CORNELL UNIVERSITY LIBRARY FROM H.F.Roberts THE DOCTRINE OF DESCENT AND DARWINISM BY OSCAR SCHMIDT PROFESSOR IN THE UNIVERSITY OF STRASBURG WITH TWENT\-SIX WOODCUTS NEW YORK D. APPLETON AND COMPANY 1900 Aegon] Authorized Edition. PREFACE. THE important chapter which closes this work was included in a public lecture which I delivered at the meeting of Naturalists and Physicians held this year at Wiesbaden; my purpose being, as I am willing to con- fess, to ascertain, by experience, whether on this signifi- cant subject I had struck the right note to suit a circle of hearers and readers not hampered by prejudice. After the reception given to this fragment, which I also issued in a separate pamphlet, under the title of “The Doctrine of Descent, in its application to Man” (Die Anwendung der Descendenzlehre auf den Mens- chen), I venture to hope that the whole may find a welcome. With the exception of the Ecclesiastico-political ques- tion, no sphere of thought agitates the educated classes of our day so profoundly as the doctrine of descent. On both subjects the cry is, “ Avow your colours!” We have, therefore, endeavoured to define our standpoint vv vi PREFACE. sharply in the introduction, and to preserve it rigidly throughout the work. This is, indeed, a case in which, as Theodor Fechner has recently said, a definite deci- sion has to be made between two fundamental alter- natives. May our exposition afford a lucid testimony to this dictum of one of the patriarchs of the philo- sophical view of nature. Oscar SCHMIDT. StrassBurG, October 18th, 1873. CONTENTS. I. INTRODUCTION—-SUMMARY OF THE RESULTS OF LINGUISTIC IN- QUIRY—POSITIVE KNOWLEDGE PRELIMINARY TO THE Doc- TRINE OF DESCENT—BELIEF IN MIRACLE—THE LIMITS OF THE INVESTIGATION OF NATURE . ‘ ‘ 3 - - II. THE ANIMAL WORLD IN ITS PRESENT STATE . 4 ‘ III. THE PHENOMENA OF REPRODUCTION IN THE ANIMAL WORLD . Iv. Tue ANIMAL WORLD IN ITS HISTORICAL AND PALAONTOLOG- ICAL DEVELOPMENT. < i ‘ * 4 Vv. THE STANDPOINT OF THE MIRACULOUS, AND THE INVESTIGATION oF NATURE—CREATION OR NATURAL DEVELOPMENT—LIN- N&US—CUVIER—AGASSIZ—EXAMINATION OF THE IDEA OF SPECIES . . ‘ 7 r ” “ ‘ . a ‘ VI. NATURAL PHILOSOPHY—GOETHE—PREDESTINED TRANSFORMA~ TION ACCORDING TO RICHARD OWEN—LAMARCK A ‘ vu PAGE 24 39 60 82 104 viii CONTENTS. VIL LYELL AND MoDERN GEOLOGY—DARWIN’s THEORY OF SELEC- TION—BEGINNING OF LIFE . 5 i % , ‘ VIII. HEREDITY— REVERSION—VARIABILITY —~ADAPTATION — RESULTS oF UsE AND DIsusE OF ORGANS—DIFFERENTIATION LEAD- ING TO PERFECTION 5 “ ‘ 7 g : ‘ é IX. THE DEVELOPMENT OF THE INDIVIDUAL (ONTOGENESIS) IS A REPETITION OF THE HISTORICAL DEVELOPMENT OF THE FAMILY (PHYLOGENESIS) . : “og 7 P 5 7 X. THE GEOGRAPHICAL DISTRIBUTION OF ANIMALS IN THE LIGHT OF THE DOCTRINE OF DERIVATION : ‘ 7 : zi XI. THE PEDIGREE OF VERTEBRATE ANIMALS . ‘. 2 s 5 XII. MAN ei ‘ 7 ‘ i P : ‘ i REFERENCES AND QUOTATIONS . : 2 7 : : i PAGE 127 165 195 222 248 252 311 LIST OF ILLUSTRATIONS. FIG, PAGE 1. Legs of Bird and Chick . . 5 : A 7 : : 9 2. Medusa, Tiaropsis Diadema . 5 i . "i - 31 3. Stauridium. Medusa, Cladonema Radiata ‘ ‘ , 43 4. Spermatozoa 3 r i ‘ * 45 5. Section of Larva of Ciieaeaes Seatige ‘ é , 7 » 51 6. Embryo of Hydrophilus Piceus 5 - . é . a 1153 7. Sessile Stage of Crinoid . ‘ ‘ P ‘ ; 7 » 56 8. Larva of Crayfish . é : : A ‘ ‘ - 56 g. Graptolites. ‘i z ‘i : s 3 5 ‘ 69 10. Trilobites remipes . ; 3 5 r i ‘ : + 70 11. Palzoniscus ‘ é ‘ * . - ‘5 , o 72 12. Larva of Bewinodern : : . ‘ 7 é : - 197 13. Larva of Sea-Snail . ji 3 5 é 5 ; ‘ . 2C0 14. Stauridium. Cladonema . : - 5 i ‘ 5 - 203 15. Hydractinea carnea . e : : ‘ F 5 5 204 16. Larva of Parasitic Crustacea . 3 5 3 ; 7 . 207 17. Axolotl . ‘ : . . 5 : - : . 208 18. Amblystoma : : ‘ - - e » 209 1g. Ammonites Humphresianus . 7 Ina paper, published in 1855, he demonstrated the dependence of the flora and fauna on the geographical position and geological nature of the district of propagation, and the close connection of the species, according to time and habitat, with kindred species previously existing; and in a second work, in the year 1858, on the inclination of varieties to deviate without limit from the original type, we find a disquisi- tion on the importance of the struggle for existence, the consequences of adaptation, the selection of the most useful, and the replacement of the earlier species by the establishment of the more valuable varieties. We shall repeatedly have occasion to draw upon the rich sup- plies of his researches. VII. Heredity—Reversion—Variability—Adaptation—Results of Use and Disuse of Organs—Differentiation leading to Perfection. THE two properties of organic being which determine and regulate the relation of the offspring to the pro- genitors, and which not only assign to individuals their position in the surrounding world, but also help them to attain it, are transmission or heredity, and adaptation. Heredity is the conservative, adaptation, the pro- gressive principle. Yet all heredity is not directed to immutability, and many cases of adaptation involve morphological and physiological retrogression. For the elucidation of the inherited peculiarities of organisms, we reconstruct their pedigree; by the characters acquired by adaptation, we test the pliability of organisms in the lapse of time, and trace the ramifications of the pedigree. Groups of organisms, in which the conservative principle predominates, certainly evince their powers of endurance in the struggle for existence, but they make no advance in physiological value, and are outstripped by the more progressive groups which yield to obstacles and profit by them, a course of which human life also affords so many examples. As the phenomena of heredity are usually more obvi- ous than the results of adaptation, the latter was almost 165 166 THE DOCTRINE OF DESCENT. entirely neglected by naturalists in former days. And indeed what comparison in organic nature can be made so frequently and universally as the resemblance of the offspring to the parent? An anatomist, it is true, quaintly attempted to work out the proposition that the resem- blance in the children is not dependent on heredity, but is the result of identical and similar influences, customs, and habits, prevalent in families. But this paradoxical theory requires no special refutation. It is quite true that similar habits and similar external impulses elicit a certain similarity of demeanour and appearance; but if the little son of the pompous millionaire apes his father, it cannot be said that he has likewise mimicked his large or small nose, &c., or has acquired it by a similar call for adaptation. We have only cursorily alluded to this quibble, in flagrant contradiction as it is with every ex- perience; and, in conformity with general opinion, we corroborate the transmission of the parental characteris- tics to the offspring. The breeders of animals in particu- lar has occasion to observe these transmissions spe- cially, and to evolve their astounding progress from the combination and reciprocal influence of the various forms and degrees of heredity. It is well known that not only are normal conditions transmitted, but monstrosities are also reproduced through several generations, and, as we have seen in the instance of the crooked-legged sheep of Massachusetts, may even be established as the characters of a race. A mere reference to the inheritability of morbid tendencies, bodily and mental, will enable us to realize this intrinsic connection of the offspring to the ancestors. Only since the theory of selection has rendered the modalities of the HEREDITY. 167 transmission of bodily characters a subject of more pro- found study, have general and national psychology been impelled to estimate the influence of heredity in the province of the mind, and demonstrate how, in the vari- ous races and families of nations, the molecular peculiari- ties of the brain, the tendency of character and intelligence of the individuals, and whole series of ideas, conform both in vigour and purport to the laws of heredity. It is manifest that the key to the phenomena of hered- ity must be looked for in the. process of reproduction. The molecular motions and disturbances, the inconceiv- ably minute mechanical transfers which take place, do not, indeed, admit of observation. They are, however, no more “obscure” and “ enigmatical,” as they are so readily termed, than the invisible, but not supernatural motions, on the control and calculation of which the stately edifice of theoretic Chemistry and Physics se- curely rests. With the advance from asexual to sexual reproduction, and from the simple to the more perfect organisms, the difficulty of representation increases, but not that of abstract comprehension. If a low organism, a monad, divides itself, the divided individuals differ from the parent individual only in their inferior bulk, and the difference of their functions is, as to quality, nil. So, too, where gemmules and germs separate from a parent organism, the dower of the offspring is so large that identity in form and function of progenitor and progeny appears self-evident and natural. But the sexual reproduction of composite organisms is, as we have known since the old doctrine of the aura seminalis was refuted, also a separation of material portions of the pa- rental organisms. It is still a mechanical process which I2 168 THE DOCTRINE OF DESCENT. is not incomprehensible, and seems inexplicable only if we make the naturally futile attempt to bring sensibly before us the infinitely minute agencies which operate both mechanically and chemically. In the ‘‘ Variation of Plants and Animals,” Darwin has set up a provisional hypothesis of Pangenesis. He says that all phenomena of heredity and reversion would thereby be rendered pos- sible, that in every elementary or cellular portion of the ‘ organism innumerable gemmules are produced, which are hoarded up in the reproductive elements, in every ovum, in every sperm corpuscule, and might remain latent during hundreds of generations, and only then exhibit their powers in reversion.*® This hypothesis, it appears, has met with no ready approbation, probably, as it seems to us, because, in the attempt to meditate upon it, the sensible representation forces itself forward only to prove inadequate. But if it be steadfastly borne in mind that in Protoplasm, as Rollet *7 appropriately terms it, the most complex phenomenal forms of life pos- sess a most persistent witness of their connection with the simplest, it follows that the general laws shown to be true or probable with reference to the simplest or- ganisms, must be applicable to the most perfect also. This holds good also in reproduction, which, in its funda- mental phenomena, offers nothing that cannot be based upon molecular physics applied to colloidal living sub- stance capable of imbibition, and thus divested of vitalistic dualism. The more highly complex is an organism, that is, the greater the differentiation in the development from the protoplasm of the germ-cell to maturity, the more heterogeneously does heredity display itself. These REVERSION. 169 modes of heredity have been defined by Darwin, and yet more systematically by Haeckel, as “laws of in- heritance,” and corroborated by them by a profusion of examples. If heredity may be termed the conserva- tive element in the life of species, we may also speak in particular of a conservative heredity, by which the old, long-established characteristics and peculiarities are transferred. The more stubbornly a character is trans- mitted, or, what amounts to the same, the greater the number of families, gcnera, and species, over which a character is extended, the more ancient must it be con- sidered, the earlier did it appear in the ancestral stock. In most cases, this conservative heredity occurs in an unbroken succession of generations, an observation on which it is needless to enlarge, as it may be daily made by every one. But conservative heredity may likewise dis- play itself intermittently, either when merely individual characters of the ancestors reappear after lying dormant for one, several, or many generations,—a phenomenon designated as Atavism, or reversion,—or when the species is composed of a regular alternation of variously con- stituted generations and individuals. ‘This particular sort of reversion is termed Alternate Generation, or Heterogenesis. No one is surprised if children exhibit the bodily or mental features of their grand-parents which were sus- pended in the parents. But most frequent and striking is the atavism of domestic animals and cultivated plants, a stubborn antagonist to breeders. Of no domestic ani- mal is the aboriginal stock known with such approximate certainty as that of the pigeon. Now there are races of pigeons purely bred for several centuries, and in 170 THE DOCTRINE OF DESCENT. colour and shape transformed into new creatures, which yet from time to time spontaneously, or by crossing with other conspicuous races, produce birds which, in colouring and characteristic pencilings of black bars on wings and tail, resemble the wild rock- pigeon. “T paired,” says Darwin,®* “a mongrel female barb- fantail with a mongrel male barb-spot, neither of which mongrels had the least blue about them. Let it be remembered that blue barbs are excessively rare; that spots, as has been already stated, were perfectly char- acterized in the year 1676, and breed perfectly true; this likewise is the case with white fantails, so much so that I have never heard of white fantails showing any other colour. Nevertheless, the offspring from the above two mongrels was of exactly the same blue tint over the whole back and wings as that of the wild rock- pigeon of the Shetland Islands; the double black wing-bars were equally. conspicuous; the tail was exactly alike in all its characters, and the croup was pure white.” Another reversion frequently to be observed is the striping of the feral domestic cat of Europe, in which it resembles the wild-cat so closely as to be scarcely distingdishable. Darwin has collected evidence from which we may infer that the wild ancestral stock of the horse was striped, and this evidence includes the appearance of striped individuals. But yet another strange phenomenon in horses may be interpreted by atavism. Foals are occasionally born with supernumerary toes. This “monstrosity” can be explained only by reversion to the three-toed historical ancestors of the present genus. These vouchers are sufficient. PROGRESSIVE HEREDITY. 