Chapter V: Arguments for the Reality of the Processes of Selection (4)

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.

FOOTNOTES:

[Footnote 75: _Life and Letters of Thomas Henry Huxley_, Vol. I. p. 171, London, 1900.]

[Footnote 76: _Ibid._, p. 363.]

[Footnote 77: No italics in original.]

[Footnote 78: _Life and Letters of Charles Darwin_, Vol. I. p. 93.]

[Footnote 79: _Ibid._ Vol. II. p. 263.]

[Footnote 80: _Ibid._ Vol. I. p. 94.]

[Footnote 81: _Life and Letters_, Vol. III. p. 175.]

[Footnote 82: _Ibid._ Vol. II. p. 109.]

[Footnote 83: _Ibid._ Vol. III. p. 112.]

[Footnote 84: _Ibid._ Vol. I. pp. 304-317.]

[Footnote 85: _Life and Letters_, Vol. I. p. 309.]

[Footnote 86: _Loc. cit._ p. 313.]

[Footnote 87: _Ibid._ Vol. II. p. 310.]

[Footnote 88: _Ibid._ Vol. III. p. 236. ["C. Ridley," Mr. Francis Darwin points out to me, should be H. N. Ridley. A.C.S.]]

[Footnote 89: _Descent of Man_ (Popular Edit., 1901), fig. 1, p. 14.]

[Footnote 90: G. Schwalbe, "Das Darwin'sche Spitzohr beim menschlichen Embryo," _Anatom. Anzeiger_, 1889, pp. 176-189, and other papers.]

[Footnote 91: _Descent of Man_, fig. 3, p. 24.]

[Footnote 92: _Descent of man_, p. 6.]

[Footnote 93: _Ibid._ p. 54.]

[Footnote 94: _Descent of Man_, p. 92.]

[Footnote 95: _Ibid._ p. 100.]

[Footnote 96: _Life and letters_, Vol. II. p. 161, June 22, 1859.]

[Footnote 97: _Ibid._ Vol. III. p. 15, March 17, 1863.]

[Footnote 98: _Descent of Man_, p. 132.]

[Footnote 99: _Ibid._ pp. 136, 137.]

[Footnote 100: _Ibid._ p. 143.]

[Footnote 101: _Ibid._ p. 193.]

[Footnote 102: _Descent of Man_, p. 231.]

[Footnote 103: _Descent of Man_, p. 308.]

[Footnote 104: _Life and Letters_, Vol. II. p. 23.]

[Footnote 105: _Loc. cit._ p. 39.]

[Footnote 106: _Loc. cit._ (1856), p. 82.]

[Footnote 107: _Ibid._ Vol. III p. 159.]

[Footnote 108: _Descent of Man_, p. 924.]

[Footnote 109: "Die Hautfarbe des Menschen," _Mitteilungen der Anthropologischen Gessellschaft in Wien_, Vol. XXXIV. pp. 331-352.]

[Footnote 110: _Ibid._ p. 947.]

[Footnote 111: _Descent of Man_, p. 240.]

[Footnote 112: "Das geologische Alter der Pithecanthropus-Schichten bei Trinil, Ost-Java." _Neues Jahrb. f. Mineralogie_. Festband, 1907.]

[Footnote 113: _Descent of Man_, p. 82.]

[Footnote 114: "La race humaine de Neanderthal ou de Canstatt en Belgique." _Arch. de Biologie_, VII. 1887.]

[Footnote 115: Gorjanovic-Kramberger. _Der diluviale Mensch van Krapina in Kroatien_, 1906.]

[Footnote 116: _Studien zur Vorgeschichte des Menschen_, 1906, pp. 154 ff.]

[Footnote 117: "On the cranial and facial characters of the Neandertal Race." _Trans. R. Soc._ London, vol. 199, 1908, p. 281.]

[Footnote 118: Since this essay was written Schoetensack has discovered near Heidelberg and briefly described an exceedingly interesting lower jaw from rocks between the Pliocene and Diluvial beds. This exhibits interesting differences from the forms of lower jaw of _Homo primigenius_. (Schoetensack, _Der Unterkiefer des Homo heidelbergensis_, Leipzig, 1908.) G. S.]

