By G. Schwalbe

Professor of Anatomy in the University of Strassburg

The problem of the origin of the human race, of the descent of man, is ranked by Huxley in his epoch-making book Man's Place in Nature, as the deepest with which biology has to concern itself, "the question of questions,"—the problem which underlies all others. In the same brilliant and lucid exposition, which appeared in 1863, soon after the publication of Darwin's Origin of Species, Huxley stated his own views in regard to this great problem. He tells us how the idea of a natural descent of man gradually grew up in his mind. It was especially the assertions of Owen in regard to the total difference between the human and the simian brain that called forth strong dissent from the great anatomist Huxley, and he easily succeeded in showing that Owen's supposed differences had no real existence; he even established, on the basis of his own anatomical investigations, the proposition that the anatomical differences between the Marmoset and the Chimpanzee are much greater than those between the Chimpanzee and Man.

But why do we thus introduce the study of Darwin's Descent of Man, which is to occupy us here, by insisting on the fact that Huxley had taken the field in defence of the descent of man in 1863, while Darwin's book on the subject did not appear till 1871? It is in order that we may clearly understand how it happened that from this time onwards Darwin and Huxley followed the same great aim in the most intimate association.

Huxley and Darwin working at the same Problema maximum! Huxley fiery, impetuous, eager for battle, contemptuous of the resistance of a dull world, or energetically triumphing over it. Darwin calm, weighing every problem slowly, letting it mature thoroughly,—not a fighter, yet having the greater and more lasting influence by virtue of his immense mass of critically sifted proofs. Darwin's friend, Huxley, was the first to do him justice, to understand his nature, and to find in it the reason why the detailed and carefully considered book on the descent of man made its appearance so late. Huxley, always generous, never thought of claiming priority for himself. In enthusiastic language he tells how Darwin's immortal work, The Origin of Species, first shed light for him on the problem of the descent of man; the recognition of a vera causa in the transformation of species illuminated his thoughts as with a flash. He was now content to leave what perplexed him, what he could not yet solve, as he says himself, "in the mighty hands of Darwin." Happy in the bustle of strife against old and deep-rooted prejudices, against intolerance and superstition, he wielded his sharp weapons on Darwin's behalf; wearing Darwin's armour he joyously overthrew adversary after adversary. Darwin spoke of Huxley as his "general agent."[75] Huxley says of himself "I am Darwin's bulldog."[76]

Thus Huxley openly acknowledged that it was Darwin's Origin of Species that first set the problem of the descent of man in its true light, that made the question of the origin of the human race a pressing one. That this was the logical consequence of his book Darwin himself had long felt. He had been reproached with intentionally shirking the application of his theory to Man. Let us hear what he says on this point in his autobiography: "As soon as I had become, in the year 1837 or 1838, convinced that species were mutable productions, I could not avoid the belief that man must come under the same law. Accordingly I collected notes on the subject for my own satisfaction, and not for a long time with any intention of publishing. Although in the 'Origin of Species' the derivation of any particular species is never discussed, yet I thought it best, in order that no honourable man should accuse me of concealing my views,[77] to add that by the work 'light would be thrown on the origin of man and his history.' It would have been useless and injurious to the success of the book to have paraded, without giving any evidence, my conviction with respect to his origin."[78]

In a letter written in January, 1860, to the Rev. L. Blomefield, Darwin expresses himself in similar terms. "With respect to man, I am very far from wishing to obtrude my belief; but I thought it dishonest to quite conceal my opinion."[79]

The brief allusion in the Origin of Species is so far from prominent and so incidental that it was excusable to assume that Darwin had not touched upon the descent of man in this work. It was solely the desire to have his mass of evidence sufficiently complete, solely Darwin's great characteristic of never publishing till he had carefully weighed all aspects of his subject for years, solely, in short, his most fastidious scientific conscience that restrained him from challenging the world in 1859 with a book in which the theory of the descent of man was fully set forth. Three years, frequently interrupted by ill-health, were needed for the actual writing of the book:[80] the first edition, which appeared in 1871, was followed in 1874 by a much improved second edition, the preparation of which he very reluctantly undertook.[81]

This, briefly, is the history of the work, which, with the Origin of Species, marks an epoch in the history of biological sciences—the work with which the cautious, peace-loving investigator ventured forth from his contemplative life into the arena of strife and unrest, and laid himself open to all the annoyances that deep-rooted belief and prejudice, and the prevailing tendency of scientific thought at the time could devise.

Darwin did not take this step lightly. Of great interest in this connection is a letter written to Wallace on Dec. 22, 1857,[82] in which he says, "You ask me whether I shall discuss 'man.' I think I shall avoid the whole subject, as so surrounded with prejudices; though I fully admit that it is the highest and most interesting problem for the naturalist." But his conscientiousness compelled him to state briefly his opinion on the subject in the Origin of Species in 1859. Nevertheless he did not escape reproaches for having been so reticent. This is unmistakably apparent from a letter to Fritz MÜller dated Feb. 22 [1869?], in which he says: "I am thinking of writing a little essay on the Origin of Mankind, as I have been taunted with concealing my opinions."[83]

It might be thought that Darwin behaved thus hesitatingly, and was so slow in deciding on the full publication of his collected material in regard to the descent of man, because he had religious difficulties to overcome.

But this was not the case, as we can see from his admirable confession of faith, the publication of which we owe to his son Francis.[84] Whoever wishes really to understand the lofty character of this great man should read these immortal lines in which he unfolds to us in simple and straightforward words the development of his conception of the universe. He describes how, though he was still quite orthodox during his voyage round the world on board the Beagle, he came gradually to see, shortly afterwards (1836-1839) that the Old Testament was no more to be trusted than the Sacred Books of the Hindoos; the miracles by which Christianity is supported, the discrepancies between the accounts in the different Gospels, gradually led him to disbelieve in Christianity as a divine revelation. "Thus," he writes,[85] "disbelief crept over me at a very slow rate, but was at last complete. The rate was so slow that I felt no distress." But Darwin was too modest to presume to go beyond the limits laid down by science. He wanted nothing more than to be able to go, freely and unhampered by belief in authority or in the Bible, as far as human knowledge could lead him. We learn this from the concluding words of his chapter on religion "The mystery of the beginning of all things is insoluble by us; and I for one must be content to remain an Agnostic."[86]

Darwin was always very unwilling to give publicity to his views in regard to religion. In a letter to Asa Gray on May 22, 1860,[87] he declares that it is always painful to him to have to enter into discussion of religious problems. He had, he said, no intention of writing atheistically.