171 All the phenomena of artificial breeding, as well as natural selection, serve to show that not only the char- acters descended from past ages, but also those subse- quently and most recently acquired, may be transmitted to posterity. This is progressive heredity. Without it, improvement and progress would be impossible; and its own possibility is the direct result of the nature of re- production. The newer a useful modification, the less has it hitherto been able to place itself in correlation with the entire organism, the less is the reproductive sys- tem as yet affected by it; the more uncertain and fluc- tuating, therefore, is the transfer by propagation; breed- ing, or natural selection, is requisite to convert the po- tentiality of progress into a fact, and gradually to enrol this fact among the conservative inheritances. Progressive heredity is naturally more complex where the sexes are separate, where sexual selection asserts its rights, and the advantages of one sex are fostered by the taste of the other, and are then either trans- ferred exclusively to the sex benefited by its secondary characters, or turned to the profit of the whole species. As a rule, the males are endowed with these advantages, and have transmitted them incompletely to the females. We will explain ourselves by a single example. In the order of insects termed Orthoptera (or straight-winged), the males, by rubbing their wing-covers together, or by stroking them with the lower portion of their hind legs, are able to make a music attractive to the females. Von Graber, a distinguished modern entomologist, has shown °° that the teeth of the stridulating instruments of these animals are merely modified hairs; that their con- struction may be explained by their use; and that in all 172 THE DOCTRINE OF DESCENT. probability they have been perfected by sexual selec- tion, the best and loudest musicians being the most favoured wooers. With one single exception, the fe- males of the Orthoptera are dumb, but many possess traces of the stridulating apparatus peculiar to the males. Contrary to the older opinion, that it was merely a case of transmission emanating from the males, Graber has made it ‘‘ more than probable that the resonant nervures of the females—of the stridulating Ephippigera vitium— have been gradually developed independently of the males, but in the same manner.” In other cases, on the contrary, the feebly developed nervures of the females, unfit to produce audible stridulations, seem to be an in- heritance from the males. Heredity at corresponding periods of life is a well- known phenomenon. The tendency to disease is trans- mitted from the father or the mother to the child to break out at the age at which they suffered. Generation after generation, the milk teeth make room for the per- manent teeth at a corresponding time. ‘But all special cases are mere results of the general law of develop- ment, by which in the individual characters appear in the sequence in which they were historically acquired and became susceptible of transmission. Heredity, at a definite age after the period at which we consider actual development to be complete, is after all only a continu- ation of the embryonic development, beginning with fission, germ and ovum, of which the ninth chapter will teach us the signification. In this development of the individual, or ontogenesis, as will be shown below in more detail, processes are frequently abridged or totallv omitted which once, while they were being acquired and MUTABILITY. i 173 after they had been established, occupied a longer period, but in the course of selection either became of less im- portance to the individual, or preserved a physiological value only as phases of transition. The second great class of characters, namely, those which have been newly acquired and depend on adapta- tion, pre-suppose the mutability of the organism. This is a fundamental phenomenon of organic bodies. It is inherent in the minutest morphological constituents, in protoplasm, and in cells, and in the morphological ele- ments evolved from them, the pervading and determining individual life of which results in the collective life of the creature. The organic morphological element is in a state of saturation; it is continually imbibing and emitting, and its stability is therefore constantly dependent on the supply of material for its functions. For nutrition, which generally and wholly determines the external appearance and the nature of the individual, is accomplished by the innumerable cells and their derivatives. Every fluctua- tion of supply to any part of the organism, nay, to a single point of the surface of a microscopic atom, must involve a modification of textural parts, or in the struc- ture of integrated textural groups or organs. Mutability is thus a character resulting from the in- trinsic nature of organism, and dependent on external conditions, which determine quantity and form, as well as the development and transformation of the ele- mentary constituents, or their abortion and _ retro- gression. These effects may be exhibited in a polype- stem, which as a whole represents the individual, in its single polypes, the cells and morphological elements. The single individuals are alike in their constitution, but 174 THE DOCTRINE OF DESCENT. are usually very different in size and development, even in those species in which the differentiation unquestion- ably produced by selection has not led to polymorphism ‘or separation into personal groups performing different functions. The weal or the woe of our polypes is greatly. dependent on the position which they occupy upon the stem; the supply of nutriment primarily furnished to the single individuals is unequally and variably appor- tioned according to currents and tides. Hence on each polype-stem there are regions where the single polypes are especially thriving, others where they are just able to maintain themselves, others where they cannot keep their balance. But as the polype-stem is traversed by a canal system conveying the nutritive fluid and con- necting the several cells, the superfluity of the well- situated cells goes to the benefit of those for whom a worse lot was prepared by their accidental position, and conversely. These relations, which, complex as they seem, are very simple for our comparison, determine the form and appearance of the polype-stem. Among a hundred thousand stems, no two will be found absolutely alike. To return to the mutability of organisms, even if two individuals of the same species are bred under the most similar conditions imaginable, it has never been possible to pronounce them absolutely alike. That mu- tability is slighter in lower than in higher organisms, is a prejudice frequently repeated and fortified by the old dogma of species. The doctrine of descent and se- lection would fare ill if the case were so. But as the shepherd unerringly knows the physiognomy of his sheep where an excursionist from the town sees only ADAPTATION. 175 a general sheep’s face, so to an attentive naturalist, in most of the lower organisms, the specific type re- solves itself into as many varieties as individuals, irre- spectively of the cases in which no specific type can be established. As modification under given conditions, adaptation is thus as little an unknown quantity as heredity, but is merely a function of the mechanical character of muta- bility, or, in the widest sense of the word, of nutrition. Adaptation takes place when the organism or its parts are pliable and plastic to external influences, when they conquer and make use of them. Climate, light, humid- ity, nutriment, are hindrances or advantages that directly or indirectly affect the organism, and are all actively concerned in it. Surrounded by organisms, we see them without exception adapting themselves to circumstances; and if our only object is to be convinced of the formative influence of the mode of life, this is most readily done in the case of domestic animals. In his studies on the pig, H. von Nathusius, perhaps the most scientific, of the celebrated breeders, shows how in the simplest cases, where the looseness of cultivated soil has facilitated the labour of grubbing, the skull of the domestic pig is ar- rested, by the softer structure of the cranium, at the im- mature form of the wild boar, and how those extreme shapes of the head in cultivated breeds, characterized by the bending and shortening of the face, and the impossi- bility of closing the jaw in front, are entirely the result of their altered mode of life. It is known that men, animals, and plants, removed far from their previous abode to a new and strange environment, after a longer or shorter effort of the organism to domesticate itself, 176 THE DOCTRINE OF DESCENT. ‘ either die out, or else accommodate themselves to the new conditions and become acclimatized. Every accli- matization is therefore an adaptation, accompanied by modifications more or less perceptible. Thus, in conse- quence of the varied conditions of life, there is a wide divergence among races of men who, by their kindred language, are of the same origin, not to mention those whose relations linguistic inquiry has not yet decided. How different is the idiosyncrasy of the Englishman from that of the Hindoo! Physically and psychically, they represent two remarkable sub-races of which the pe- culiarities must be ascribed to adaptation,—in the latter, to a climate which requires a vegetable diet, and, eliciting neither bodily nor mental energy, favours a dreamy sensuality; in the former, to a country which is in every particular the opposite of the Indian original home. Similarly, the annual alternation in the vital phenomena of so many organisms, designated as hybernating ani- mals, is a case of adaptation. It is changed the moment the organism is exposed to another climate, or rather acclimatization is essentially the accommodation of the hybernating animals to the new climate. In all these examples we have the results of direct adaptation, in which the power of resistance in the in- dividual comes into play, as does cumulative adaptation in artificial, and the survival of the fittest in natural, selection. In all cases of adaptation, one or several or- gans are primarily concerned, either actively or passively; and only in consequence of the resulting modifications are the other organs drawn into sympathy. This may be termed correlative adaptation. It might be supposed that the most perspicuous examples would be afforded CORRELATIVE ADAPTATION. 177 by parasitic animals, in which, with the alteration of the aliment and of the alimentary apparatus, especially of the manducatory portions, is usually combined a trans- formation and retrogression, often extending to the total extinction of the locomotive organs, and of the entire segmentation of the body. But, although the limits are difficult to define, the cause of these associated modifi- cations in the locomotive and alimentary apparatus con- sists less in their reciprocal sympathetic influence than in their simultaneous disuse. It is, however, by correlative adaptation that, for in- stance, in the short-beaked races of pigeons, the middle toe and astragalus are shortened, and that in the long- beaked races these organs have shared in the elongation. In the case, however, in which short beaks are combined with short feet, a certain share in the shortening of the feet is also owing to disuse; while where the pigeon- fancier took pleasure in the elongation of the beak by cumulative selection, the correlative elongation of the foot took place in spite of disuse. The most important group of correlative modifications or adaptations, always using this word in its widest acceptation, relates to the sphere of the sexes. Direct attacks on the generative organs manifest their effects on all the rest of the or- ganism, as is best shown in animals of both sexes cas- trated for the market or for labour. We have already seen that the degree of perfection attained in the orders of the Articulata, Annulosa, Vertebrata, and partially in the Radiata also, depends on the integration of the originally similar parts lying behind or by the side of one another; hence on the di- vision of labour. This Haeckel has designated divergent 178 THE DOCTRINE OF DESCENT. adaptation. It gives rise to the remarkable poly- morphism, which appears especially in the marvellous forms of the Hydra tuba; and, higher up, in the organization of the communities of the Termites and the Bee, &c. So far as modification coincides with adaptation, the direct adaptations hitherto discussed may be contrasted with a series of indirect adaptations. Among these may be comprised a series of phenomena of which the causes do not fall within the life of the individual, but are to be sought in influences by which the parents were af- fected. It is obvious that we here come into contact with the province of heredity, which is well known to breeders. Thus H. v. Nathusius, in his studies on the formation of the pig’s skull,*t says:—‘‘ From the facts here col- lected, it is plain that the transmission, the transfer of the form of head from the parents to the offspring, does not unconditionally ensue. If the form of skull, which we will briefly term the cultivated form, be the product of nutrition and mode of life, hence of external influ- ence,—if it can be differently formed in the same indi- vidual, and is therefore not constant,—in that case the heredity of this form cannot be spoken of without quali- fication. The form itself will not be transmitted to the offspring, but only the tendency to the form. This may be inferred from the circumstance that from generation to generation, and to a certain degree, the form increases in peculiarity. If we rear a common with a thorough- bred pig, and if we allow exactly the same influences of nutriment and keeping to operate upon both, and in equal measure, we shall not obtain the same form of head. The development of the form of head must there- MIMICRY. 179 fore be aided by a pre-existing tendency, and we must hence regard it as hereditary.” Haeckel likewise propounds a law of individual adaptation, which expresses the fact that, notwithstand- ing the closest kinship, individuals diverge in many ways. The cause of this difference, chiefly conspicuous in the individuals of the same litter or brood, is, so far as it is not due to adaptation, inherent in the germs, and is transferred to them by fluctuations and differentiations in the conditions of nutrition in the parents, mostly be-. yond our ken. Other phenomena of indirect adapta- tion are exhibited in the occurrence of malformations, of which the causes must be looked for only in dis- turbances of nutrition in the parental organisms by which the progenitors themselves were not percepti- bly affected. Here also belong the cases, in which influences which have affected one sex only are mani- fested exclusively in posterity in the same sex. As may be seen, these processes, of which the initia- tion is entirely withdrawn from observation, are closely connected with the most obscure province of heredity. An extremely interesting and important form of adaptation is the so-called mimicry, or protection by means of colouring and form. The first discoveries on this subject were made by Bates, the well-known “Naturalist on the Amazon;” the greater part were subsequently added by Wallace. In South America, the family of butterflies named Heliconida is extraor- dinarily extensive; they are remarkable for their elongated wings, body, and antenne, and for the beauty of their colours. It might be imagined they were ex- 180 THE DOCTRINE OF DESCENT. posed to the persecutions of insectivorous birds and other animals; but this is not the case, for they have a disagreeable smell, which, in all likelihood, renders them obnoxious. Their smell and flavour are thus a protec- tion, as the birds and lizards who have once seized them by mistake are certain, ever after, to leave them unmo- lested. Now, as the insectivora do not test the individual case, but have adopted a general repugnance to the as- pect of the Heliconidz, if other butterflies resembled the -Heliconide without possessing the bad smell, they would participate in the security to life enjoyed by the Heli- conide in proportion as they approach their external appearance. The case has actually occurred, for Bates discovered a number of species of the otherwise very different genus, Leptalis, of which each almost undis- tinguishably resembles one of the Heliconidz both in colour and form. The Leptalide have also adopted the flight of the Heliconide, share their habitats, and, al- though without the offensive smell, fly about with im- punity. This state of things would be impossible if the Leptalidze were not considerably in the minority, so as to be in a measure hidden by the Heliconidz. Wallace has proved that species protected by mimicry of other animals are invariably in the minority, and often very rare in comparison with the species which they imi- tate. Neither the explanation that like conditions of life produced like results, nor the hypothesis that, in some cases at least, the mimicry consists in reversion to a common original species, is in any way satisfactory. Many cases can be interpreted only by natural selec- tion, those, namely, where from the first, before the imi- tation had begun, such a resemblance already existed MIMICRY. 