[Footnote 119: _Life and Letters of Thomas Henry Huxley_, Vol. II. p. 394.]

[Footnote 120: _Descent of Man_, p. 229.]

[Footnote 121: _Loc. cit._]

[Footnote 122: Klaatsch in his last publications speaks in the main only of an ancestral form common to men and anthropoid apes.]

[Footnote 123: Haeckels latest genealogical tree is to be found in his most recent work, _Unsere Ahnenreihe_. Jena, 1908.]

[Footnote 124: Sergi, G. _Europa_, 1908.]

[Footnote 125: _See_ Ameghino's latest paper, "_Notas preliminaries sobre el Tetraprothomo argentinus_," etc. _Anales del Museo nacional de Buenos Aires_, XVI. pp. 107-242, 1907.]

[Footnote 126: "Nouvelles recherches sur la formation pampeenne et l'homme fossile de la Republique Argentine." _Rivista del Museo de la Plata_, T. XIV. pp. 193-488.]

V

CHARLES DARWIN AS AN ANTHROPOLOGIST

BY ERNST HAECKEL

_Professor of Zoology in the University of Jena_

The great advance that anthropology has made in the second half of the nineteenth century is due, in the first place, to Darwin's discovery of the origin of man. No other problem in the whole field of research is so momentous as that of "Man's place in nature," which was justly described by Huxley (1863) as the most fundamental of all questions. Yet the scientific solution of this problem was impossible until the theory of descent had been established.

It is now a hundred years since the great French biologist Jean Lamarck published his _Philosophie Zoologique_. By a remarkable coincidence the year in which that work was issued, 1809, was the year of the birth of his most distinguished successor, Charles Darwin. Lamarck had already recognised that the descent of man from a series of other Vertebrates--that is, from a series of Ape-like Primates--was essentially involved in the general theory of transformation which he had erected on a broad inductive basis; and he had sufficient penetration to detect the agencies that had been at work in the evolution of the erect bimanous man from the arboreal and quadrumanous ape. He had, however, few empirical arguments to advance in support of his hypothesis, and it could not be established until the further development of the biological sciences--the founding of comparative embryology by Baer (1828) and of the cell-theory by Schleiden and Schwann (1838), the advance of physiology under Johannes Mueller (1833), and the enormous progress of palaeontology and comparative anatomy between 1820 and 1860--provided this necessary foundation. Darwin was the first to coordinate the ample results of these lines of research. With no less comprehensiveness than discrimination he consolidated them as a basis of a modified theory of descent, and associated with them his own theory of natural selection, which we take to be distinctive of "Darwinism" in the stricter sense. The illuminating truth of these cumulative arguments was so great in every branch of biology that, in spite of the most vehement opposition, the battle was won within a single decade, and Darwin secured the general admiration and recognition that had been denied to his forerunner, Lamarck, up to the hour of his death (1829).

Before, however, we consider the momentous influence that Darwinism has had in anthropology, we shall find it useful to glance at its history in the course of the last half century, and notice the various theories that have contributed to its advance. The first attempt to give extensive expression to the reform of biology by Darwin's work will be found in my _Generelle Morphologie_ (1866)[127] which was followed by a more popular treatment of the subject in my _Natuerliche Schoepfungsgeschichte_ (1868),[128] a compilation from the earlier work. In the first volume of the _Generelle Morphologie_ I endeavoured to show the great importance of evolution in settling the fundamental questions of biological philosophy, especially in regard to comparative anatomy. In the second volume I dealt broadly with the principle of evolution, distinguishing ontogeny and phylogeny as its two coordinate main branches, and associating the two in the Biogenetic Law. The Law may be formulated thus: "Ontogeny (embryology or the development of the individual) is a concise and compressed recapitulation of phylogeny (the palaeontological or genealogical series) conditioned by laws of heredity and adaptation." The "Systematic introduction to general evolution," with which the second volume of the _Generelle Morphologie_ opens, was the first attempt to draw up a natural system of organisms (in harmony with the principles of Lamarck and Darwin) in the form of a hypothetical pedigree, and was provisionally set forth in eight genealogical tables.

