size, bodily strength, hairyness, &c., as well as in mind, in the same manner as do the two sexes of many mammals. It is, in short, scarcely possible to exaggerate the close correspondence in general structure, in the minute structure of the tissues, in chemical composition and in constitution, between man and the higher animals, especially the anthropomorphous apes.
Embryonic Development.—Man is developed from an ovule, about the 125th of an inch in diameter, which differs in no respect from the ovules of other animals. The embryo itself at a very early period can hardly be distinguished from that of other members of the vertebrate kingdom. At this period the arteries run in arch-like branches, as if to carry the blood to branchiæ which are not present in the higher vertebrata, though the slits on the sides of the neck still remain (f, g, fig. 1), marking their former position. At a somewhat later period, when the extremities are developed, “the feet of lizards and mammals,” as the illustrious Von Baer remarks, “the wings and feet of birds, no less than the hands and feet of man, all arise from the same fundamental form.” It is, says Prof. Huxley,12 “quite in the later stages of development that the young human being presents marked differences from the young ape, while the latter departs as much from the dog in its developments, as the man does. Startling as this last assertion may appear to be, it is demonstrably true.”
As some of my readers may never have seen a drawing of an embryo, I have given one of man and another of a dog, at about the same early stage of development, carefully copied from two works of undoubted accuracy.13
Fig. 1. Upper figure human embryo, from Ecker. Lower figure that of a dog, from Bischoff.
a. Fore-brain, cerebral hemispheres, &c. g. Second visceral arch. b. Mid-brain, corpora quadrigemina. H. Vertebral columns and muscles in process of development. c. Hind-brain, cerebellum, medulla oblongata. i. Anterior ┐│ extremities
┘ d. Eye. K. Posterior e. Ear. L. Tail or os coccyx. f. First visceral arch.
After the foregoing statements made by such high authorities, it would be superfluous on my part to give a number of borrowed details, shewing that the embryo of man closely resembles that of other mammals. It may, however, be added that the human embryo likewise resembles in various points of structure certain low forms when adult. For instance, the heart at first exists as a simple pulsating vessel; the excreta are voided through a cloacal passage; and the os coccyx projects like a true tail, “extending considerably beyond the rudimentary legs.”14 In the embryos of all air-breathing vertebrates, certain glands called the corpora Wolffiana, correspond with and act like the kidneys of mature fishes.15 Even at a later embryonic period, some striking resemblances between man and the lower animals may be observed. Bischoff says that the convolutions of the brain in a human fœtus at the end of the seventh month reach about the same stage of development as in a baboon when adult.16 The great toe, as Prof. Owen remarks,17 “which forms the fulcrum when standing or walking, is perhaps the most characteristic peculiarity in the human structure;” but in an embryo, about an inch in length, Prof. Wyman18 found “that the great toe was shorter than the others, and, instead of being parallel to them, projected at an angle from the side of the foot, thus corresponding with the permanent condition of this part in the quadrumana.” I will conclude with a quotation from Huxley,19 who after asking, does man originate in a different way from a dog, bird, frog or fish? says, “the reply is not doubtful for a moment; without question, the mode of origin and the early stages of the development of man are identical with those of the animals immediately below him in the scale: without a doubt in these respects, he is far nearer to apes, than the apes are to the dog.”
Rudiments.—This subject, though not intrinsically more important than the two last, will for several reasons be here treated with more fullness.20 Not one of the higher animals can be named which does not bear some part in a rudimentary condition; and man forms no exception to the rule. Rudimentary organs must be distinguished from those that are nascent; though in some cases the distinction is not easy. The former are either absolutely useless, such as the mammæ of male quadrupeds, or the incisor teeth of ruminants which never cut through the gums; or they are of such slight service to their present possessors, that we cannot suppose that they were developed under the conditions which now exist. Organs in this latter state are not strictly rudimentary, but they are tending in this direction. Nascent organs, on the other hand, though not fully developed, are of high service to their possessors, and are capable of further development. Rudimentary organs are eminently variable; and this is partly intelligible, as they are useless or nearly useless, and consequently are no longer subjected to natural selection. They often become wholly suppressed. When this occurs, they are nevertheless liable to occasional reappearance through reversion; and this is a circumstance well worthy of attention.