181 between the imitating and imitated forms as to render confusion possible; where, therefore, the resemblance so conducive to the preservation of those in which it was the strongest, needed only to be increased by natural selection. Darwin * is also of opinion “that the process probably has never commenced with forms widely dis- similar in colour.” A peculiar, simpler and long known mimicry, is when animals have accommodated themselves in colour to their habitats in such a manner as not to attract the attention of their enemies, and likewise to deceive their prey. Who, in the days when he chased butterflies, did not learn how difficult it is to recognize certain evening and nocturnal flyers on the bark of trees, as they quietly sit with their dusky brown or gray-striped or speckled wings, outspread in a roof-like shape? The tree locusts and Mantidz can look so deceptively like leaves or twigs, that it is only by the touch that one can be as- sured of their real nature. Wallace relates that one of the Phasmide (Ceroxylus laceratus), which he obtained at Borneo, was so covered with pale olive-green ex- crescences, that it looked like a stick covered with moss. The Dyak who brought him the animal declared that, although alive, it was really overgrown with moss, and the naturalist himself was only convinced of the con- trary by the closest examination. A remarkable example of advantageous colouring, within easy reach of many of our readers, is exhibited in most species of the flat-fish (Pleuronectidz), now so frequently kept in aquaria. Observe the gray or brown- ish speckled creatures, as with a few strokes of their fins they partially cover their upper surface with sand. 182 THE DOCTRINE OF DESCENT. They need not bury themselves entirely, for it is only by close examination that their bare skin can be distin- guished from the sandy bottom, and under this partly artificial, partly natural veil and mask, the animal waits for its prey. In many animals provided with protective colouring the phenomena are more complex, and explanation by natural selection is far more difficult; for they are able voluntarily to adapt their colour to circumstances, or else their colour changes by involuntary reflexes. Verany’s unsurpassable observations on the Cephalopoda have ac- quainted us with the range of colours at the disposal of these Molluscs, and to this may be joined Brehm’s description of the changes of colour in the chameleon. On these highly complex cases some light is thrown by the simpler instances in which the manifestly protective colouring has become fixed in skin and plumage, and the concurrence of other circumstances scarcely admits of any other explanation than selection. On this point, Wallace’s interesting researches on bird’s-nests are especially instructive. The great ma- jority of female birds which sit in open nests possess brown or gray, in short, unobtrusive plumage. No con- tradiction will be offered to the statement that any casual modifications of plumage, which would more readily be- tray the sitting bird to its enemies, would have no pros- pect of becoming constant. The converse follows naturally with regard to colouring which brings the bird into harmony with its environment; and an important guarantee of the correctness of this interpretation of facts is afforded by the other observation, that most female birds with gaily coloured or speckled plumage sit in cov- USE AND DISUSE OF ORGANS. 183 ered and concealed nests. It must be added, that the construction of nests is not determined by the absolute rules of a blind instinct, but is modified by the experience of the animals, an experience of which we are indeed scarcely able to perceive the development, except with the age of the individual, but which, at least in several cases, has been proved to be the progress of the species. Natural selection has an important accessory in the modifications produced by the use or disuse of organs. Compulsion to more diligent use, inducements to dis- use, are involved in the varying conditions of life. In both cases it is therefore a question of adaptation. Look- ing at nature, profound modifications are most readily demonstrated as the consequence of disuse; but artificial selection gives numerous examples of both sorts, espe- cially where disproportionate use of certain organs is combined with simultaneous disuse of others. Such products of selection with disproportionate use are the racer and the dray-horse. The blindness of cave animals admits of no explana- tion, but that, with the increasing uselessness of the eyes during accommodation to cave life, the exchange of ma- terial in the less active organs gradually diminished, and atrophy was initiated. The accuracy of these theoretical observations is enforced by the observation that the near- est kin of many blind cave animals, especially of insects and spiders, reside in the vicinity of the cave, and that those cave animals which inhabit passages only partially obscure, possess less atrophied optic apparatus. A singular gradation occurs among the burrowing mam- mals, and Darwin ® cites an example admirably illus- tating the loss of sight in consequence of the mode of 13, 184 THE DOCTRINE OF DESCENT. life. “In South America a burrowing rodent, the Tuco- tuco, or Ctenomys, ‘is even more subterranean in its habits than the mole; and I was assured by a Spaniard, who had often caught them, that they were frequently blind; one which I kept alive was certainly in this con- dition, the cause, as appeared on dissection, having been the inflammation of the nictilating membrane. As fre- quent inflammation of the eyes must be injurious to any animal, and as eyes are certainly not necessary to ani- mals having subterranean habits, a reduction in their size, with the adhesion of the eyelids and growth of fur over them, might in such case be an advantage; and, if so, natural selection would constantly aid the effects of disuse.” In the classes of flying animals, a large number have left off flying; and we find their flying apparatus in an aborted or incomplete condition, which perverse judg- ment and reasoning alone can regard as a state of pro- gressive development from yet simpler rudiments. If throughout the great family of the Coleoptera, genera and species are to be found with imperfect flying ap- paratus, consolidated wing covers, &c., if the whole fam- ily of Staphylinze does not possess the power of flight, no one dreams of considering them as arrested forms; but it is conceivable that the mode of life in which they differ from the other members of their order and class, gradually superinduced in their flying ancestry the habit of not flying, and at the same time the atrophy of the organs of flight. With this was combined, as these beetles show, no degradation of organization, but, on the contrary, a higher and extremely advantageous de- velopment of other organs, the manducatory and loco- USE AND DISUSE OF ORGANS. 185 motive apparatus. A general reduction of the power of flight has been shown in the beetle fauna of many islands. Thus in Madeira, of 550 species, over 200 fly imperfectly or not at all, and for this there is no ex- planation but natural selection. Here the less good and enterprising flyers had the advantage, while the others were blown into the sea and eliminated. The non-application of a previously attained special per- fection is advantageous in the “struggle for exist- ence.” In several families of lizards, some genera are ser- pentine, as they are termed, which, with elongated bodies, possess either fore-legs only (Chirotes), or merely rudi- mentary hind-legs (Pseudopus), or no vestiges of legs (Anguis). They bear the same relation to the great class of normally four-legged lizards as the non-flying insects to their own class. They have not been arrested in their development, nor are they animals in process of evolv- ing four legs; but, as Fiirbringer has demonstrated from the history of development and comparative anatomy, their limbs, and—if these are entirely absent—the remains of the pectoral and pelvic arches and the sternum bear indubitable marks of the abortion of a once complete apparatus. Further comparison shows that this atrophy reaches its climax in the snakes, but that it is compen- sated for by the ribs and intercostal muscles having un- dertaken the work of the limbs. Here, again, disuse and adaptation coincide as well as differentiation. In the class of birds is repeated the spectacle we have just witnessed in beetles and reptiles. In some few families and smaller groups, individual species are de- prived of the power of flight, and one whole large sys- 186 THE DOCTRINE OF DESCENT. tematic group is characterized by the incapacity of fly- ing. In our opinion, there was a direct connection be- tween the inducements to disuse and its consequences in the case of the dodo, which, with its few con- geners, so promptly fell a sacrifice to its helplessness on the discovery of the lonely islands which they had prob- ably inhabited for thousands of years without disturb- ance. In no other way has the northern penguin (Alca impennis) at some time obtained the curtailment of its wings; and the scanty but wide-spread remains of the order of flightless birds indicate a period at which, in a more peaceful environment, their far more numerous -wingless ancestry made less use of their pinions, and natural selection endowed them with greater strength and nimbleness of leg. The effects of disuse of the or- gans of locomotion are likewise directly exhibited by ’ artificial selection. : Use and disuse, combined with selection, elucidate the separation of the sexes, and the existence, otherwise totally incomprehensible, of rudimentary sexual organs. In the Vertebrata especially, each sex possesses such dis- tinct traces of the reproductive apparatus characteristic of the other, that even antiquity assumed hermaphrodit- ism as a natural primeval condition of mankind. The technical proofs of the homologies concerning these partly manifest, partly internal and hidden relations, are given in the manuals of comparative anatomy. We shall merely indicate the manner in which the theory of selec- tion is here borne out. It is self-evident that in her- maphrodite animals, fluctuations in the sexual sphere must take place, in which one half or the other will predominate. Should these fluctuations be sufficiently PARASITES. 187 strong for natural selection to take possession of them, the productive power of the less active portion will gradu- ally decrease, and finally, with the extinction of the physiological character and the function, nothing will be transmitted but the morphological remains, as a mock- ery to the theory of special design or teleology. Here and there only occurs a reversion more or less striking, connected, however, almost exclusively with the adjunc- tive organs, and the secondary sexual characters, by which we mean, not those acquired by either sex, but originally common to both. The tenacity with which these rudiments of sexual organs are inherited is very remarkable. In the class of mammals actual hermaphro- ditism is unheard of, although through the whole period of their development they drag along with them these residues, borne by their unknown ancestry no one can say how long. Unless we suppose that parasitic animals were created simultaneously with their hosts from the dust of the earth,—man and his tapeworm, and other disagreeable guests,—and thus put an end to the discussion, this en- tire province has to be explained by descent, with the special co-operation of disuse. The proposition to be demonstrated in the next chapter, that the evolution- ary history of the individual represents the history of the species, will show the influence of the disuse of particular organs on the configuration of the various parasites. The parasitic Crustacea are perhaps the most instructive, as they present the most complete systematic series, exhibiting the gradual atrophy of the organs which accompanies the ever-increasing connection of the para- site with his host. In several orders of intestinal worms, 188 THE DOCTRINE OF DESCENT. the alimentary canal has become entirely unnecessary; but they exhibit neither intermediate forms nor phases of development. It is different, however, with the para- sitic Crustacean, for here the young, locomotive, and well-integrated being as its protype in definitive ge- neric forms permanently locomotive, from which, after adhesion, it deteriorates into a mere motionless sac. All these animals, including the intestinal worms, have ac- quired their position and status (and this is the true significance of parasitic life) by the apparent degrada- tion of their organization. They are, almost without exception, distinguished by their reproductive power; and on this, owing to the easy supply of nutriment, without any exertion of the other parts of the or- ganic system, the whole bodily activity could be concen- trated. We have hitherto demonstrated that organisms are urged to continual differentiation by the unremitting struggle for existence. For the cultivation of morpho- logical species, natural selection, moreover, seizes on the modifications arising from the mere variability of the organism, and implying no physiological advance. But sooner or later these are also inevitably drawn into the vortex of competition. After what has been already said, this fact is so self-evident as to need no further proof. Even did we not see the infinite variety of or- ganisms, a divergence into novelty must needs be in- ferred on @ priori grounds, from the existence of the simple and uniform, and the necessity of adaptation to altered external conditions. But with development in various directions, under the guidance of natural selec- tion, progress is necessarily combined. It is one THEOLOGICAL. INFLUENCES. 