In the nineteenth chapter of the _Generelle Morphologie_--a part of which has been republished, without any alteration, after a lapse of forty years--I made a critical study of Lamarck's theory of descent and of Darwin's theory of selection, and endeavoured to bring the complex phenomena of heredity and adaptation under definite laws for the first time. Heredity I divided into conservative and progressive: adaptation into indirect (or potential) and direct (or actual). I then found it possible to give some explanation of the correlation of the two physiological functions in the struggle for life (selection), and to indicate the important laws of divergence (or differentiation) and complexity (or division of labor), which are the direct and inevitable outcome of selection. Finally, I marked off dysteleology as the science of the aimless (vestigial, abortive, atrophied, and useless) organs and parts of the body. In all this I worked from a strictly monistic standpoint, and sought to explain all biological phenomena on the mechanical and naturalistic lines that had long been recognised in the study of inorganic nature. Then (1866), as now, being convinced of the unity of nature, the fundamental identity of the agencies at work in the inorganic and the organic worlds, I discarded vitalism, teleology, and all hypotheses of a mystic character.

It was clear from the first that it was essential, in the monistic conception of evolution, to distinguish between the laws of conservative and progressive heredity. Conservative heredity maintains from generation to generation the enduring characters of the species. Each organism transmits to its descendants a part of the morphological and physiological qualities that it has received from its parents and ancestors. On the other hand, progressive heredity brings new characters to the species--characters that were not found in preceding generations. Each organism may transmit to its offspring a part of the morphological and physiological features that it has itself acquired, by adaptation, in the course of its individual career, through the use or disuse of particular organs, the influence of environment, climate, nutrition, etc. At that time I gave the name of "progressive heredity" to this inheritance of acquired characters, as a short and convenient expression, but have since changed the term to "transformative heredity" (as distinguished from conservative). This term is preferable, as inherited regressive modifications (degeneration, retrograde metamorphosis, etc.) come under the same head.

Transformative heredity--or the transmission of acquired characters--is one of the most important principles in evolutionary science. Unless we admit it most of the facts of comparative anatomy and physiology are inexplicable. That was the conviction of Darwin no less than of Lamarck, of Spencer as well as Virchow, of Huxley as well as Gegenbaur, indeed of the great majority of speculative biologists. This fundamental principle was for the first time called in question and assailed in 1885 by August Weismann of Freiburg, the eminent zoologist to whom the theory of evolution owes a great deal of valuable support, and who has attained distinction by his extension of the theory of selection. In explanation of the phenomena of heredity he introduced a new theory, the "theory of the continuity of the germ-plasm." According to him the living substance in all organisms consists of two quite distinct kinds of plasm, somatic and germinal. The permanent germ-plasm, or the active substance of the two germ-cells (egg-cell and sperm-cell), passes unchanged through a series of generations, and is not affected by environmental influences. The environment modifies only the soma-plasm, the organs and tissues of the body. The modifications that these parts undergo through the influence of the environment or their own activity (use and habit), do not affect the germ-plasm, and cannot therefore be transmitted.

This theory of the continuity of the germ-plasm has been expounded by Weismann during the last twenty-four years in a number of able volumes, and is regarded by many biologists, such as Mr. Francis Galton, Sir E. Ray Lankester, and Professor J. Arthur Thomson (who has recently made a thorough-going defence of it in his important work _Heredity_),[129] as the most striking advance in evolutionary science. On the other hand, the theory has been rejected by Herbert Spencer, Sir W. Turner, Gegenbaur, Koelliker, Hertwig, and many others. For my part I have, with all respect for the distinguished Darwinian, contested the theory from the first, because its whole foundation seems to me erroneous, and its deductions do not seem to be in accord with the main facts of comparative morphology and physiology. Weismann's theory in its entirety is a finely conceived molecular hypothesis, but it is devoid of empirical basis. The notion of the absolute and permanent independence of the germ-plasm, as distinguished from the soma-plasm, is purely speculative; as is also the theory of germinal selection. The determinants, ids, and idants, are purely hypothetical elements. The experiments that have been devised to demonstrate their existence really prove nothing.