Finally, let us cite one characteristic sentence from a letter from Darwin to C. Ridley[88] (Nov. 28, 1878). A clergyman, Dr. Pusey, had asserted that Darwin had written the Origin of Species with some relation to theology. Darwin writes emphatically, "Many years ago when I was collecting facts for the 'Origin,' my belief in what is called a personal God was as firm as that of Dr. Pusey himself, and as to the eternity of matter I never troubled myself about such insoluble questions." The expression "many years ago" refers to the time of his voyage round the world, as has already been pointed out. Darwin means by this utterance that the views which had gradually developed in his mind in regard to the origin of species were quite compatible with the faith of the Church.

If we consider all these utterances of Darwin in regard to religion and to his outlook on life (Weltanschauung), we shall see at least so much, that religious reflection could in no way have influenced him in regard to the writing and publishing of his book on The Descent of Man. Darwin had early won for himself freedom of thought, and to this freedom he remained true to the end of his life, uninfluenced by the customs and opinions of the world around him.

Darwin was thus inwardly fortified and armed against the host of calumnies, accusations, and attacks called forth by the publication of the Origin of Species, and to an even greater extent by the appearance of the Descent of Man. But in his defence he could rely on the aid of a band of distinguished auxiliaries of the rarest ability. His faithful confederate, Huxley, was joined by the botanist Hooker, and, after longer resistance, by the famous geologist Lyell, whose "conversion" afforded Darwin peculiar satisfaction. All three took the field with enthusiasm in defence of the natural descent of man. From Wallace, on the other hand, though he shared with him the idea of natural selection, Darwin got no support in this matter. Wallace expressed himself in a strange manner. He admitted everything in regard to the morphological descent of man, but maintained, in a mystic way, that something else, something of a spiritual nature must have been added to what man inherited from his animal ancestors. Darwin, whose esteem for Wallace was extraordinarily high, could not understand how he could give utterance to such a mystical view in regard to man; the idea seemed to him so "incredibly strange" that he thought some one else must have added these sentences to Wallace's paper.

Even now there are thinkers who, like Wallace, shrink from applying to man the ultimate consequences of the theory of descent. The idea that man is derived from ape-like forms is to them unpleasant and humiliating.

So far I have been depicting the development of Darwin's work on the descent of man. In what follows I shall endeavour to give a condensed survey of the contents of the book.

It must at once be said that the contents of Darwin's work fall into two parts, dealing with entirely different subjects. The Descent of Man includes a very detailed investigation in regard to secondary sexual characters in the animal series, and on this investigation Darwin founded a new theory, that of sexual selection. With astonishing patience he gathered together an immense mass of material, and showed, in regard to Arthropods and Vertebrates, the wide distribution of secondary characters, which develop almost exclusively in the male, and which enable him, on the one hand, to get the better of his rivals in the struggle for the female by the greater perfection of his weapons, and, on the other hand, to offer greater allurements to the female through the higher development of decorative characters, of song, or of scent-producing glands. The best equipped males will thus crowd out the less well-equipped in the matter of reproduction, and thus the relevant characters will be increased and perfected through sexual selection. It is, of course, a necessary assumption that these secondary sexual characters may be transmitted to the female, although perhaps in rudimentary form.

As we have said, this story of sexual selection takes up a great deal of space in Darwin's book, and it need only be considered here in so far as Darwin applied it to the descent of man. To this latter problem the whole of Part I is devoted, while Part III contains a discussion of sexual selection in relation to man, and a general summary. Part II treats of sexual selection in general, and may be disregarded in our present study. Moreover, many interesting details must necessarily be passed over in what follows, for want of space.

The first part of the Descent of Man begins with an enumeration of the proofs of the animal descent of man taken from the structure of the human body. Darwin chiefly emphasises the fact that the human body consists of the same organs and of the same tissues as those of the other mammals; he shows also that man is subject to the same diseases and tormented by the same parasites as the apes. He further dwells on the general agreement exhibited by young embryonic forms, and he illustrates this by two figures placed one above the other, one representing a human embryo, after Ecker, the other a dog embryo, after Bischoff.[89]

Darwin finds further proofs of the animal origin of man in the reduced structures, in themselves extremely variable, which are either absolutely useless to their possessors, or of so little use that they could never have developed under existing conditions. Of such vestiges he enumerates: the defective development of the panniculus carnosus (muscle of the skin) so widely distributed among mammals, the ear-muscles, the occasional persistence of the animal ear-point in man, the rudimentary nictitating membrane (plica semilunaris) in the human eye, the slight development of the organ of smell, the general hairiness of the human body, the frequently defective development or entire absence of the third molar (the wisdom tooth), the vermiform appendix, the occasional reappearance of a bony canal (foramen supracondyloideum) at the lower end of the humerus, the rudimentary tail of man (the so-called taillessness), and so on. Of these rudimentary structures the occasional occurrence of the animal ear-point in man is most fully discussed. Darwin's attention was called to this interesting structure by the sculptor Woolner. He figures such a case observed in man, and also the head of an alleged orang-foetus, the photograph of which he received from Nitsche.

Darwin's interpretation of Woolner's case as having arisen through a folding over of the free edge of a pointed ear has been fully borne out by my investigations on the external ear.[90] In particular, it was established by these investigations that the human foetus, about the middle of its embryonic life, possesses a pointed ear somewhat similar to that of the monkey genus Macacus. One of Darwin's statements in regard to the head of the orang-foetus must be corrected. A large ear with a point is shown in the photograph,[91] but it can easily be demonstrated—and Deniker has already pointed this out—that the figure is not that of an orang foetus at all, for that form has much smaller ears with no point; nor can it be a gibbon-foetus, as Deniker supposes, for the gibbon ear is also without a point. I myself regard it as that of a Macacus-embryo. But this mistake, which is due to Nitsche, in no way affects the fact recognised by Darwin, that ear-forms showing the point characteristic of the animal ear occur in man with extraordinary frequency.