189 of the greatest services rendered by the theory of se- lection, that it has finally broken with the notion of de- sign, which hitherto invested the organic world with perfection externally bestowed, and even in the province of intelligence and morality, where it is said with Schiller, So grows the Man as grow his greater aims,* has secured admittance for the uniform method of natural science. It is highly remarkable how the teleological view of nature could be so long upheld, and is still in part up- held, by theological influence although in the whole or- ganic world we behold a merely relative perfection, and the manifest and multifarious arrangements adverse to design in every grade of organisms, bear a bad testi- mony to the external directing Power. The perfection exhibited by comparative anatomy, and the estimate of physiological functions is, under all circumstances, the result of adaptation and selection. In the struggle of all against all, those individuals win who in any degree excel their fellows in the division of labour, which, if the direction of activity be altered, often obliges them to disuse organs which were once of service, but in the new conditions are useless, and, it may be generally said, have become injurious. Artificial selection—and here we may speak of design —produces perfection when, by mechanical and physio- logical labour (the latter especially by means of suit- able nutriment), it exercises the particular parts which are to be perfected, and propagates the advantages ob- tained. What we term natural selection is the epitome * Es wachst der Mensch mit seinen gréssern Zwecken. 190 THE DOCTRINE OF DESCENT. of the improvements acquired by specialization in the process of adaptation. The most faithful image of this gradually acquired specialization is afforded by the de- velopment of the individual, where from the undifferen- tiated, by constantly increasing differentiation, the mature animal is evolved in the plenitude of its physiological functions. That in the various animal groups certain grades of perfection are attained, is an uncontroverted fact; but every closer investigation shatters the idol of design. The organism of the bird might induce us to consider it, in the abstract, as modified for the purpose of flight. But if design be allowed to watch over the good flyers, the idea of design must be abandoned with respect to the non-flyers, and, if some idea is indispensa- ble, adaptation must have its due. Herewith the whole theory is broken down, and it will be the same in every other case. How organic perfection stands with reference to the idea of design, has been acutely and clearly expressed by the author of the “ Unconscious ” (“‘ Das Unbewussten ”). The theory of descent teaches that there is no inde- pendence of the conditions co-operating, in an organic phenomenon; rather that its increasing divergence from a common neutral point was an effect of the same causes. The theory of selection makes us acquainted with one of these causes, and unquestionably the most important as one, which, by purely mechanical compensative phe- nomena, produces advantageous results. The theory of descent merely casts doubts on the teleological principle by withdrawing the ‘basis for positive proof, but the doctrine of selection sets it directly aside, so far as it is able to extend its explanation. For natural selection ‘FITNESS WITHOUT DESIGN. IgI in the struggle for existence, the extermination of the less appropriate, and the survival and perpetuation of the fittest and most appropriate, is a process of mechanical causality of which the steady conformity to law is no- where infringed by any teleological controlling meta- physical principle. This, however, produces a result essentially corresponding to design; that is to say, it naturally bestows on organisms the highest capacity for life under given circumstances. Natural selection solves the apparently insoluble problem of explaining fitness as a result, without calling in the aid of design as a principle. In each family—for, as we have seen, what zoologists once designated type, has in the doctrine of Descent become the family—in each family lies the potentiality of a certain grade of perfection; and when the main outline of the family character is established, we see a development taking place, of which the potentiality is inherent in the tendency of the character, the realization and necessity in the external conditions. Hence to us also, progress is development, but not towards a pre- destined and pre-established harmony. Karl Ernst v. Baer,** anxious to rescue design, or at least the “ pur- pose ”—in short, predestiny, in the evolutionary series of Nature, says: ‘‘ Every cause engenders a process which again works on towards another purpose.” But why purpose? Ought it not rather to be: Every cause en- genders a process which again works on towards another process? The further we go back, the deeper and more general is the grade, and the various ramifications at their peripheral ends have either halted, or arrived at very different. grades. 192 THE DOCTRINE OF DESCENT. An objection frequently made against this result of the doctrine of Descent is, that if all are pressing for- ward towards perfection, how is it that, besides the higher, so many lower members of the family are able to main- tain themselves, and how can the lower families hold their own against the higher, in the struggle for exist- ence? In presence of the irrefutable facts of progress, it is enough to point out that the lower forms could and can continue to exist wherever they could find space as well as the other necessaries of life. While they here underwent only slight modifications, elsewhere the need- ful selection led to more profound metamorphoses; and on a subsequent geographical displacement, the newly transformed beings, accustomed to other conditions of ex- istence, were again able to share sea and land with the stationary species. For as diversity is restored now by selection, and the demands for nutriment and other neces- saries are likewise different, a partial remission in the struggle must take place. The preservation of a great many inferior organisms is evidently favoured by the circumstance that just be- cause they are simpler, their propagation is more easily effected. Hence although, especially in limited districts, amid violent competition of superior varieties, countless species must suffer extirpation, yet the struggle for ex- istence and.perfection do not exclude the existence of lower forms. But teleology, as it seems to us, still fails to explain what has béen explained by the theory of selec- tion. The retardation of the lower organisms, notwith- standing the internal pressure and the appointed purpose, is incomprehensible. But, it is frequently asked, if you will not hear of CHANCE. 193 a “principle of perfectibility” inherent in organisms (Nageli), of the “divine breath as the inward impulse in the evolutionary history of nature” (Braun), of “tendency to perfectibility” implanted by the Creator (R. Owen), even of the “ striving towards the purpose ” (v. Baer), can chance be supposed to have produced these marvellous higher organizations? To this it may be plainly answered, that this chance, to which purblind humanity allots so great a part wherever the personal interference of a superior Being or the universal “ crea- tive and productive principle” is not at hand, has no existence in nature, and that our conviction of the truth of the doctrine of derivation is due to its adjustment of the phenomenal series as causes and effects. Let us remember, and fancy ourselves in possession of, the formula of the universe of Laplace, by the aid of which all future evolutions might be computed in advance. With our limited powers, it is true, it is retrospectively alone that certainty can be approached in the calcu- lation and discrimination of the series. In this we must obliterate the word chance, for causality, as we under- ‘stand it, makes chance entirely superfluous. Any one who transports himself to the commencement of an evo- lution, who, for instance, fancies himself present at the genesis of the reptiles, may, from his antediluvian observatory, look upon the development of the reptile into the bird as a “ chance,” if he does not peradventure regard it as predestined. To us, who trace the bird backwards to its origin, it seems the result of mechanical causes. Let us now recapitulate what we have gained by the doctrine of Descent, based on the theory of selec- 194 THE DOCTRINE OF DESCENT. tion; it is the knowledge of the connection of organisms as consanguineous beings. The greater the accordance of internal and external characteristics, the closer is the kinship. The further we trace the pedigree to its origin, the fewer become the characters persisting to these roots, the more do these characters reveal themselves as acquisitions in the lapse of time. It is by eliminating these acquisitions and restricting the inherited charac- ters more and more as we feel our way backward, that _ we are enabled to reconstruct the pedigrees of the various groups.® We do the very thing which in linguistic inquiry is deemed extremely natural and scientific. The ideas and words common to the individuals of a linguistic family are the inheritance from the intellectual and linguistic property of the original people, from which the pedigree of the family has ramified. This so-called “ chance” prevailed in the formation of the derived languages neither more nor less than in the evolution of organisms from their original forms. IX. The Development of the Individual (Ontogenesis) is a Repetition of the Historical Development of the Family (Phylogenesis). ALTHOUGH the paleontological record is full of gaps, it is nevertheless unmistakable, as even most of the op- ponents of the doctrine of Descent are ready to admit, that from the older to the more recent period, a progress takes place from the lower to the higher grades of or- ganisms, which is likewise exhibited in the system of the present vegetal and animal world; and that in many ways embryonic development as well as metamorphosis and heterogenesis,—in a word, individual development (“ Ontogenesis,” Haeckel) suggests a comparison with these palzontological series, as well as with the sys- tematic order of succession. The parallelism of the palzontological and the systematic series is either a mir- acle, or it may be accounted for by the doctrine of Descent. There is no other alternative. And the doc- trine of Descent fully bears the test; it shows how the derivation of the present organisms from those pre- viously existing rests on the transmission of the char- acters of the progenitors to the offspring and the ac- quisitions of the individuals. The phenomena of indi- vidual development or Ontogenesis admit of no other choice; either they remain incomprehensive, or they 195 196 ‘THE DOCTRINE OF DESCENT. stand the test of the doctrine of Descent and submit to the great general principle. If we scrutinize the countless facts of reproduction and development, they certainly admit of classification; they range themselves in analogous and homologous groups; types of development become apparent; we speak of development without metamorphosis, of transformation, and heterogenesis. But what necessary relation the al- ternating forms, the shapes appearing in heterogenesis, bear to the complete animal or the sexually developed chief representative of the species? why so many animals undergo no transformations, but emerge ‘ complete” from the egg? why the species belonging to the same class or “type” possess the same type of development and process of construction?—these and similar questions as to the interpretation of the tangled mass of facts press themselves upon us. And they are also tests of our theory of derivation. The doctrine does as much as has been done by any great hypothesis in its special appli- cation; and if it gives a satisfactory reply to all, or at least to nearly all, pertinent questions, these are so many witnesses and proofs of its truth, which, according to all scientific custom and justice and philosophic method, will remain valid until the falsity of the inductions and inferences has been demonstrated and a better hypothesis substituted in its stead. The first proposition derived from the doctrine of Descent in explanation of the facts of individual de- velopment may run thus: accordance in the outlines of development is based on similar derivation; or, somewhat differently stated: accordance in the out- lines of individual development is accounted for by DEVELOPMENT OF ECHINODERMS. 197 similarity of derivation. As we already know, C. E. v. Baer first demonstrated that the members of the great divisions of the animal kingdom agreeing in the out- lines of their organization testify their coherence by a special “type of development.” This fact was always looked upon as self-evident, although, if it were not de- rived from descent, it would be the greatest miracle. This is therefore the place for us to review some of the fundamental forms of development which we partially considered in the third chapter, and at the same time to elucidate the meaning of these types with the aid of the doctrine of derivation. We will take the Echinoderm as our first example. Although from the anatomical comparison of a crinoid, a star-fish, a sea-urchin, and a sea-cucumber or holothuria, the close kindred of these various divisions of echinoderms is easily deduced, they yet deviate won- derfully from one another in outward shape and in the construction of the skeleton. The relative value of the difference between a holothuria and a star-fish, a sea- urchin and a comatula, may be compared to the differ- ence between a mammal and a bird, an amphibian and a fish. Nevertheless, with some few exceptions which have a special meaning, these various echinoderms leave the egg ina larval state almost identical. The larva (Fig. 12) is boat-like in form, with a curved lt iy margin bent over at both ends like a ‘Y= deck. This border is edged with a continuous row of cilia, by the agency of which the little boat is moved. A short digestive canal, provided with a gastric enlargement, is the first essential organ of this 198 THE DOCTRINE OF DESCENT. body. We will not describe the highly complex trans- formations of the larva here into an ophiura, there into a sea-urchin, and here again into a sea-cucumber; but we will only inquire what can be the cause of this accord- ance in the earlier stages of individual development. There is no reasonable answer but the derivation of all echinoderms known to us from an older form, in the development of which our larva likewise appeared, and from which this common phase of development was transmitted to the whole family. But it is allowable to ask, further, how from a bilateral larva, one, that is, symmetric on two sides, should be evolved in animals of radiate structure, as are the greater number of mature echinoderms? On this point Haeckel instituted a conjecture which at first exasperated the systematizers of the old school, but which now gains more and more footing, and is supported by the most recent comparative investigations, such as those of Hoffmann “On the Minute Anatomy of the Starfish” (“ Ueber die feinere Anatomie der See- sterne”). The boat-shaped larva of the Echinoderms, especially a modification occurring in the star-fish, strik- ingly resembles a certain larval type of the marine Annelida. And as in the structure and distribution of the parts of the rays of the echinoderms, especially of the star-fish, an unmistakable resemblance with the rela- tive distribution and succession of parts of the Annelids is observable, Haeckel regards the Echinoderms as an offshoot of the Annelids. He corisiders that the oldest, and to us unknown, echinoderms originated as annelid stems; the anterior end of the bilateral annulose parent-animal budding out gemmules in a radiate ar- DEVELOPMENT OF MOLLUSCS. 