It seems to me quite improper to describe this hypothetical structure as "Neodarwinism." Darwin was just as convinced as Lamarck of the transmission of acquired characters and its great importance in the scheme of evolution. I had the good fortune to visit Darwin at Down three times and discuss with him the main principles of his system, and on each occasion we were fully agreed as to the incalculable importance of what I may call transformative inheritance. It is only proper to point out that Weismann's theory of the germ-plasm is in express contradiction to the fundamental principles of Darwin and Lamarck. Nor is it more acceptable in what one may call its "ultradarwinism"--the idea that the theory of selection explains everything in the evolution of the organic world. This belief in the "omnipotence of natural selection" was not shared by Darwin himself. Assuredly, I regard it as of the utmost value, as the process of natural selection through the struggle for life affords an explanation of the mechanical origin of the adapted organisation. It solves the great problem: how could the finely adapted structure of the animal or plant body be formed unless it was built on a preconceived plan? It thus enables us to dispense with the teleology of the metaphysician and the dualist, and to set aside the old mythological and poetic legends of creation. The idea had occurred in vague form to the great Empedocles 2000 years before the time of Darwin, but it was reserved for modern research to give it ample expression. Nevertheless, natural selection does not of itself give the solution of all our evolutionary problems. It has to be taken in conjunction with the transformism of Lamarck, with which it is in complete harmony.

The monumental greatness of Charles Darwin, who surpasses every other student of science in the nineteenth century by the loftiness of his monistic conception of nature and the progressive influence of his ideas, is perhaps best seen in the fact that not one of his many successors has succeeded in modifying his theory of descent in any essential point or in discovering an entirely new standpoint in the interpretation of the organic world. Neither Naegeli nor Weismann, neither De Vries nor Roux, has done this. Naegeli, in his _Mechanisch-Physiologische Theorie der Abstammungslehre_[130] which is to a great extent in agreement with Weismann, constructed a theory of the idioplasm, that represents it (like the germ-plasm) as developing continuously in a definite direction from internal causes. But his internal "principle of progress" is at the bottom just as teleological as the vital force of the Vitalists, and the micella structure of the idioplasm is just as hypothetical as the "dominant" structure of the germ-plasm. In 1889 Moritz Wagner sought to explain the origin of species by migration and isolation, and on that basis constructed a special "migration-theory." This, however, is not out of harmony with the theory of selection. It merely elevates one single factor in the theory to a predominant position. Isolation is only a special case of selection, as I had pointed out in the fifteenth chapter of my _Natural history of creation_. The "mutation-theory" of De Vries,[131] that would explain the origin of species by sudden and saltatory variations rather than by gradual modification, is regarded by many botanists as a great step in advance, but it is generally rejected by zoologists. It affords no explanation of the facts of adaptation, and has no causal value.

Much more important than these theories is that of Wilhelm Roux[132] of "the struggle of parts within the organism, a supplementation of the theory of mechanical adaptation." He explains the functional autoformation of the purposive structure by a combination of Darwin's principle of selection with Lamarck's idea of transformative heredity, and applies the two in conjunction to the facts of histology. He lays stress on the significance of functional adaptation, which I had described in 1866, under the head of cumulative adaptation, as the most important factor in evolution. Pointing out its influence in the cell-life of the tissues, he puts "cellular selection" above "personal selection," and shows how the finest conceivable adaptations in the structure of the tissue may be brought about quite mechanically, without preconceived plan. This "mechanical teleology" is a valuable extension of Darwin's monistic principle of selection to the whole field of cellular physiology and histology, and is wholly destructive of dualistic vitalism.