Finally, there is a discussion of those rudimentary structures which occur only in one sex, such as the rudimentary mammary glands in the male, the vesicula prostatica, which corresponds to the uterus of the female, and others. All these facts tell in favour of the common descent of man and all other vertebrates. The conclusion of this section is characteristic: "It is only our natural prejudice, and that arrogance which made our forefathers declare that they were descended from demi-gods, which leads us to demur to this conclusion. But the time will before long come, when it will be thought wonderful that naturalists, who were well acquainted with the comparative structure and development of man, and other mammals, should have believed that each was the work of a separate act of creation."[92]

In the second chapter there is a more detailed discussion, again based upon an extraordinary wealth of facts, of the problem as to the manner in which, and the causes through which, man evolved from a lower form. Precisely the same causes are here suggested for the origin of man, as for the origin of species in general. Variability, which is a necessary assumption in regard to all transformations, occurs in man to a high degree. Moreover, the rapid multiplication of the human race creates conditions which necessitate an energetic struggle for existence, and thus afford scope for the intervention of natural selection. Of the exercise of artificial selection in the human race, there is nothing to be said, unless we cite such cases as the grenadiers of Frederick William I, or the population of ancient Sparta. In the passages already referred to and in those which follow, the transmission of acquired characters, upon which Darwin does not dwell, is taken for granted. In man, direct effects of changed conditions can be demonstrated (for instance in regard to bodily size), and there are also proofs of the influence exerted on his physical constitution by increased use or disuse. Reference is here made to the fact, established by Forbes, that the Quechua Indians of the high plateaus of Peru show a striking development of lungs and thorax, as a result of living constantly at high altitudes.

Such special forms of variation as arrests of development (microcephalism) and reversion to lower forms are next discussed. Darwin himself felt[93] that these subjects are so nearly related to the cases mentioned in the first chapter, that many of them might as well have been dealt with there. It seems to me that it would have been better so, for the citation of additional instances of reversion at this place rather disturbs the logical sequence of his ideas as to the conditions which have brought about the evolution of man from lower forms. The instances of reversion here discussed are microcephalism, which Darwin wrongly interpreted as atavistic, supernumerary mammae, supernumerary digits, bicornuate uterus, the development of abnormal muscles, and so on. Brief mention is also made of correlative variations observed in man.

Darwin next discusses the question as to the manner in which man attained to the erect position from the state of a climbing quadruped. Here again he puts the influence of Natural Selection in the first rank. The immediate progenitors of man had to maintain a struggle for existence in which success was to the more intelligent, and to those with social instincts. The hand of these climbing ancestors, which had little skill and served mainly for locomotion, could only undergo further development when some early member of the Primate series came to live more on the ground and less among trees.

A bipedal existence thus became possible, and with it the liberation of the hand from locomotion, and the one-sided development of the human foot. The upright position brought about correlated variations in the bodily structure; with the free use of the hand it became possible to manufacture weapons and to use them; and this again resulted in a degeneration of the powerful canine teeth and the jaws, which were then no longer necessary for defence. Above all, however, the intelligence immediately increased, and with it skull and brain. The nakedness of man, and the absence of a tail (rudimentariness of the tail vertebrae) are next discussed. Darwin is inclined to attribute the nakedness of man, not to the action of natural selection on ancestors who originally inhabited a tropical land, but to sexual selection, which, for aesthetic reasons, brought about the loss of the hairy covering in man, or primarily in woman. An interesting discussion of the loss of the tail, which, however, man shares with the anthropoid apes, some other monkeys and lemurs, forms the conclusion of the almost superabundant material which Darwin worked up in the second chapter. His object was to show that some of the most distinctive human characters are in all probability directly or indirectly due to natural selection. With characteristic modesty he adds:[94] "Hence, if I have erred in giving to natural selection great power, which I am very far from admitting, or in having exaggerated its power, which is in itself probable, I have at least, as I hope, done good service in aiding to overthrow the dogma of separate creations." At the end of the chapter he touches upon the objection as to man's helpless and defenceless condition. Against this he urges his intelligence and social instincts.

The two following chapters contain a detailed discussion of the objections drawn from the supposed great differences between the mental powers of men and animals. Darwin at once admits that the differences are enormous, but not that any fundamental difference between the two can be found. Very characteristic of him is the following passage: "In what manner the mental powers were first developed in the lowest organisms, is as hopeless an enquiry as how life itself first originated. These are problems for the distant future, if they are ever to be solved by man."[95]

After some brief observations on instinct and intelligence, Darwin brings forward evidence to show that the greater number of the emotional states, such as pleasure and pain, happiness and misery, love and hate are common to man and the higher animals. He goes on to give various examples showing that wonder and curiosity, imitation, attention, memory and imagination (dreams of animals), can also be observed in the higher mammals, especially in apes. In regard even to reason there are no sharply defined limits. A certain faculty of deliberation is characteristic of some animals, and the more thoroughly we know an animal the more intelligence we are inclined to credit it with. Examples are brought forward of the intelligent and deliberate actions of apes, dogs and elephants. But although no sharply defined differences exist between man and animals, there is, nevertheless, a series of other mental powers which are characteristics usually regarded as absolutely peculiar to man. Some of these characteristics are examined in detail, and it is shown that the arguments drawn from them are not conclusive. Man alone is said to be capable of progressive improvement; but against this must be placed as something analogous in animals, the fact that they learn cunning and caution through long continued persecution. Even the use of tools is not in itself peculiar to man (monkeys use sticks, stones and twigs), but man alone fashions and uses implements designed for a special purpose. In this connection the remarks taken from Lubbock in regard to the origin and gradual development of the earliest flint implements will be read with interest; these are similar to the observations on modern eoliths, and their bearing on the development of the stone industry. It is interesting to learn from a letter to Hooker,[96] that Darwin himself at first doubted whether the stone implements discovered by Boucher de Perthes were really of the nature of tools. With the relentless candour as to himself which characterised him, he writes four years later in a letter to Lyell in regard to this view of Boucher de Perthes' discoveries: "I know something about his errors, and looked at his book many years ago, and am ashamed to think that I concluded the whole was rubbish! Yet he has done for man something like what Agassiz did for glaciers."[97]

To return to Darwin's further comparisons between the higher mental powers of man and animals; He takes much of the force from the argument that man alone is capable of abstraction and self-consciousness by his own observations on dogs. One of the main differences between man and animals, speech, receives detailed treatment. He points out that various animals (birds, monkeys, dogs) have a large number of different sounds for different emotions, that, further, man produces in common with animals a whole series of inarticulate cries combined with gestures, and that dogs learn to understand whole sentences of human speech. In regard to human language, Darwin expresses a view contrary to that held by Max MÜller:[98] "I cannot doubt that language owes its origin to the imitation and modification of various natural sounds, the voices of other animals, and man's own instinctive cries, aided by signs and gestures." The development of actual language presupposes a higher degree of intelligence than is found in any kind of ape. Darwin remarks on this point:[99] "The fact of the higher apes not using their vocal organs for speech no doubt depends on their intelligence not having been sufficiently advanced."