199 rangement. This gemmation, or, in other words, this stem structure, still occurs in Echinoderms, inasmuch as some species of star-fish possess such powers of repro- duction as to enable a single arm or ray, when torn off, to complete itself into a whole animal. Nay, Kowalewsky’s observations render it highly probable that the separa- tion of rays, and their completion by gemmation, is in some species a normal process. Haeckel’s hypothesis is thus laughed at only by those who are afraid to think or reason. In the family of the Mollusca, the so-called navicula larva testifies the kinship of at least two of the great classes. The third and most advanced class, that of the cuttle-fish, had perhaps lost their distinctive badge even in those primzeval times when, under the somewhat lower forms of the Tetrabranchiata, they left their shells in the Silurian strata. But the bivalve shells, or Lamelli- branchiata, and the snails, widely differing in anatomical development, and constituting two natural classes, have a common larval form, or, if the larvz display different shapes, a highly distinctive common larval organ, the velum. The accompanying diagram gives on the right the navicula of a cockle-shell as seen from behind. At the anterior end, two fleshy lobes have been formed, edged with cilia, by the vibrations of which the young animal, even in the egg, performs spiral twisting mo- tions; in the midst of the cilia rises a little prominence, furnished with a longer filament. These ciliated lobes or. vela, merging into one another, are shown on the left in the larva of a sea-snail (Pterotrachea), as seen nearly in profile, and in the phase in which the eyes and auditory apparatus, the foot and operculum, as well as a delicate 14 200 -THE DOCTRINE OF DESCENT. shell, have made their appearance. Here also, from the plane of the velum, a small fleshy protuberance juts out, without any special purport. The distribution of the velum, the period at which this larval organ makes its FIG. 13. appearance, its position towards the testa, head, mouth, and foot, and its subsequent effacement, one and all co- incide exactly in the two classes. It is as yet of only a relatively small number of marine shells and slugs that we know the evolutionary history; yet we may infer that in these animals remaining in their original home, this heirloom has been generally preserved. Even gen- era which in their mature state scarcely recall the type of the Mollusca, as the boring mollusks (Dentalium Tere- do), have preserved the phase of the navicula. On the other hand, in the branchiate fresh-water snails (Paludina) the velum is little developed, and in the land snails, which differ most widely from their marine kindred, the velum is entirely obliterated, as it is also among fresh-water mussels. If in these animals adaptation and migration to land has had this effect on embryonic and post- embryonic development, we must suppose that in OTHER FORMS OF DEVELOPMENT. 201 the Cephalopoda, notwithstanding their continued so- journ in salt water, other causes have produced the loss of the velum phase, and the course of development pe- ‘culiar to it. With respect to the other fundamental forms of de- velopment, we may refer to the third chapter. The con- struction of the higher Articulata points to annulose pro- genitors, more or less corresponding to the annelids of present times; and, again, the gradual increase of the segments of the larval annelids, which may be compared to the process of gemmation, leads from these higher Vermes to the lower ones with unsegmented bodies. All vertebrate animals, man included, if they do not pre- serve through life an unsegmented vertebral column, not separable into single vertebre, are raised as embryos from this condition into their higher and definitive phase. The fact that they pass through this common embryonic condition excludes all other mechanical causes but that of a common derivation from primordial forms which possessed an unsegmented vertebral column, no cranium or an imperfect one, and either no brain or one little differentiated from the spinal cord. Karl Ernst v. Baer, who, while we write these pages, raises his voice against the doctrine of Descent, has established the fact of types of development, and the course, within these types, from the undifferentiated to the special; but by the words “type of development,” the fact is para- phrased, not explained; and, as we cannot repeat too often, we prefer the distinct idea of derivation to the supposition of an unknown higher Power manifesting itself after an incomprehensible fashion in the types of development. 202 THE DOCTRINE OF DESCENT. If the concatenation of the series by direct derivation and heredity be disallowed, it is absolutely inconceiv- able‘ why the supreme creative Power, Nature, or the personal God, should have bound all higher animals to the same common stages of early development, and here- by exposed them to such manifold purposeless arrange- ments and great dangers. Of the millions of young oysters which annually escape from the egg, the ma- jority perish under the disadvantages of external condi- tions, because the oyster has not yet divested itself of the ancient heirloom of the roving navicula. It has been able to compete successfully in the struggle for existence, only because, like most of its congeners, it is enormously prolific. This may be understood; but that a personal Creator, merely on principle, in order to keep the oyster within the type of development, should have. endowed it with the phase of the navicula, in this case so extremely unpractical, can be accepted, like much other nonsense, only as matter of faith. If accordance in the outlines of development has gen- erally shown itself derivable from similarity of descent, we may now proceed to the explanation of those phe- nomena of development known to us as heterogenesis and metamorphosis. In these, the historical stages of development of whole classes and orders are inherited in the development of the individual; a proposition which is merely the corollary and application of what has been already intimated. The Hydromedure offer numbers of such instances of heterogenesis, of which Gegenbaur’s early views gives the following account: We have already (p. 43) become acquainted with the origin of the Cladonema from the polype-like Stauridium. The ALTERNATION OF GENERATIONS. 203 Medusa is the sexually mature form of the cycle of the species; its ova develope into polypes, which constitute the intermediate form in their develop- ment; that is to say, it is not transformed into the FIG, 14. animal from which it is derived, but produces gemmules. Only in this generation does the species revert to the sexual form. We shall understand this alternation of generations if we begin with the simplest Medusa polypes. Such a one is the annexed Hydractinea carnea, of which the female individual is portrayed. Compared with the intermediate form, Stauridium, the preliminary phase of the Cladonema, reproducing itself asexually, 204 THE DOCTRINE OF DESCENT. the Hydractinea seems superior, inasmuch as it is itself a sexual form. The zone of spherical protuberances in the middle of the body are the ovaries or egg capsules corresponding to the sperm capsules of the male indi- vidual. Heterogenesis does not take place in our Hy- dractinia, but, as in the development of the ovum of the Cladonema into the Stauridium, there is a transformation of a ciliated lava into a sessile polype. But it is obvious that the part which in the Hydractinia is played, by the male and female sexual organs is per- formed in the generative cycle of the Cladonema by the sexual animals. By following the transition from the de- pendent organ into the independent animal, we find the solution and ex- planation of the process termed he- terogenesis. Between the genera reproduced like the Hydractinia, and those reproduced like the Cladonema, there are many others, of which the propagation shows the gradual transition of the rudimentary sexual organs into the sexual animal. We may so arrange the genera of the “ Medusa polypes” as to exhibit how the parts which in the Hydractinia are mere capsules, generating and enclosing the ova, become more and more perfect. They acquire a special branch of the alimentary canal and blood-vessels, and are provided with the marginal papilla characteristic of the Medusz, and constituting their peculiar sensory organs. In short, what in one member of the systematic series may be termed an or- gan, is, in the next, the Medusa separating itself and FIG. 15. PARASITIC WORMS. 205 becoming a new generation; the sexual organ has be- come the sexual animal. Now as the individual development of the Cladonema, and other Meduse similarly propagated, corresponds with the systematic series of the Medusa polypes, the only reasonable and creditable explanation of the onto- genesis of those Medusz in which heterogenesis occurs, is that, in them, the historical development of the genus has become fixed. Neither the egg nor the hen was created. Before the delicately tinted Medusze populated the primzeval ocean in lonely splendour, the Medusa. polypes on the constantly changing shores were the sole representatives of the still infant class. Another and. very reasonable view is that which Weismann’s latest re- searches tend to support: That polypes were indeed the precursors of the Medusz; that individual polypes (not: merely organs) became Medusz, and that the im- perfectly developed intermediate forms were degener- ate Medusa, degraded to the condition of apparently mere organs. The evolutionary history of the intestinal worms leads to the same reflections and results. These animals, widely. differing in their structure, were either created in or with their hosts, or else they have become habituated to them in a natural and direct manner. We may surely disregard the third alternative, that they were led by an innate “ obscure impulse.” According to our doctrine, the worms now passing the whole or a portion of their lives as parasites on or in other organisms, are descended from free and independent animals, and the periods oc- curring in their development, during which parasitic life is exchanged for independent phases, signifies a rever- 206 THE DOCTRINE OF DESCENT. sion taking place systematically in all individuals to the once permanent condition of their progenitors. Of the Trematoda or Flukes, and Cestoda or Tapeworms, be- ‘longing to the class of the Platelmintha Suctoria, the latter have diverged the most from their starting-point; their adaptation to life within other animals has rendered the alimentary canal superfluous, and their generations and transformations hence point less to their progenitors than is the case with a number of other Trematoda, with which many anatomical characters prove them to be closely related. Both, moreover, share the characters of their class with the free-living Turbellaria. From such as these, that is to say, from forms approximate to the present Turbellaria, the Trematoda and Cestoda must be descended, and with this agrees the free roving phase which the larva of the Fluke (Distomum) undergoes as the so-called Cercaria, and previously as a rotating spher- ical body. Many of the ciliated Nematoids, or thread-worms, too,—the division which includes the Ascarides among others,—have in their infancy a stage of independent life, during which they cannot be distinguished from the infantine forms of their more numerous kindred, which never adopt a parasitic life, and chiefly inhabit the sea. The transition to parasitism, as recapitulated by ontogenesis, was nothing more than an extension to a new territory offering advantages of nutriment; and on this point it is highly instructive to compare the Nematodes with the systematic series of the leech- like Suctoria (Trematoda), so excellently described by Van Beneden. We here find all the transitions from independent predatory genera to others occasion- PARASITIC CRUSTACEA. 207 ally parasitic, and again from these to others which on leaving the egg immediately attach themselves for life. Here, as elsewhere, parasitism seems an adaptation to new habitats, which is recorded in the biography of the individual with a reminiscence of the previous form. The circumstances of the parasitic worms are repeated by the parasitic Crustacea, as, moreover, a probably primordial form of the crab family is preserved in the metamorphoses of several orders of this large and diversified, though coherent class. The larva, which, it may safely be assumed, approximates closely to the primordial form, was at one time taken for an independent genus and re- ceived the name of Nau- plius. Hence a Nauplius phase is spoken of, which obtains especially among the lower Crustacea, the Copepoda, parasitical Crus- tacea and Cirripedes, and the remarkable Rhizopoda connected with them; but is not wanting in the highest order, the decapodous stalk-eyed crab. We shall later have to make ac- quaintance with the so-called curtailed development which among the crabs has been adopted by the decapods, and it was formerly supposed by all. Were this actually the case, we should still, by analogy, infer their connec- tion with the other orders repeating the Nauplius phase in the course of their development; but it was a welcome discovery of Fritz Miiller’s that a shrimp (Peneus) still FIG, 16, 208 THE DOCTRINE OF DESCENT. begins its development as a Nauplius; whereas all the other members of the order, as far as they are known,. leave the egg in the higher Zoea phase (p. 50). As of the hundreds of stalk-eyed crabs, scarcely a dozen have been hitherto examined as to their development, it will not be doubted that, with regard to the Nauplius phase, some resemble the Peneus of the Brazilian coast. But Fic. 17, Axolotl, even were this case to prove unique in the order, it would suffice as a living witness of the connection be- tween the presence of the decapods and the primordial crabs. There can be no other view of this subject. The Nauplius phase in the development of the Peneus is either a shining testimony in favour of the doctrine of Descent, or a senseless paradox. After what has gone before, the transformation of the AMPHIBIANS. 