The most important advance that evolution has made since Darwin and the most valuable amplification of his theory of selection is, in my opinion, the work of Richard Semon: _Die Mneme als erhaltendes Prinzip im Wechsel des organischen Geschehens_.[133] He offers a psychological explanation of the facts of heredity by reducing them to a process of (unconscious) memory. The physiologist Ewald Hering had shown in 1870 that memory must be regarded as a general function of organic matter, and that we are quite unable to explain the chief vital phenomena, especially those of reproduction and inheritance, unless we admit this unconscious memory. In my essay _Die Perigenesis der Plastidule_[134] I elaborated this far-reaching idea, and applied the physical principle of transmitted motion to the plastidules, or active molecules of plasm. I concluded that "heredity is the memory of the plastidules, and variability their power of comprehension." This "provisional attempt to give a mechanical explanation of the elementary processes of evolution" I afterwards extended by showing that sensitiveness is (as Carl Naegeli, Ernst Mach, and Albrecht Rau express it) a general quality of matter. This form of panpsychism finds its simplest expression in the "trinity of substance."

To the two fundamental attributes that Spinoza ascribed to substance--Extension (matter as occupying space) and Cogitation (energy, force)--we now add the third fundamental quality of Psychoma (sensitiveness, soul). I further elaborated this trinitarian conception of substance in the nineteenth chapter of my _Die Lebenswunder_ (1904),[135] and it seems to me well calculated to afford a monistic solution of many of the antitheses of philosophy.

This important Mneme-theory of Semon and the luminous physiological experiments and observations associated with it not only throw considerable light on transformative inheritance, but provide a sound physiological foundation for the biogenetic law. I had endeavoured to show in 1874, in the first chapter of my _Anthropogenie_,[136] that this fundamental law of organic evolution holds good generally, and that there is everywhere a direct causal connection between ontogeny and phylogeny. "Phylogenesis is the mechanical cause of ontogenesis;" in other words, "The evolution of the stem or race is--in accordance with the laws of heredity and adaptation--the real cause of all the changes that appear, in a condensed form, in the development of the individual organism from the ovum, in either the embryo or the larva."

It is now fifty years since Charles Darwin pointed out, in the thirteenth chapter of his epoch-making _Origin of Species_, the fundamental importance of embryology in connection with his theory of descent:

"The leading facts in embryology, which are second to none in importance, are explained on the principle of variations in the many descendants from some one ancient progenitor, having appeared at a not very early period of life, and having been inherited at a corresponding period."[137]

He then shows that the striking resemblance of the embryos and larvae of closely related animals, which in the mature stage belong to widely different species and genera, can only be explained by their descent from a common progenitor. Fritz Mueller made a closer study of these important phenomena in the instructive instance of the Crustacean larva, as given in his able work _Fuer Darwin_[138] (1864). I then, in 1872, extended the range so as to include all animals (with the exception of the unicellular Protozoa) and showed, by means of the theory of the Gastraea, that all multicellular, tissue-forming animals--all the Metazoa--develop in essentially the same way from the primary germ-layers.

I conceived the embryonic form, in which the whole structure consists of only two layers of cells, and is known as the gastrula, to be the ontogenetic recapitulation, maintained by tenacious heredity, of a primitive common progenitor of all the Metazoa, the Gastraea. At a later date (1895) Monticelli discovered that this conjectural ancestral form is still preserved in certain primitive Coelenterata--Pemmatodiscus, Kunstleria, and the nearly-related Orthonectida.

The general application of the biogenetic law to all classes of animals and plants has been proved in my _Systematische Phylogenie_.[139] It has, however, been frequently challenged, both by botanists and zoologists, chiefly owing to the fact that many have failed to distinguish its two essential elements, palingenesis and cenogenesis. As early as 1874 I had emphasised, in the first chapter of my _Evolution of Man_, the importance of discriminating carefully between these two sets of phenomena:

"In the evolutionary appreciation of the facts of embryology we must take particular care to distinguish sharply and clearly between the primary, palingenetic evolutionary processes and the secondary, cenogenetic processes. The palingenetic phenomena, or embryonic _recapitulations_, are due to heredity, to the transmission of characters from one generation to another. They enable us to draw direct inferences in regard to corresponding structures in the development of the species (e.g. the chorda or the branchial arches in all vertebrate embryos). The cenogenetic phenomena, on the other hand, or the embryonic _variations_, cannot be traced to inheritance from a mature ancestor, but are due to the adaption of the embryo or the larva to certain conditions of its individual development (e.g. the amnion, the allantois, and the vitelline arteries in the embryos of the higher vertebrates). These cenogenetic phenomena are later additions; we must not infer from them that there were corresponding processes in the ancestral history, and hence they are apt to mislead."