The sense of beauty, too, has been alleged to be peculiar to man. In refutation of this assertion Darwin points to the decorative colours of birds, which are used for display. And to the last objection, that man alone has religion, that he alone has a belief in God, it is answered "that numerous races have existed, and still exist, who have no idea of one or more gods, and who have no words in their languages to express such an idea."[100]

The result of the investigations recorded in this chapter is to show that, great as the difference in mental powers between man and the higher animals may be, it is undoubtedly only a difference "of degree and not of kind."[101]

In the fourth chapter Darwin deals with the moral sense or conscience, which is the most important of all differences between man and animals. It is a result of social instincts, which lead to sympathy for other members of the same society, to non-egoistic actions for the good of others. Darwin shows that social tendencies are found among many animals, and that among these love and kin-sympathy exist, and he gives examples of animals (especially dogs) which may exhibit characters that we should call moral in man (e.g. disinterested self-sacrifice for the sake of others). The early ape-like progenitors of the human race were undoubtedly social. With the increase of intelligence the moral sense develops farther; with the acquisition of speech public opinion arises, and finally, moral sense becomes habit. The rest of Darwin's detailed discussions on moral philosophy may be passed over.

The fifth chapter may be very briefly summarised. In it Darwin shows that the intellectual and moral faculties are perfected through natural selection. He inquires how it can come about that a tribe at a low level of evolution attains to a higher, although the best and bravest among them often pay for their fidelity and courage with their lives without leaving any descendants. In this case it is the sentiment of glory, praise and blame, the admiration of others, which bring about the increase of the better members of the tribe. Property, fixed dwellings, and the association of families into a community are also indispensable requirements for civilisation. In the longer second section of the fifth chapter Darwin acts mainly as recorder. On the basis of numerous investigations, especially those of Greg, Wallace, and Galton, he inquires how far the influence of natural selection can be demonstrated in regard to civilised nations. In the final section, which deals with the proofs that all civilised nations were once barbarians, Darwin again uses the results gained by other investigators, such as Lubbock and Tylor. There are two sets of facts which prove the proposition in question. In the first place, we find traces of a former lower state in the customs and beliefs of all civilised nations, and in the second place, there are proofs to show that savage races are independently able to raise themselves a few steps in the scale of civilisation, and that they have thus raised themselves.

In the sixth chapter of the work, Morphology comes into the foreground once more. Darwin first goes back, however, to the argument based on the great difference between the mental powers of the highest animals and those of man. That this is only quantitative, not qualitative, he has already shown. Very instructive in this connection is the reference to the enormous difference in mental powers in another class. No one would draw from the fact that the cochineal insect (Coccus) and the ant exhibit enormous differences in their mental powers, the conclusion that the ant should therefore be regarded as something quite distinct, and withdrawn from the class of insects altogether.

Darwin next attempts to establish the specific genealogical tree of man, and carefully weighs the differences and resemblances between the different families of the Primates. The erect position of man is an adaptive character, just as are the various characters referable to aquatic life in the seals, which, notwithstanding these, are ranked as a mere family of the carnivores. The following utterance is very characteristic of Darwin:[102] "If man had not been his own classifier, he would never have thought of founding a separate order for his own reception." In numerous characters not mentioned in systematic works, in the features of the face, in the form of the nose, in the structure of the external ear, man resembles the apes. The arrangement of the hair in man has also much in common with the apes; as also the occurrence of hair on the forehead of the human embryo, the beard, the convergence of the hair of the upper and under arm towards the elbow, which occurs not only in the anthropoid apes, but also in some American monkeys. Darwin here adopts Wallace's explanation of the origin of the ascending direction of the hair in the forearm of the orang,—that it has arisen through the habit of holding the hands over the head in rain. But this explanation cannot be maintained when we consider that this disposition of the hair is widely distributed among the most different mammals, being found in the dog, in the sloth, and in many of the lower monkeys.

After further careful analysis of the anatomical characters Darwin reaches the conclusion that the New World monkeys (Platyrrhine) may be excluded from the genealogical tree altogether, but that man is an offshoot from the Old World monkeys (Catarrhine) whose progenitors existed as far back as the Miocene period. Among these Old World monkeys the forms to which man shows the greatest resemblance are the anthropoid apes, which, like him, possess neither tail nor ischial callosities. The platyrrhine and catarrhine monkeys have their primitive ancestor among extinct forms of the Lemuridae. Darwin also touches on the question of the original home of the human race and supposes that it may have been in Africa, because it is there that man's nearest relatives, the gorilla and the chimpanzee, are found. But he regards speculation on this point as useless. It is remarkable that, in this connection, Darwin regards the loss of the hair-covering in man as having some relation to a warm climate, while elsewhere he is inclined to make sexual selection responsible for it. Darwin recognises the great gap between man and his nearest relatives, but similar gaps exist at other parts of the mammalian genealogical tree: the allied forms have become extinct. After the extermination of the lower races of mankind, on the one hand, and of the anthropoid apes on the other, which will undoubtedly take place, the gulf will be greater than ever, since the baboons will then bound it on the one side, and the white races on the other. Little weight need be attached to the lack of fossil remains to fill up this gap, since the discovery of these depends upon chance. The last part of the chapter is devoted to a discussion of the earlier stages in the genealogy of man. Here Darwin accepts in the main the genealogical tree, which had meantime been published by Haeckel, who traces the pedigree back through Monotrems, Reptiles, Amphibians, and Fishes, to Amphioxus.

Then follows an attempt to reconstruct, from the atavistic characters, a picture of our primitive ancestor who was undoubtedly an arboreal animal. The occurrence of rudiments of parts in one sex which only come to full development in the other is next discussed. This state of things Darwin regards as derived from an original hermaphroditism. In regard to the mammary glands of the male he does not accept the theory that they are vestigial, but considers them rather as not fully developed.