Disuse at that period of life, when an organ is chiefly used, and this is generally during maturity, together with inheritance at a corresponding period of life, seem to have been the chief agents in causing organs to become rudimentary. The term “disuse” does not relate merely to the lessened action of muscles, but includes a diminished flow of blood to a part or organ, from being subjected to fewer alternations of pressure, or from becoming in any way less habitually active. Rudiments, however, may occur in one sex of parts normally present in the other sex; and such rudiments, as we shall hereafter see, have often originated in a distinct manner. In some cases organs have been reduced by means of natural selection, from having become injurious to the species under changed habits of life. The process of reduction is probably often aided through the two principles of compensation and economy of growth; but the later stages of reduction, after disuse has done all that can fairly be attributed to it, and when the saving to be effected by the economy of growth would be very small,21 are difficult to understand. The final and complete suppression of a part, already useless and much reduced in size, in which case neither compensation nor economy can come into play, is perhaps intelligible by the aid of the hypothesis of pangenesis, and apparently in no other way. But as the whole subject of rudimentary organs has been fully discussed and illustrated in my former works,22 I need here say no more on this head.
Rudiments of various muscles have been observed in many parts of the human body;23 and not a few muscles, which are regularly present in some of the lower animals can occasionally be
Some few persons have the power of contracting the superficial muscles on their scalps; and these muscles are in a variable and partially rudimentary condition. M. A. de Candolle has communicated to me a curious instance of the long-continued persistence or inheritance of this power, as well as of its unusual development. He knows a family, in which one member, the present head of a family, could, when a youth, pitch several heavy books from his head by the movement of the scalp alone; and he won wagers by performing this feat. His father, uncle, grandfather, and all his three children possess the same power to the same unusual degree. This family became divided eight generations ago into two branches; so that the head of the above-mentioned branch is cousin in the seventh degree to the head of the other branch. This distant cousin resides in another part of France, and on being asked whether he possessed the same faculty, immediately exhibited his power. This case offers a good illustration how persistently an absolutely useless faculty may be transmitted.
The extrinsic muscles which serve to move the whole external ear, and the intrinsic muscles which move the different parts, all of which belong to the system of the panniculus, are in a rudimentary condition in man; they are also variable in development, or at least in function. I have seen one man who could draw his ears forwards, and another who could draw them backwards;25
and from what one of these persons told me, it is probable that most of us by often touching our ears and thus directing our attention towards them, could by repeated trials recover some power of movement. The faculty of erecting the ears and of directing them to different points of the compass, is no doubt of the highest service to many animals, as they thus perceive the point of danger; but I have never heard of a man who possessed the least power of erecting his ears,—the one movement which might be of use to him. The whole external shell of the ear may be considered a rudiment, together with the various folds and prominences (helix and anti-helix, tragus and anti-tragus, &c.) which in the lower animals strengthen and support the ear when erect, without adding much to its weight. Some authors, however, suppose that the cartilage of the shell serves to transmit vibrations to the acoustic nerve; but Mr. Toynbee,26 after collecting all the known evidence on this head, concludes that the external shell is of no distinct use. The ears of the chimpanzee and orang are curiously like those of man, and I am assured by the keepers in the Zoological Gardens that these animals never move or erect them; so that they are in an equally rudimentary condition, as far as function is concerned, as in man. Why these animals, as well as the progenitors of man, should have lost the power of erecting their ears we cannot say. It may be, though I am not quite satisfied with this view, that owing to their arboreal habits and great strength they were but little exposed to danger, and so during a lengthened period moved their ears but little, and thus gradually lost the power of moving them. This would be a parallel case with that of those large and heavy birds, which from inhabiting oceanic islands have not been exposed to the attacks of beasts of prey, and have consequently lost the power of using their wings for flight.