209 Amphibians needs no elucidation. Their predecessors were water-breathers, whose form and mode of life are more faithfully preserved by the long-tailed Amphibians, the tritons, and salamanders, than by the frogs. In our tritons, sexual maturity not rarely commences in the larval state, hence in a phase which was definitive in the progenitors of the present genera. There is, in- deed, one species, the Mexican Axolotl, which normally propagates itself during the larval phase. Auguste Dumeril’s observation is highly interesting, that of the FIG. 18,—Amblystoma. thousands of Axolotls that he bred at Paris, some few advanced beyond the grade of development hitherto known in them, #.e. they lost their gills, changed the shape of their bodies not inconsiderably, and from gill- 210 THE DOCTRINE OF DESCENT. breathers, and aquatic animals became lung-breathers and terrestrial animals. It needs further observation to ascertain whether (what is, however, very improbable), in their home, all Axolotls, after having propagated them- selves in their larval state, undergo the_ metamorphosis into salamander-like animals (Amblystoma), or whether the transfer to Europe and the consequent entire change of the circumstances of life gave the impulse to a pro- gressive transformation of these few individuals, which, by the continuance of these conditions, would in future generations extend to more and more individuals, and finally become the characteristic of a new species. The examples of Ontogenesis, or individual develop- ment, hitherto examined, had the peculiarity that the sexual animal does not issue directly from its egg like the Phoenix from its ashes, but had to pass through various forms and existences in which the progenitors of the species again become alive and palpable. We must now inquire how this development is related to that form of reproduction which the systematizers, completely in accordance with the facts, yet without any correspond- ing meaning, have termed “direct development,” or “development without heterogenesis or metamorpho- sis?” The ciliated embryos of many Medusz are not converted into polype-like intermediate forms, but pass directly into Medusz. The greater number of higher crabs do not leave the egg as Nauplia, but as more or less perfect decapods. The bird, the mammal, and man are all at birth “similar to their parents.” Consider- ing that the processes of heterogenesis are in themselves by no means advantageous to or “in harmony with design ”—we have only to remember the fate of the tape- DIRECT DEVELOPMENT. 211 worm’s eggs—that by the larval state the period of in- fancy and weakness is prolonged, and the period of ma- turity and efficient care for the continuance of the species delayed, it follows that curtailments and reductions, con- sequent on adaptation have, ‘as advantageous modifica- tions, a prospect of perpetuation. As in Amphibians the prolongation of the larval phase may be effected by natural circumstances and artificial experiments, so in like manner a compression of the phases of transforma- tion, and a general curtailment of the metamorphosis is imaginable. In the class of Amphibians we have, in fact, several examples of curtailed and modified meta- morphosis which bridge over the apparent chasm be- tween development with and without transformation, and render direct development comprehensible as being gradually acquired. Amphibians will endeavour to ex- tend themselves wherever they are invited by a sufficient supply of insects, and the black salamander of the moun- tains (Salamandra atra) has even overcome the impedi- ment which might have been deemed insurmountable, the absence of water for its larvee. It does not lay its eggs like its congeners, but only two are received into the oviduct, and the fluids secreted from its walls replace the marsh to them and to the larve which emerge from them. Here, and not when separated from the parent, do the gills make their appearance, while the other eggs, gradually following, are devoured by the hungry larve. The metamorphosis of the black salamander, which has lately been the subject of Fr. Marie Chauvin’s remark- able experiments, can be effected after exclusion, and observation places it beyond a doubt that this is an in- stance of adaptation to unusual conditions. . If the mode 212 THE DOCTRINE OF DESCENT. of life of the marsupial frog, which carries its young in a membranous fold of the back, and the Surinam toad, of which the larve live singly in the chambers of a kind of honeycomb on the back, were better known than they are, we should assuredly arrive at the same results as with the black salamander. In the absence of other knowledge, the observations of M. Bavey, Marine Pharmaceutist at Guadaloupe, first published in 1873, are of the highest importance.** A trog of those parts (Hylodon Martinicensis) goes through its whole metamorphosis in the egg. In the egg it has gills and tail; and from the brief remark that the island con- tains only rapid running streams, and nowhere stag- nant waters or marshes, it appears that this is also a case in which adaptation modifies and curtails devel- opment. If, after this introduction, we now examine the so- called direct development with more attention, it may in every way be compared to the metamorphosis of the Hylodes of Guadaloupe. Direct development is a trans- formation in the ovum; and in the cases in which it oc- curs, the phases of embryonic development are repeti- tions, more or less distinct, of the historic development of the family. We will only particularize in the embry- onic life of the Vertebrata (in which metamorphosis does not take place), some phases that are stages of curtailed transformation, and recapitulate the permanent condi- tion of their progenitors. It has been repeatedly men- tioned that in all vertebrate animals, the vertebral column is first laid out as an unsegmented cord and an unseg- mented sheath for the spinal cord. This is the permanent state of the lower fishes. In the higher Vertebrata also, DEVELOPMENT OF AMMONITES. 213 the brain at first consists of vesicles, lying one behind the other, which is the persistent form of the lower groups. The embryonic heart of mammals and birds begins in the form of a tube, and subsequently ac- ‘quires the communications between the chambers, which in the reptiles never close. In the Amphibians, the branchial arches really bear gills during the lar- val state. They are not wanting in the embryos: of reptiles, birds, and mammals, any more than the fissures through which, in fish and the larve of Amphibians, the water passes off after being inhaled. Must we again set forth the only possible explanation of these facts? Before referring to the phenomena which testify the emanation of families from a common root, we will cite one of the most important evidences of recent times, which traces the genesis of species through a great geological period, and exhibits in detail the relations of the development of the individuals to that of the species, -genus, and family. We mean L. Wiirten- berger’s contribution to the geological evidence of the Darwinian theory, to which we have already appealed ‘(p. 97). It relates to the two families of Ammonites, the Planulata and Armata; of which, according to Wiirten- berger’s researches, the latter are developed from the former, as the ribs of the Planulata gradually pass into the spines of the Armata. Of special interest to us are the following passages of the preliminary communication on the discoveries obtained from thousands of specimens, and which will probably not be made public, with all the vouchers, for some years to come. “It gave me particu- lar pleasure,” says Wiirtenberger, ‘‘ when, after divers 214 THE DOCTRINE OF DESCENT. careful comparative studies, I at last detected an inter- esting and simple conformity to law in the variations of the Ammonites. Namely, on the first appearance of a modification which subsequently attains essential im- portance in an entire group, it is only slightly indicated on a portion of the last convolution. Towards more recent deposits, this modification is more and more plainly shown, and then advances, following the spiral course of the shell; that is to say, it gradually takes possession of FIG. 19.—Ammonites Humphresianus. A form analogous to the Planulata. the central turns also, as we trace the forms to higher strata. This reproduction in younger stages of life of modifications first occurring at a more advanced age, makes but slow progress, so that we see the older forms repeated with great persistency in the central turns. Fre- quently a modification of this sort has taken possession of only a small part of the convolutions, when a new one already appears at the outside, and follows the first. Thus searching through the strata from below DEVELOPMENT OF AMMONITES. 215 upwards, we see modification after modification begin- ning at the outer part of the Ammonites, and advancing towards the centre of the discs. The innermost convo- lutions often resist these innovations with great per- sistency, so that we usually find upon their surface sev- eral of these states of development in close juxta-position, as the shell of the individual Ammonite begins with the old morphological type, and then adopts the modifi- cations in the same order in which they follow in vast periods in the geological development of the groups con- cerned.” “The Ammonites,” he says moreover, “thus obtain at an advanced and maturer age—only when they have gone through the development inherited from their parents, and as much as possible in the same manner as their parents—the power of modifying themselves in a new direction, that is to say, of adapting themselves to new conditions; yet these modifications may then be: transmitted to the offspring, so as to appear in each subsequent generation a trifle earlier, until this phase of development in its turn characterizes the greater portion of the period of growth. But this last. and longest. phase of development scarcely ever suffers itself to be supplanted by new ones, formed in like manner; heredity operates so powerfully, that a period of development thus once predominant, is repeated in the infancy of the Ammonites, even though but slightly indicated. Hence in an individual Ammonite from a recent stratum, the periods of development compressed and forced back upon the innermost convolutions, must appear in the same succession in which they. wrested the dominion from one another. It is extremely interesting to study 15 216 THE DOCTRINE OF DESCENT. the development of the Inflata of the upper white Jura (which follow the Ammonites liparus, whose externally visible convolutions display only one row of spines), carefully breaking off convolution by convolution. Towards the middle there is a region in which there are always two rows of spines; nearer the centre the inner- most row disappears; soon afterwards the outer one also; and the nucleus, some millimetres in diameter, now ap- ‘pears for about half a turn as a Planulatum, with dis- tinct ribs, which, towards the beginning, likewise disap- pear. Thus even the Planulate ribs, which prevailed among the Liassic ancestors of these Inflata, and were supplanted by the spines as early as in the brown Jura, still distinguish these later and essentially modified de- scendants during a short period of their youth.” Wiurtenberger further shows how these relations can be simply ex- plained by the Darwinian theory alone; “without it we should have only an extraordinary problem.” It was natural to test the applica- bility of the theory of selection also on the forms alliedto the Ammonites, such as the Ancyloceras; namely, the genera in which the convolutions do not touch and partially conceal one another, as in genuine Ammonites, and which, as late comers and side L shoots of the group, seemed des- FIG. 20.—Ancyloceras. —_ tined to decay. Selection and decay? Wirtenberger shows how the abandonment of contact in the convolutions was to the spinous Ammonites .an FORMS ALLIED TO AMMONITES. 217 advantage which would be established by selection. If other paleontologists consider the fluctuations of form accompanying the relaxation of the closed spiral as evinc- ing the decline of the group, no contradiction seems to be implied, for what was originally used as an ad- vantage by natural selection, proved injurious in its con- sequences. As we have seen, the earliest states are obliterated to WN » (of 00 i Ty-4 e SS i =a FIG. 21. such a degree by curtailment of development that the indication of the nature of the progenitors continually diminishes. But our theory necessarily leads to the conviction that the families within which we have as yet been able to compare Ontogenesis with Phylogenesis, constantly approximate in their origin, and vindicate the 218 THE DOCTRINE OF DESCENT. expectation that at least here and there, in the indi- vidual development of single representatives of the vari- ous families, witnesses of their common derivation should come to light. This likewise occurs, and to such a de- gree that in the earliest larval stages a link is estab- lished between the lowest and the highest animals. If a number of groups of the lowest living beings, in which the various vital functions of nutrition, irritability, mo- tion, and reproduction are supplied by amorphous proto- plasm,—if these be separated, as by Haeckel, into a neutral kingdom, owing to the absence of sexual repro- duction, we must likewise agree with him in attributing to the Spongiadz ranking next to the Protista, the name of animals, on account of their sexual propagation and the nature of their embryonic development and first larval phases. Haeckel has bestowed on one larval stage of the calcareous sponges the title of Gastrula, wherein the animal represents a sac, or, in other words, a stomach provided with a mouth-like orifice. The walls are formed of two rows of cells, the outer one consisting of ciliated cells; that is to say, each cell is furnished with a long filament. At the orifice of the sac, the outer row merges into the inner one, and from these two mem- branes the body of the sponge is constructed in a definite manner. Now, if this Gastrula larva reappears in the Coelenterata, Polypes, and Medusz, in which the gradual development from the two membranes, the entoderm and ectoderm, into the most complex forms has long been known; and if, as Haeckel has further shown, the osculum, or larger opening of the spongiadz may be closely compared with the mouth of the polype and GERMINAL MEMBRANES. 