The last chapter of Part I deals with the question whether the different races of man are to be regarded as different species, or as sub-species of a race of monophyletic origin. The striking differences between the races are first emphasised, and the question of the fertility or infertility of hybrids is discussed. That fertility is the more usual is shown by the excessive fertility of the hybrid population of Brazil. This, and the great variability of the distinguishing characters of the different races, as well as the fact that all grades of transition stages are found between these, while considerable general agreement exists, tell in favour of the unity of the races and lead to the conclusion that they all had a common primitive ancestor.

Darwin therefore classifies all the different races as sub-species of one and the same species. Then follows an interesting inquiry into the reasons for the extinction of human races. He recognises as the ultimate reason the injurious effects of a change of the conditions of life, which may bring about an increase in infantile mortality, and a diminished fertility. It is precisely the reproductive system, among animals also, which is most susceptible to changes in the environment.

The final section of this chapter deals with the formation of the races of mankind. Darwin discusses the question how far the direct effect of different conditions of life, or the inherited effects of increased use or disuse may have brought about the characteristic differences between the different races. Even in regard to the origin of the colour of the skin he rejects the transmitted effects of an original difference of climate as an explanation. In so doing he is following his tendency to exclude Lamarckian explanations as far as possible. But here he makes gratuitous difficulties from which, since natural selection fails, there is no escape except by bringing in the principle of sexual selection, to which, he regarded it as possible, skin-colouring, arrangement of hair, and form of features might be traced. But with his characteristic conscientiousness he guards himself thus: "I do not intend to assert that sexual selection will account for all the differences between the races."[103]

I may be permitted a remark as to Darwin's attitude towards Lamarck. While, at an earlier stage, when he was engaged in the preliminary labours for his immortal work, The Origin of Species, Darwin expresses himself very forcibly against the views of Lamarck, speaking of Lamarckian "nonsense,"[104] and of Lamarck's "absurd, though clever work"[105] and expressly declaring, "I attribute very little to the direct action of climate, etc."[106] yet in later life he became more and more convinced of the influence of external conditions. In 1876, that is, two years after the appearance of the second edition of The Descent of Man, he writes with his usual candid honesty: "In my opinion the greatest error which I have committed, has been not allowing sufficient weight to the direct action of the environment, i.e. food, climate, etc. independently of a natural selection."[107] It is certain from this change of opinion that, if he had been able to make up his mind to issue a third edition of The Descent of Man, he would have ascribed a much greater influence to the effect of external conditions in explaining the different characters of the races of man than he did in the second edition. He would also undoubtedly have attributed less influence to sexual selection as a factor in the origin of the different bodily characteristics, if indeed he would not have excluded it altogether.

In Part III of the Descent two additional chapters are devoted to the discussion of sexual selection in relation to man. These may be very briefly referred to. Darwin here seeks to show that sexual selection has been operative on man and his primitive progenitor. Space fails me to follow out his interesting arguments. I can only mention that he is inclined to trace back hairlessness, the development of the beard in man, and the characteristic colour of the different human races to sexual selection. Since bareness of the skin could be no advantage, but rather a disadvantage, this character cannot have been brought about by natural selection. Darwin also rejected a direct influence of climate as a cause of the origin of the skin-colour. I have already expressed the opinion, based on the development of his views as shown in his letters, that in a third edition Darwin would probably have laid more stress on the influence of external environment. He himself feels that there are gaps in his proofs here, and says in self-criticism: "The views here advanced, on the part which sexual selection has played in the history of man, want scientific precision."[108] I need here only point out that it is impossible to explain the graduated stages of skin-colour by sexual selection, since it would have produced races sharply defined by their colour and not united to other races by transition stages, and this, it is well known, is not the case. Moreover, the fact established by me,[109] that in all races the ventral side of the trunk is paler than the dorsal side, and the inner surface of the extremities paler than the outer side, cannot be explained by sexual selection in the Darwinian sense.

With this I conclude my brief survey of the rich contents of Darwin's book. I may be permitted to conclude by quoting the magnificent final words of The Descent of Man: "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."[110]

What has been the fate of Darwin's doctrines since his great achievement? How have they been received and followed up by the scientific and lay world? And what do the successors of the mighty hero and genius think now in regard to the origin of the human race?

At the present time we are incomparably more favourably placed than Darwin was for answering this question of all questions. We have at our command an incomparably greater wealth of material than he had at his disposal. And we are more fortunate than he in this respect, that we now know transition-forms which help to fill up the gap, still great, between the lowest human races and the highest apes. Let us consider for a little the more essential additions to our knowledge since the publication of The Descent of Man.

Since that time our knowledge of animal embryos has increased enormously. While Darwin was obliged to content himself with comparing a human embryo with that of a dog, there are now available the youngest embryos of monkeys of all possible groups (Orang, Gibbon, Semnopithecus, Macacus), thanks to Selenka's most successful tour in the East Indies in search of such material. We can now compare corresponding stages of the lower monkeys and of the Anthropoid apes with human embryos, and convince ourselves of their great resemblance to one another, thus strengthening enormously the armour prepared by Darwin in defence of his view on man's nearest relatives. It may be said that Selenka's material fills up the blanks in Darwin's array of proofs in the most satisfactory manner.

The deepening of our knowledge of comparative anatomy also gives us much surer foundations than those on which Darwin was obliged to build. Just of late there have been many workers in the domain of the anatomy of apes and lemurs, and their investigations extend to the most different organs. Our knowledge of fossil apes and lemurs has also become much wider and more exact since Darwin's time: the fossil lemurs have been especially worked up by Cope, Forsyth Major, Ameghino, and others. Darwin knew very little about fossil monkeys. He mentions two or three anthropoid apes as occurring in the Miocene of Europe,[111] but only names Dryopithecus, the largest form from the Miocene of France. It was erroneously supposed that this form was related to Hylobates. We now know not only a form that actually stands near to the gibbon (Pliopithecus), and remains of other anthropoids (Pliohylobates and the fossil chimpanzee, Palaeopithecus), but also several lower catarrhine monkeys, of which Mesopithecus, a form nearly related to the modern Sacred Monkeys (a species of Semnopithecus) and found in strata of the Miocene period in Greece, is the most important. Quite recently, too, Ameghino's investigations have made us acquainted with fossil monkeys from South America (Anthropops, Homunculus), which, according to their discoverer, are to be regarded as in the line of human descent.

