[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 _h.o.m.o primigenius_. (Schoetensack, _Der Unterkiefer des h.o.m.o 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 Tetraproth.o.m.o argentinus_," etc. _a.n.a.les 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 Muller (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 a.s.sociated 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 c.u.mulative 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 _Naturliche Schopfungsgeschichte_ (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 a.s.sociating 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 ident.i.ty 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 a.s.sailed 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), pa.s.ses 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, Kolliker, 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.

a.s.suredly, 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 surpa.s.ses 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 Nageli nor Weismann, neither De Vries nor Roux, has done this. Nageli, 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 c.u.mulative 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 Nageli, 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 s.p.a.ce) 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 ant.i.theses of philosophy.

This important Mneme-theory of Semon and the luminous physiological experiments and observations a.s.sociated 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 Muller made a closer study of these important phenomena in the instructive instance of the Crustacean larva, as given in his able work _Fur 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 cla.s.ses 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 cla.s.sic work, _The Descent of Man and Selection in Relation to s.e.x_ (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 a.n.a.logous facts--all point in the plainest manner to the conclusion that man is the co-descendant with other mammals of a common progenitor."