219 medusa, and the great central cavity of the sponge with the stomach of the others, of the canal system with the canals and cavities of the Ccelenterata,—then, in combination with the host of other facts, implying and supporting the doctrine of Descent, the inference is in- evitable that in the Gastrula we have a testimony of the consanguinity of the Spongiade and Ccelenterata. But this Gastrula reappears in the Holothuria; hence in the Echinoderms, in the Sagitta, in the Ascidians, which will be more narrowly examined in the pedigree of the Vertebrata, and finally in the Lancelet; and we, there- fore, hold ourselves justified in regarding this coinci- dence of the earliest states of development in different families, as the remnant of the common root, which in other families, as in the Articulata, for example, has been lost in the curtailment of development. The significance of the “germinal membranes” in the Vertebrata was recognized even by Pander, and in the suggestive works of v. Baer; the extension and application of this observa- tion to the whole animal kingdom, for which we are especially indebted to Kowalewsky, marks one of. the greatest advances in the science of comparative develop- ment. The reader unacquainted with the detailed researches of our science, has already been called upon to observe that there are opponents of the theory of selection, such as Owen, who nevertheless accept the doctrine of Descent as incontestable. Even rejecting natural selection, the parellelism of Ontogenesis with Phylogenesis may also be brought into the natural connection maintained by us, on the assumption of an unnatural or supernatural guidance which converts this apparently natural unity 220 THE DOCTRINE OF DESCENT. into a miracle. Quite recently, A. Braun has pointed out the accordance of the botanical system, and therewith of paleontological succession, with the development of the individual plant, when he says: *7—‘In the further elaboration of the natural system, the gradation of the vegetal kingdom, and, at the same time, the relation of the system to the history of development, becomes more and more spontaneously and incontrovertibly manifest. The Acotyledons are verified as Cryptogams, as they were already considered by the old botanists of pre- Linnen times, and their relation to the Phzenogams is thus more clearly pronounced. The Cryptogams are separated into two essentially different divisions, in which gradation is likewise distinctly pronounced (cellular and vascular Cryptogams, Thallophytes and Kormophytes); between the perfect Phaenogams and the Cryptogams an intermediate grade has been shown, that of the Gym- nosperms. But most important of all is the circumstance that the four chief grades ascertained in the vegetal king- dom accurately correspond with the grades of develop- ment occurring in the individuals of all the higher plants; —the germ, the vegetative stem, the blossom and the fruit.” But why this parallelism is to be most important of all, if it is not to lead us to the knowledge of true causality, is beyond our comprehension. We can well imagine that the “inherent causes” and the “ Principle of Perfection” may be welcomed as the refugium ig- norantie, but not that they can really satisfy inquiry. For our own standpoint, the accordance oftheresults of botan- ical investigation must be extremely important, but it is for the palpable reason that the theory thereby gains the support and corroboration of another great series of facts. THE BEGINNING OF LIFE. 221 If the accordance of the evolution of families has once been followed up to the Gastrula, we shall not pause there, but must regard the similarity of the sperm cor- puscules and germ cells from the Spongiadz to the Ver- tebrata as a primordial common property, connecting the animal and vegetal world; and prior to the acquisi- tion of which, only those modes of reproduction took place which have been maintained among Protista and in heterogenesis. As the common basis of sexual reproduction in the various families argues a common origin, asexual re- production, directly connected as we have seen it to be with sexual propagation, by means of unfecundated eggs and germs, leads us constantly further towards the be- ginning of life. But the cell furnished with a nucleus and sheath is inseparable from the protoplasmic cor- puscule devoid of nucleus or sheath, on the growth and fission of which rests the reproduction of the lowest liv- ing beings. Their origin from inorganic matter, as we have set forth above, is a postulate of sound human understand- ing. To this beginning we are led, not, as the opponents of the doctrine of Descent are wont to say, by a dog- matic after-philosophy, but by the unprejudiced consid- eration and computation of the facts of individual devel- opment.*® 222 THE DOCTRINE OF DESCENT. X. The Geographical Distribution of Animals in the light of the Doctrine of Derivation. ALTHOUGH ever since the century of the great geo- graphical discoveries, material has been accumulating for a geography of plants and animals, the foundations of scientific botanical geography (apart from George Forster’s observations) were first contained in Hum- boldt’s celebrated “ Ideas on the Physiognomy of Plants” (Ideen zu einer Physiognomik der Gewachse). It is the first description of vegetal forms, comprising the entire area of the earth, and the manner in which, singly or combined, they lend a characteristic impress to the land- scape of their region of distribution, and again on their side harmonize with the other factors of the scene. The celebrated founder of Climatology, who circled the ter- restrial globe with lines of equal temperature, of equal inclination and declination of the magnetic needle, and divided it into dry and rainy zones, knew better than any of his contemporaries that the animal and vegetal world depended on all these factors. Yet neither he nor his followers, before Darwin, rose higher than the description of Nature, which had already checked Buffon in his grand picture of Nature, “Les Epoques de la Nature.” A natural result of the extraordinary extension of VICARIOUS FORMS. 223 the geographical horizon and the profundity of special research was the more careful ascertainment of .the regions of distribution of animal and vegetal families, and of their more prominent species, in which, as we have already said, either no questions were asked as to the causes of distribution, or the matter was facilitated, as by Louis Agassiz, who did not, like Linnzus, derive each species from a pair, but supposed them to be cre- ated in suitable numbers of individuals in their own regions of distribution. It cannot be expected that any solution was hereby given to the questions which now force themselves upon us, such as why, under like natural conditions, like species are not always to be found, and conversely? Why very similar species frequently appear under external conditions entirely dissimilar? What is to be thought of the mutual relations of the so-called vicarious forms? &c. As Riitimeyer has recently observed, in his excellent treatise “On the Derivation of the Animal World of Switzerland ” (“ Ueber die Herkunft der schweizerischen Thierwelt ” *°), Buffon had already remarked the repeti- tion of the African in the American fauna; how, for ex- ample, the lama is a juvenescent and feeble copy of the camel; and how the puma of the New represents the lion of the Old World. Still, by the mere word “ repre- sentative” or “vicarious form” nothing is gained, and a true apprehension of these facts is obtained singly and solely if we meet the inquiry with the assumption that camel and lama, puma and lion, are of common deriva- tion, and that their diverse development was in the lapse of time favoured and determined by the separation of the habitats of their progenitors. 224 THE DOCTRINE OF DESCENT. Another example of so-called vicarious or “ analo- gous” species, affording an easier basis for induction, is provided by the comparison of the snails of Southern Europe, and especially of Spain, with those of North Africa, on which we are indebted to Bourguignat for some excellent observations. In accordance with other botanical and zooligical facts, he has established that the shell fauna of Spain and North Africa forms a whole, so that the Algerian snails appear a mere appendage to those of Southern Europe, notwithstanding the separa- tion by the Straits of Gibraltar. Now it is proved that, in geologically recent times, this region of North Africa was in fact a peninsula of Spain, and that its union with Africa was effected on the north by the rupture of the Straits of Gibraltar, and on the south by an up- heaval to which the Sahara owes its existence. The shores of the former Sea of Sahara are still marked by the shells of the same snails that live on the shores of the Mediterranean. But all North African species are not identical with those of Spain; of many African sorts, only “ analogous” species are found on our side. Now if certain Spanish species do not themselves occur in Africa, but are yet replaced by very similar forms, our standpoint at once connects with the otherwise unmean- ing word “analogous” species the idea of the common derivation of the forms replacing one another, and of the local variations superinduced by isolation and altered conditions. A severe test is applied to those who believe that species were separately created, by the air-breathing land snails (pulmo-gasteropoda), when it is seen that in isolated islands and island groups these earth-bound ani- ANALOGOUS FORMS. 225 mals, migrating with so much difficulty, have attained an extraordinary diversity. In the Madeira Islands, 134 species of pulmo-gasteropoda were reckoned about ten years ago, of which only 21 were to be found in the Africo-European fauna. These and the 113 other spe- cies are mostly confined to narrow districts and single valleys. Are we to suppose that the 113 species for Madeira, and the 21 species for Madeira and Africa with Europe, were each separately created? Must we not much rather infer that a connection at one time ex- isted between Europe and the present island group of Madeira, and that these 21 species remained what they were before the separation; while from unknown species still appearing in analogous forms upon the continent emanated the remarkable profusion of new species? They, and their comrades on other isolated islands, were spared a conflict many sided, and they doubtless af- ford a favourable example of Wagner’s law of mi- gration, as with the difficulties of locomotion, and the improbability of a large subsequent arrival, the se- cluded individuals, under even slightly different influ- ences, had had a prospect of diverging from the parent species. The unscientific opinion, that under like, or nearly like, ‘external conditions, like or similar organisms were created in great numbers, receives a severe blow by the perception that the direct reverse has frequently occurred. Why has America no horses in the present era, although it is proved that the horses introduced, thrive capitally? It is not necessary for us to explain why the fossil horses which existed in America, as well as in the Eastern hemisphere, became extinct without 226 THE DOCTRINE OF DESCENT. leaving any progeny—we do not know the cause, though we may yet be able to fathom it; but in this and all similar cases the adherents of the doctrine of Creation must confess the inadequacy of their theory of belief. Our exposition has shown that the species now extant are the progeny of organisms previously existing; the present apportionment on the earth is therefore a con- sequence of the distribution of the progenitors of the present organisms, and of the manifold displacements of land and water by which they were indirectly or di- rectly affected. We cannot hope ever to picture to our- selves a faithful representation of the perpetual transfor- mations of the surface of the earth. Only, if this could be accomplished, and if we, moreover, had an accurate register of the animals at each period inhabiting the former islands, continents, and oceans—only then could the distribution of the present organisms be thoroughly fathomed and established. But in thus acknowledging the incompleteness of our statistical means, we are at least able to lay down with certainty the course of in- quiry. We must, in the first place, proceed in the method of the older vegetal and animal geography to ascertain the natural limits and regions of distribution; and, secondly, to collate these facts with the facts. of the distribution of the former progenitors of the present animate world as it was determined by the geological conditions of those times. It is needless to say that Darwin has furnished the outlines for this work also. But among his followers two are specially worthy of dis- tinction: Wallace, with his researches on the Malay Archipelago,” abounding in subtle observation; and DISTRIBUTION OF ANIMALS. 227 Riitimeyer, in his treatise already cited. In what follows we may essentially adhere to the latter. Our knowledge of the regions of distribution of the animal world is still extraordinarily deficient. What do we know, for instance, of the occurrence of marine ani- mals? Few years only have elapsed since the depths of the sea were rendered accessible to research, and the result has almost entirely upset our earlier notions of the geological significance of the sea-bottom and its habitability. After the strong impulse given by Maury to the investigation of the physical condition of the sea, we are now occupied in ascertaining the submarine tem- peratures and currents, the constitution of the sea-bottom, the occurrence of deep-sea organisms, and the conditions of their existence. We are therefore just beginning to collect the material for a future geography of marine organisms. Among terrestrial animals, certain groups of which the actual distribution can be defined, are use- less for our general purpose. Butterflies, for instance, which are an easy prey to currents of air, defy geological barriers, and, above all, that important partition which from the tertiary era has been erected, or rather excavated in the bottom of the sea, between Australia and India.71 It is the same with bats, and also with migratory, predatory, and aquatic birds; while, as Wallace shows, the other orders of this class are in tropical regions very reliable and stable in- habitants of their often limited districts, seemingly sug- gestive of migration. Exclusive of these, there remains therefore little more than the Mammalia, whose extrac- tion may be inferred with certainty from a comparison of their present cantonments (Cantonirung),—an expres- 228 THE DOCTRINE OF DESCENT. sion which we borrow from Ritimeyer,—with the en- campments of their former kindred, whence are derived general points of view as to the causes of the present geographical apportionment of organisms. If in the preliminary establishment of facts we there- fore confine ourselves to the Mammalia, exclusive of whales and bats, a superficial survey is enough to show that not only single species, but families also, have each a certain region of greatest density of occurrence, a focus of distribution, and that from thence radiations have taken place according to the convenience and fit- ness of the territory. Lion and tiger, elephant and camel, range over a definite era; the monkeys of the New World differ from those of the Old World not only geographically, but also in family characteristics. Mar- supials are chiefly concentrated in Australia; sloths and armadilloes in South America. And these examples, easy to multiply, indicate how individuals of widely dis- persed species, and the species themselves, emanated from single points of the earth’s surface and flowed over the territory of distribution now occupied. When to this observation is added the other, that in past eras also the same groups had the same centres of distri- bution,—for instance, Brazil not only harbours sloths and armadilloes now, but was once peopled by more numerous and partly colossal species of these families, and Australia has furnished the most numerous and im- portant fossil remains of Marsupials,—the cognizance of this persistent localization becomes very significant, and we account for the “repetition” of these forms by de- rivation. Now if the centres of distribution, at the first glance ORIGIN OF ISLANDS. 