What Darwin missed most of all—intermediate forms between apes and man—has been recently furnished. E. Dubois, as is well known, discovered in 1893, near Trinil in Java, in the alluvial deposits of the river Bengawan, an important form represented by a skull-cap, some molars, and a femur. His opinion—much disputed as it has been—that in this form, which he named Pithecanthropus, he has found a long-desired transition-form is shared by the present writer. And although the geological age of these fossils, which, according to Dubois, belong to the uppermost Tertiary series, the Pliocene has recently been fixed at a later date (the older Diluvium), the morphological value of these interesting remains, that is, the intermediate position of Pithecanthropus, still holds good. Volz says with justice,[112] that even if Pithecanthropus is not the missing link, it is undoubtedly a missing link.

As on the one hand there has been found in Pithecanthropus a form which, though intermediate between apes and man, is nevertheless more closely allied to the apes, so on the other hand, much progress has been made since Darwin's day in the discovery and description of the oldest human remains. Since the famous roof of a skull and the bones of the extremities belonging to it were found in 1856 in the Neandertal near DÜsseldorf, the most varied judgments have been expressed in regard to the significance of the remains and of the skull in particular. In Darwin's Descent of Man there is only a passing allusion to them[113] in connection with the discussion of the skull-capacity, although the investigations of Schaaffhausen, King, and Huxley were then known. I believe I have shown, in a series of papers, that the skull in question belongs to a form different from any of the races of man now living, and, with King and Cope, I regard it as at least a different species from living man, and have therefore designated it Homo primigenius. The form unquestionably belongs to the older Diluvium, and in the later Diluvium human forms already appear, which agree in all essential points with existing human races.

As far back as 1886 the value of the Neandertal skull was greatly enhanced by Fraipont's discovery of two skulls and skeletons from Spy in Belgium. These are excellently described by their discoverer,[114] and are regarded as belonging to the same group of forms as the Neandertal remains. In 1899 and the following years came the discovery by Gorjanovic-Kramberger of different skeletal parts of at least ten individuals in a cave near Krapina in Croatia.[115] It is in particular the form of the lower jaw which is different from that of all recent races of man, and which clearly indicates the lowly position of Homo primigenius, while, on the other hand, the long-known skull from Gibraltar, which I[116] have referred to Homo primigenius, and which has lately been examined in detail by Sollas,[117] has made us acquainted with the surprising shape of the eye-orbit, of the nose, and of the whole upper part of the face. Isolated lower jaws found at La Naulette in Belgium, and at Malarnaud in France, increase our material which is now as abundant as could be desired. The most recent discovery of all is that of a skull dug up in August of this year [1908] by Klaatsch and Hauser in the lower grotto of the Le Moustier in Southern France, but this skull has not yet been fully described. Thus Homo primigenius must also be regarded as occupying a position in the gap existing between the highest apes and the lowest human races, Pithecanthropus, standing in the lower part of it, and Homo primigenius in the higher, near man. In order to prevent misunderstanding, I should like here to emphasise that in arranging this structural series—anthropoid apes, Pithecanthropus, Homo primigenius, Homo sapiens—I have no intention of establishing it as a direct genealogical series. I shall have something to say in regard to the genetic relations of these forms, one to another, when discussing the different theories of descent current at the present day.[118]

In quite a different domain from that of morphological relationship, namely in the physiological study of the blood, results have recently been gained which are of the highest importance to the doctrine of descent. Uhlenhuth, Nuttall, and others have established the fact that the blood-serum of a rabbit which has previously had human blood injected into it, forms a precipitate with human blood. This biological reaction was tried with a great variety of mammalian species, and it was found that those far removed from man gave no precipitate under these conditions. But as in other cases among mammals all nearly related forms yield an almost equally marked precipitate, so the serum of a rabbit treated with human blood and then added to the blood of an anthropoid ape gives almost as marked a precipitate as in human blood; the reaction to the blood of the lower Eastern monkeys is weaker, that to the Western monkeys weaker still; indeed in this last case there is only a slight clouding after a considerable time and no actual precipitate. The blood of the Lemuridae (Nuttall) gives no reaction or an extremely weak one, that of the other mammals none whatever. We have in this not only a proof of the literal blood relationship between man and apes, but the degree of relationship with the different main groups of apes can be determined beyond possibility of mistake.

Finally, it must be briefly mentioned that in regard to remains of human handicraft also, the material at our disposal has greatly increased of late years, that, as a result of this, the opinions of archaeologists have undergone many changes, and that, in particular, their views in regard to the age of the human race have been greatly influenced. There is a tendency at the present time to refer the origin of man back to Tertiary times. It is true that no remains of Tertiary man have been found, but flints have been discovered which, according to the opinion of most investigators, bear traces either of use, or of very primitive workmanship. Since Rutot's time, following Mortillet's example, investigators have called these "eoliths," and they have been traced back by Verworn to the Miocene of the Auvergne, and by Rutot even to the upper Oligocene. Although these eoliths are even nowadays the subject of many different views, the preoccupation with them has kept the problem of the age of the human race continually before us.

Geology, too, has made great progress since the days of Darwin and Lyell, and has endeavoured with satisfactory results to arrange the human remains of the Diluvial period in chronological order (Penck). I do not intend to enter upon the question of the primitive home of the human race; since the space at my disposal will not allow of my touching even very briefly upon all the departments of science which are concerned in the problem of the descent of man. How Darwin would have rejoiced over each of the discoveries here briefly outlined! What use he would have made of the new and precious material, which would have prevented the discouragement from which he suffered when preparing the second edition of The Descent of Man! But it was not granted to him to see this progress towards filling up the gaps in his edifice of which he was so painfully conscious.