The fundamental importance of these facts of comparative anatomy, atavism, and the rudimentary organs, was pointed out by Darwin in the first part of his classic work, _The Descent of Man and Selection in Relation to Sex_ (1871).[140] In the "General summary and conclusion" (chap. xxi.) he was able to say, with perfect justice: "He who is not content to look, like a savage, at the phenomena of nature as disconnected, cannot any longer believe that man is the work of a separate act of creation. He will be forced to admit that the close resemblance of the embryo of man to that, for instance, of a dog--the construction of his skull, limbs, and whole frame on the same plan with that of other mammals, independently of the uses to which the parts may be put--the occasional reappearance of various structures, for instance of several muscles, which man does not normally possess, but which are common to the Quadrumana--and a crowd of analogous facts--all point in the plainest manner to the conclusion that man is the co-descendant with other mammals of a common progenitor."

These few lines of Darwin's have a greater scientific value than hundreds of those so-called "anthropological treatises," which give detailed descriptions of single organs, or mathematical tables with series of numbers and what are claimed to be "exact analyses," but are devoid of synoptic conclusions and a philosophical spirit.

Charles Darwin is not generally recognised as a great anthropologist, nor does the school of modern anthropologists regard him as a leading authority. In Germany, especially, the great majority of the members of the anthropological societies took up an attitude of hostility to him from the very beginning of the controversy in 1860. _The Descent of Man_ was not merely rejected, but even the discussion of it was forbidden on the ground that it was "unscientific."

The centre of this inveterate hostility for thirty years--especially after 1877--was Rudolph Virchow of Berlin, the leading investigator in pathological anatomy, who did so much for the reform of medicine by his establishment of cellular pathology in 1858. As a prominent representative of "exact" or "descriptive" anthropology, and lacking a broad equipment in comparative anatomy and ontogeny, he was unable to accept the theory of descent. In earlier years, and especially during his splendid period of activity at Wuerzburg (1848-1856), he had been a consistent free-thinker, and had in a number of able articles (collected in his _Gesammelte Abhandlungen_)[141] upheld the unity of human nature, the inseparability of body and spirit. In later years at Berlin, where he was more occupied with political work and sociology (especially after 1866), he abandoned the positive monistic position for one of agnosticism and scepticism, and made concessions to the dualistic dogma of a spiritual world apart from the material frame.

In the course of a Scientific Congress at Munich in 1877 the conflict of these antithetic views of nature came into sharp relief. At this memorable Congress I had undertaken to deliver the first address (September 18th) on the subject of "Modern evolution in relation to the whole of science." I maintained that Darwin's theory not only solved the great problem of the origin of species, but that its implications, especially in regard to the nature of man, threw considerable light on the whole of science, and on anthropology in particular. The discovery of the real origin of man by evolution from a long series of mammal ancestors threw light on his place in nature in every respect, as Huxley had already shown in his excellent lectures of 1863. Just as all the organs and tissues of the human body had originated from those of the nearest related mammals, certain ape-like forms, so we were bound to conclude that his mental qualities also had been derived from those of his extinct primate ancestor.

This monistic view of the origin and nature of man, which is now admitted by nearly all who have the requisite acquaintance with biology, and approach the subject without prejudice, encountered a sharp opposition at that time. The opposition found its strongest expression in an address that Virchow delivered at Munich four days afterwards (September 22nd), on "The freedom of science in the modern State." He spoke of the theory of evolution as an unproved hypothesis, and declared that it ought not to be taught in the schools, because it was dangerous to the State. "We must not," he said, "teach that man has descended from the ape or any other animal." When Darwin, usually so lenient in his judgment, read the English translation of Virchow's speech, he expressed his disapproval in strong terms. But the great authority that Virchow had--an authority well founded in pathology and sociology--and his prestige as president of the German Anthropological Society, had the effect of preventing any member of the Society from raising serious opposition to him for thirty years. Numbers of journals and treatises repeated his dogmatic statement: "It is quite certain that man has descended neither from the ape nor from any other animal." In this he persisted till his death in 1902. Since that time the whole position of German anthropology has changed. The question is no longer whether man was created by a distinct supernatural act or evolved from other mammals, but to which line of the animal hierarchy we must look for the actual series of ancestors. The interested reader will find an account of this "battle of Munich" (1877) in my three Berlin lectures (April, 1905), _Der Kampf um die Entwickelungs-Gedanken_.[142]

The main points in our genealogical tree were clearly recognised by Darwin in the sixth chapter of the _Descent of Man_. Lowly organised fishes, like the lancelot (Amphioxus), are descended from lower invertebrates resembling the larvae of an existing Tunicate (Appendicularia). From these primitive fishes were evolved higher fishes of the ganoid type and others of the type of Lepidosiren (Dipneusta). It is a very small step from these to the Amphibia:

"In the class of animals the steps are not difficult to conceive which led from the ancient Monotremata to the ancient Marsupials; and from these to the early progenitors of the placental mammals. We may thus ascend to the Lemuridae; and the interval is not very wide from these to the Simiadae. The Simiadae then branched off into two great stems, the New World and Old World monkeys; and from the latter, at a remote period, Man, the wonder and glory of the Universe, proceeded."[143]

In these few lines Darwin clearly indicated the way in which we were to conceive our ancestral series within the vertebrates. It is fully confirmed by all the arguments of comparative anatomy and embryology, of palaeontology and physiology; and all the research of the subsequent forty years have gone to establish it. The deep interest in geology which Darwin maintained throughout his life and his complete knowledge of palaeontology enabled him to grasp the fundamental importance of the palaeontological record more clearly than anthropologists and zoologists usually do.

There has been much debate in subsequent decades whether Darwin himself maintained that man was descended from the ape, and many writers have sought to deny it. But the lines I have quoted _verbatim_ from the conclusion of the sixth chapter of the _Descent of Man_ (1871) leave no doubt that he was as firmly convinced of it as was his great precursor Jean Lamarck in 1809. Moreover, Darwin adds, with particular explicitness, in the "general summary and conclusion" (chap. xxi.) of that standard work:[144]

"By considering the embryological structure of man--the homologies which he presents with the lower animals,--the rudiments which he retains,--and the reversions to which he is liable, we can partly recall in imagination the former condition of our early progenitors; and can approximately place them in their proper place in the zoological series. We thus learn that man is descended from a hairy, tailed quadruped, probably arboreal in its habits, and an inhabitant of the Old World. This creature, if its whole structure had been examined by a naturalist, would have been classed amongst the Quadrumana, as surely as the still more ancient progenitor of the Old and New World monkeys."

These clear and definite lines leave no doubt that Darwin--so critical and cautious in regard to important conclusions--was quite as firmly convinced of the descent of man from the apes (the Catarrhinae, in particular) as Lamarck was in 1809 and Huxley in 1863.

It is to be noted particularly that, in these and other observations on the subject, Darwin decidedly assumes the monophyletic origin of the mammals, including man. It is my own conviction that this is of the greatest importance. A number of difficult questions in regard to the development of man, in respect of anatomy, physiology, psychology, and embryology, are easily settled if we do not merely extend our _progonotaxis_ to our nearest relatives, the anthropoid apes and the tailed monkeys from which these have descended, but go further back and find an ancestor in the group of the Lemuridae, and still further back to the Marsupials and Monotremata. The essential identity of all the Mammals in point of anatomical structure and embryonic development--in spite of their astonishing differences in external appearance and habits of life--is so palpably significant that modern zoologists are agreed in the hypothesis that they have all sprung from a common root, and that this root may be sought in the earlier Palaeozoic Amphibia.