He did, however, have the satisfaction of seeing his ideas steadily gaining ground, notwithstanding much hostility and deep-rooted prejudice. Even in the years between the appearance of The Origin of Species and of the first edition of the Descent, the idea of a natural descent of man, which was only briefly indicated in the work of 1859, had been eagerly welcomed in some quarters. It has been already pointed out how brilliantly Huxley contributed to the defence and diffusion of Darwin's doctrines, and how in Man's Place in Nature he has given us a classic work as a foundation for the doctrine of the descent of man. As Huxley was Darwin's champion in England, so in Germany Carl Vogt, in particular, made himself master of the Darwinian ideas. But above all it was Haeckel who, in energy, eagerness for battle, and knowledge may be placed side by side with Huxley, who took over the leadership in the controversy over the new conception of the universe. As far back as 1866, in his Generelle Morphologie, he had inquired minutely into the question of the descent of man, and not content with urging merely the general theory of descent from lower animal forms, he drew up for the first time genealogical trees showing the close structural relationships of the different animal groups; the last of these illustrated the relationships of Mammals, and among them of all groups of the Primates, including man. It was Haeckel's genealogical trees that formed the basis of the special discussion of the relationships of man, in the sixth chapter of Darwin's Descent of Man.

In the last section of this essay I shall return to Haeckel's conception of the special descent of man, the main features of which he still upholds, and rightly so. Haeckel has contributed more than any one else to the spread of the Darwinian doctrine.

I can only allow myself a few words as to the spread of the theory of the natural descent of man in other countries. The Parisian anthropological school, founded and guided by the genius of Broca, took up the idea of the descent of man, and made many notable contributions to it (Broca, Manouvrier, Mahoudeau, Deniker and others). In England itself Darwin's work did not die. Huxley took care of that, for he, with his lofty and unprejudiced mind, dominated and inspired English biology until his death on June 29, 1895. He had the satisfaction shortly before his death of learning of Dubois' discovery, which he illustrated by a humourous sketch.[119] But there are still many followers in Darwin's footsteps in England. Keane has worked at the special genealogical tree of the Primates; Keith has inquired which of the anthropoid apes has the greatest number of characters in common with man; Morris concerns himself with the evolution of man in general, especially with his acquisition of the erect position. The recent discoveries of Pithecanthropus and Homo primigenius are being vigorously discussed; but the present writer is not in a position to form an opinion of the extent to which the idea of descent has penetrated throughout England generally.

In Italy independent work in the domain of the descent of man is being produced, especially by Morselli; with him are associated, in the investigation of related problems, Sergi and Giuffrida-Ruggeri. From the ranks of American investigators we may single out in particular the eminent geologist Cope, who championed with much decision the idea of the specific difference of Homo neandertalensis (primigenius) and maintained a more direct descent of man from the fossil Lemuridae. In South America too, in Argentina, new life is stirring in this department of science. Ameghino in Buenos Ayres has awakened the fossil primates of the Pampas formation to new life; he even believes that in his Tetraprothomo, represented by a femur, he has discovered a direct ancestor of man. Lehmann-Nitsche is working at the other side of the gulf between apes and man, and he describes a remarkable first cervical vertebra (atlas) from Monte Hermoso as belonging to a form which may bear the same relation to Homo sapiens in South America as Homo primigenius does in the Old World. After a minute investigation he establishes a human species Homo neogaeus, while Ameghino ascribes this atlas vertebra to his Tetraprothomo.

Thus throughout the whole scientific world there is arising a new life, an eager endeavour to get nearer to Huxley's problema maximum, to penetrate more deeply into the origin of the human race. There are to-day very few experts in anatomy and zoology who deny the animal descent of man in general. Religious considerations, old prejudices, the reluctance to accept man, who so far surpasses mentally all other creatures, as descended from "soulless" animals, prevent a few investigators from giving full adherence to the doctrine. But there are very few of these who still postulate a special act of creation for man. Although the majority of experts in anatomy and zoology accept unconditionally the descent of man from lower forms, there is much diversity of opinion among them in regard to the special line of descent.

In trying to establish any special hypothesis of descent, whether by the graphic method of drawing up genealogical trees or otherwise, let us always bear in mind Darwin's words[120] and use them as a critical guiding line: "As we have no record of the lines of descent, the pedigree can be discovered only by observing the degrees of resemblance between the beings which are to be classed." Darwin carries this further by stating "that resemblances in several unimportant structures, in useless and rudimentary organs, or not now functionally active, or in an embryological condition, are by far the most serviceable for classification."[121] It has also to be remembered that numerous separate points of agreement are of much greater importance than the amount of similarity or dissimilarity in a few points.

The hypotheses as to descent current at the present day may be divided into two main groups. The first group seeks for the roots of the human race not among any of the families of the apes—the anatomically nearest forms—nor among their very similar but less specialised ancestral forms, the fossil representatives of which we can know only in part, but, setting the monkeys on one side, it seeks for them lower down among the fossil Eocene Pseudo-lemuridae or Lemuridae (Cope), or even among the primitive pentadactylous Eocene forms, which may either have led directly to the evolution of man (Adloff), or have given rise to an ancestral form common to apes and men (Klaatsch,[122] Giuffrida-Ruggeri). The common ancestral form, from which man and apes are thus supposed to have arisen independently, may explain the numerous resemblances which actually exist between them. That is to say, all the characters upon which the great structural resemblance between apes and man depends must have been present in their common ancestor. Let us take an example of such a common character. The bony external ear-passage is in general as highly developed in the lower Eastern monkeys and the anthropoid apes as in man. This character must, therefore, have already been present in the common primitive form. In that case it is not easy to understand why the Western monkeys have not also inherited the character, instead of possessing only a tympanic ring. But it becomes more intelligible if we assume that forms with a primitive tympanic ring were the original type, and that from these were evolved, on the one hand, the existing New World monkeys with persistent tympanic ring, and on the other an ancestral form common to the lower Old World monkeys, the anthropoid apes and man. For man shares with these the character in question, and it is also one of the "unimportant" characters required by Darwin. Thus we have two divergent lines arising from the ancestral form, the Western monkeys (Platyrrhine) on the one hand, and an ancestral form common to the lower Eastern monkeys, the anthropoid apes, and man, on the other. But considerations similar to those which showed it to be impossible that man should have developed from an ancestor common to him and the monkeys, yet outside of and parallel with these, may be urged also against the likelihood of a parallel evolution of the lower Eastern monkeys, the anthropoid apes, and man. The anthropoid apes have in common with man many characters which are not present in the lower Old World monkeys. These characters must therefore have been present in the ancestral form common to the three groups. But here, again, it is difficult to understand why the lower Eastern monkeys should not also have inherited these characters. As this is not the case, there remains no alternative but to assume divergent evolution from an indifferent form. The lower Eastern monkeys are carrying on the evolution in one direction—I might almost say towards a blind alley—while anthropoids and men have struck out a progressive path, at first in common, which explains the many points of resemblance between them, without regarding man as derived directly from the anthropoids. Their many striking points of agreement indicate a common descent, and cannot be explained as phenomena of convergence.

I believe I have shown in the above sketch that a theory which derives man directly from lower forms without regarding apes as transition-types leads ad absurdum. The close structural relationship between man and monkeys can only be understood if both are brought into the same line of evolution. To trace man's line of descent directly back to the old Eocene mammals, alongside of, but with no relation to these very similar forms, is to abandon the method of exact comparison, which, as Darwin rightly recognised, alone justifies us in drawing up genealogical trees on the basis of resemblances and differences. The farther down we go the more does the ground slip from beneath our feet. Even the Lemuridae show very numerous divergent conditions, much more so the Eocene mammals (Creodonta, Condylarthra), the chief resemblance of which to man consists in the possession of pentadactylous hands and feet! Thus the farther course of the line of descent disappears in the darkness of the ancestry of the mammals. With just as much reason we might pass by the Vertebrates altogether, and go back to the lower Invertebrates, but in that case it would be much easier to say that man has arisen independently, and has evolved, without relation to any animals, from the lowest primitive form to his present isolated and dominant position. But this would be to deny all value to classification, which must after all be the ultimate basis of a genealogical tree. We can, as Darwin rightly observed, only infer the line of descent from the degree of resemblance between single forms. If we regard man as directly derived from primitive forms very far back, we have no way of explaining the many points of agreement between him and the monkeys in general, and the anthropoid apes in particular. These must remain an inexplicable marvel.

I have thus, I trust, shown that the first class of special theories of descent, which assumes that man has developed, parallel with the monkeys, but without relation to them, from very low primitive forms cannot be upheld, because it fails to take into account the close structural affinity of man and monkeys. I cannot but regard this hypothesis as lamentably retrograde, for it makes impossible any application of the facts that have been discovered in the course of the anatomical and embryological study of man and monkeys, and indeed prejudges investigations of that class as pointless. The whole method is perverted; an unjustifiable theory of descent is first formulated with the aid of the imagination, and then we are asked to declare that all structural relations between man and monkeys, and between the different groups of the latter, are valueless,—the fact being that they are the only true basis on which a genealogical tree can be constructed.

So much for this most modern method of classification, which has probably found adherents because it would deliver us from the relationship to apes which many people so much dislike. In contrast to it we have the second class of special hypotheses of descent, which keeps strictly to the nearest structural relationship. This is the only basis that justifies the drawing up of a special hypothesis of descent. If this fundamental proposition be recognised, it will be admitted that the doctrine of special descent upheld by Haeckel, and set forth in Darwin's Descent of Man, is still valid to-day. In the genealogical tree, man's place is quite close to the anthropoid apes; these again have as their nearest relatives the lower Old World monkeys, and their progenitors must be sought among the less differentiated Platyrrhine monkeys, whose most important characters have been handed on to the present day New World monkeys. How the different genera are to be arranged within the general scheme indicated depends in the main on the classificatory value attributed to individual characters. This is particularly true in regard to Pithecanthropus, which I consider as the root of a branch which has sprung from the anthropoid ape root and has led up to man; the latter I have designated the family of the Hominidae.

For the rest, there are, as we have said, various possible ways of constructing the narrower genealogy within the limits of this branch including men and apes, and these methods will probably continue to change with the accumulation of new facts. Haeckel himself has modified his genealogical tree of the Primates in certain details since the publication of his Generelle Morphologie in 1866, but its general basis remains the same.[123] All the special genealogical trees drawn up on the lines laid down by Haeckel and Darwin—and that of Dubois may be specially mentioned—are based, in general, on the close relationship of monkeys and men, although they may vary in detail. Various hypotheses have been formulated on these lines, with special reference to the evolution of man. Pithecanthropus is regarded by some authorities as the direct ancestor of man, by others as a side-track failure in the attempt at the evolution of man. The problem of the monophyletic or polyphyletic origin of the human race has also been much discussed. Sergi[124] inclines towards the assumption of a polyphyletic origin of the three main races of man, the African primitive form of which has given rise also to the gorilla and chimpanzee, the Asiatic to the Orang, the Gibbon, and Pithecanthropus. Kollmann regards existing human races as derived from small primitive races (pigmies), and considers that Homo primigenius must have arisen in a secondary and degenerative manner.

But this is not the place, nor have I the space to criticise the various special theories of descent. One, however, must receive particular notice. According to Ameghino, the South American monkeys (Pitheculites) from the oldest Tertiary of the Pampas are the forms from which have arisen the existing American monkeys on the one hand, and on the other, the extinct South American Homunculidae, which are also small forms. From these last, anthropoid apes and man have, he believes, been evolved. Among the progenitors of man, Ameghino reckons the form discovered by him (Tetraprothomo), from which a South American primitive man, Homo pampaeus, might be directly evolved, while on the other hand all the lower Old World monkeys may have arisen from older fossil South American forms (Clenialitidae), the distribution of which may be explained by the bridge formerly existing between South America and Africa, as may be the derivation of all existing human races from Homo pampaeus.[125] The fossil forms discovered by Ameghino deserve the most minute investigation, as does also the fossil man from South America of which Lehmann-Nitsche[126] has made a thorough study.

It is obvious that, notwithstanding the necessity for fitting man's line of descent into the genealogical tree of the Primates, especially the apes, opinions in regard to it differ greatly in detail. This could not be otherwise, since the different Primate forms, especially the fossile forms, are still far from being exhaustively known. But one thing remains certain,—the idea of the close relationship between man and monkeys set forth in Darwin's Descent of Man. Only those who deny the many points of agreement, the sole basis of classification, and thus of a natural genealogical tree, can look upon the position of Darwin and Haeckel as antiquated, or as standing on an insufficient foundation. For such a genealogical tree is nothing more than a summarised representation of what is known in regard to the degree of resemblance between the different forms.

Darwin's work in regard to the descent of man has not been surpassed; the more we immerse ourselves in the study of the structural relationships between apes and man, the more is our path illumined by the clear light radiating from him, and through his calm and deliberate investigation, based on a mass of material in the accumulation of which he has never had an equal. Darwin's fame will be bound up for all time with the unprejudiced investigation of the question of all questions, the descent of the human race.