229 extremely numerous, can be brought into closer union and reduced to the smallest number possible, as by our theory the Mammalia have but one point of derivation, and if we can herewith harmonize the geological succes- sion of the organisms examined, or, in other words, harmonize the horizontal distribution with the vertical or historical sequence, animal geography will then ap- proach the solution of its task. Wallace and Ritimeyer’s works are therefore an important advance, as the former has given detailed evidence that the fauna of the com- plex and extensive Australio-Indian Archipelago is by no means self-dependent, but consists merely of off- shoots of the continents; and the latter, in a grand sur- vey of the entire surface of the earth, has reduced the centres of distribution to the simplest proportions as yet possible. The comparison of insular and continental faunas is naturally of great interest. For should it appear that, with respect to the animal world, islands are one and all mere appendages of the continents, the problem would at once be vastly simplified. If we follow Peschel’s luminous exposition of the origin of islands,’? we have first to deal with the fragments of continents. A great number of islands, such as Great Britain and the great Asiatic islands, may be recognized at once as fragments of still existing continents. On the other hand, Mada- gascar and the Seychelles are not, as might be con- jectured, a segment of Africa, but the remnant of a former continent very peculiar in its flora and fauna. Other islands originate either from submarine volcanoes or from corals, and in the latter case the structure is founded on sinking land. It naturally follows that on 230 THE DOCTRINE OF DESCENT. volcanic and coral islands only such animals will be en- countered as reached them by swimming or flying. The presence of Mammals pre-supposes human agency or extraordinary accidents. The older the islands, the richer are they ini organisms. Islands detached from continents will, on the contrary, be rich in proportion as they are recent, of which Great Britain bears witness. The more divergent is their fauna, the longer must be the time which has elapsed since their separation. Thus, for instance, we may view the relations of Tasmania and Australia; and if New Zealand was ever connected with the old Australian continent, the separation oc- curred at an epoch so remote that it throws no light upon the physiognomy of the animal world of New Zealand, and vice versa. In the account of his travels in the Malay Archipel- ago, Wallace has given a pattern of animal-geographical research. Years before, G. Windsor Earl had pointed out that the great islands of Sumatra, Borneo, and Java, are connected with the Asiatic continent by a shallower sea; while a similar shallow sea assigns New Guinea and several adjacent islands to Australia, with which they have a common characteristic in the Marsupials. Wallace has defined this partition more minutely with a line marked by a deeper submergence of the sea-bot- tom. It is drawn below the Philippine Islands, and, having Celebes to the south, passes through the straits of Macassar and separates the two small islands of Bali and Lombok. We will now follow Wallace’s description (“Malay Archipelago”), with various omis- sions. “Tt is now generally admitted that the present dis- MALAY ARCHIPELAGO. 231 tribution of living things on the surface of the earth is mainly the result of the last series of changes that it has undergone. Geology teaches us that the surface of the land and the distribution of land and water is every- where slowly changing. It further teaches us that the forms of life which inhabit that surface have, during every period of which we possess any record, been also slowly changing. As to the Malay Archipelago, we find that all the wide expanse of sea which divides Java, Sumatra, and Borneo from each other, and from Malacca and Siam, is so shallow that ships can anchor in any part of it, since it rarely exceeds forty fathoms in depth: ‘and if we go as far as the line of a hundred fathoms, ‘we shall include the Philippine Islands and Bali, east of Java. If, therefore, these islands have been separated from each other and the continent, by subsidence of the intervening tracts of land, we should conclude that the separation has been comparatively recent, since the depth to which the land has subsided is so small.—But it is when we examine the zoology of these countries that we find what we most require—evidence of a very striking character that these great islands must have once formed a part of the continent, and could only have been separated at a very recent geological epoch. The elephant and tapir of Sumatra and Borneo, the rhi- noceros of Sumatra and the allied species of Java, the wild cattle of Borneo and the kind long supposed to be peculiar to Java, are now all known to inhabit some part or other of Southern Asia. None of these large ani- mals could possibly have passed over the arms of the sea which now separate these countries, and their presence plainly indicates that a land communication must have 16 232 THE DOCTRINE OF DESCENT. existed since the origin of the species. Among the smaller mammals, a considerable portion are common to each island and the continent; but the vast physical changes that must have occurred during the breaking up and subsidence of such extensive regions have led to the extinction of some in one or more of the islands, and in some cases there seems also to have beén time for a change of species to have taken place. Birds and insects illustrate the same view, for every family, and almost every genus of these groups found in any of the islands, occurs also on the Asiatic continent, and in a great num- ber of cases the spccies are exactly identical. Birds of- fer us one of the best means of determining the law of distribution; for though at first sight it would appear that the watery boundaries which keep out the land quad- rupeds could be easily passed over by birds, yet prac- tically it is not so; for if we leave out the aquatic tribes which are pre-eminently wanderers, it is found that the others (and especially the passeres, or true perching birds, which form the vast majority) are generally as strictly limited by straits and arms of the sea as are quadrupeds themselves. As an instance, among the islands of which I am now speaking, it is a remarkable fact that Java possesses numerous birds which never pass over to Sumatra, though they are separated by a strait only fifteen miles wide, and with islands in mid-channel. Java, in fact, possesses more birds and insects peculiar to itself than either Sumatra or Borneo, and this would indicate that it was earliest separated from the con- tinent; next in organic individuality is Borneo; while Sumatra is so nearly identical in all its animal forms with the peninsula of Malacca, that we may safely MALAY ARCHIPELAGO. 233 conclude it to have been the most recently dismembered island. “The Philippine Islands agree in many respects with Asia and the other islands, but present some anomalies to indicate that they were separated at an earlier period, and have since been subject to many revolutions in their physical geography. , “Turning our attention now to the remaining portion of the Archipelago, we shall find that all the islands, from Celebes to Lombock eastward, exhibit almost as close a resemblance to Australia and New Guinea as the Western Islands do to Asia. It is well known that the natural productions of Australia differ from those of Asia more than those of any of the four ancient quarters of the world differ from each other. Australia, in fact, stands alone; it possesses no apes or monkeys, no cats or tigers, wolves, bears, or hyenas, no deer or antelopes, sheep or oxen, no elephant, horse, squirrel or rabbit; none, in short, of those familiar types of quadruped which are met with in every other part of the world. Instead of these, it has Marsupials only, kangaroos and opossums, wombats and the duck-billed platypus. In birds it is almost as peculiar. It has no woodpeckers and no pheasants, families which exist in every other part of the world; but instead of them it has the mound-making brush-turkeys, the honeysuckers, the cockatoos, and the brush-tongued lories, which are found nowhere else upon the globe. All these striking peculiarities are found also in those islands which form the Austro-Malayan division of the Archipelago. “The great contrast between the two divisions of the Archipelago is nowhere so abruptly exhibited as on 234 THE DOCTRINE OF DESCENT. passing from the island of Bali to that of Lombock, where the two regions are in closest proximity. In Bali we have barbets, fruit-thrushes, and woodpeckers; on passing over to Lombock these are seen no more, but we have abundance of cockatoos, honeysuckers, and brush-turkeys, which are equally unknown in Bali or in any island further west. The strait is here fifteen miles wide, so that we may pass in two hours from one great division of the earth to another, differing as essentially in their animal life as Europe does from America.* Ii we travel from Java or Borneo to Celebes or the Moluc- cas, the difference is still more striking. In the first, the forests abound in monkeys of many kinds, wild cats, deer, civets and others, and numerous varieties of squir- rels are constantly met with. In the latter, none of these occur, but the prehensile-tailed cuscus is almost the only terrestrial mammal seen, except wild pigs, which are found in all the islands, and deer (which have probably been recently introduced) in the Celebes and the Moluc-. cas. The birds which are most abundant in the Western islands are woodpeckers, barbets, trogons, fruit-thrushes, and leaf-thrushes; they are seen daily, and form the great ornithological features of the country. In the Eastern islands these are absolutely unknown, honeysuckers and small lories being the most common birds; so that the naturalist feels himself in a new world, and can hardly realize that he had passed from the one region to the other in a few days, without ever being out of sight of land. “The inference that we must draw from these facts is undoubtedly that the whole of the islands eastwards, * This is to vaguely expressed. It would be nearer the mark to say, as Europe does from South America. (O. SCHMIDT.) FORMER PACIFIC CONTINENT. 235 beyond Java and Borneo, do essentially form a part of a former Australian or Pacific continent, although some of them may never have been actually joined to it. This continent must have been broken up not only before the Western islands were separated from Asia, but proba- bly before the extreme south-eastern portion of Asia was raised above the waters of the ocean; for a great part of the land of Borneo and Java is known to be geologically of quite recent formation; while the very great difference of species, and in many cases of genera also, between the productions of the Eastern Malay islands and Australia, as well as the great depth of the sea now separating them, all point to a comparatively long period of isolation.” “Jt is interesting to observe among the islands them- selves how a shallow sea always intimates a recent land connection. The Aru islands, Maisol and Waigiou, as well as Jobic, agree with New Guinea in their species of mammalia and birds much more closely than they do with the Moluccas, and we find that they are all united to New Guinea by a shallow sea. In fact, the 1oo-fathom line round New Guinea marks out accurately the range of the true Paradise birds. “Tt is further to be noted—and this is a very interest- ing point in connection with theories of the dependence of special forms of life on external conditions—that this division of the Archipelago into two regions character- ized by a striking diversity in their natural productions, does not in any way correspond to the main physical or climatal divisions,of the surface.” We will further quote only the following: ‘Borneo and New Guinea, as alike physically as two distinct countries can be, are 236 THE DOCTRINE OF DESCENT. zoologically wide as the poles asunder; while Australia, with its dry winds, its open plains, its stony deserts, and its temperate climate, yet produces birds and quadru- peds which are closely related to those inhabiting the hot, damp, luxuriant forests which everywhere clothe the plains and mountains of New Guinea.” Wallace gives the most specific proofs that, as the parts of this Archipelago approach one another like sepa- rated extremities of two continents, they bring with them two entirely different fauna. Similarly, the Mediter- ranean and West Indian Archipelagos are devoid of any peculiar character, and are completely dependent on the adjacent continents for their animal life and vegetation. We have already discussed Madeira and its land snails. Insular faunas therefore do not require the hypothesis of more centres of creation than are offered by the continents; and Rttimeyer has endeavoured to trace the extraction of birds and mammals to two centres of derivation. A great series of animal-geographical facts is explicable only on the hypothesis of the former ex- istence of a southern continent, of which the Australian mainland is a remnant. The present Marsupials are concentrated in Australia. Their occurrence in the south-western portion of the Malay Archipelago, includ- ing New Guinea, seems like a radiation from that centre. No single token makes it appear that the Marsupials existing in former periods in the northern hemisphere, from the Jura forwards, had migrated to meet those which were pressing on from the southern continent towards the equator. Only as to the oppossum, so widely extended in South America, could a question arise, which is however solved by the examination of a 1 SOUTHERN FAUNA. 237 host of congeners, one and all alien to the population predominant in America, and indicating importation — probably in the Tertiary period; unless it be assumed, with Rtitimeyer, ‘that implacental mammals were cre- ated out of Australia as well as in it.” Among the first to be mentioned are the wingless birds, that is, those which are anatomically and syste- matically connected, and which we now find scattered over continents and some of the larger islands. The cassowary of New Holland and America, the extinct giant birds of Madagascar and New Zealand, the African ostrich, which has advanced from the south northwards, cannot have originated in their present isolation. The same considerations are forced upon us by the mammals named Bruta by Linneus, and by modern zoologists termed Edentata, by reason of their imperfect dentition, among which, accepting the latter definition, must be included the Ornithorhyncus, or duck-mole of Tasmania. These duck-moles incon- testibly occupy the lowest grade among the mammals now extant; but the other true Edentata are no less alien to the higher orders, and their occurrence in South Amer- ica on the one hand, and in South Africa and South Asia on the other, as well as ‘the impossibility of tracing them from a common centre in the northern hemisphere, points to the vanished land of the south, where perhaps the home of the progenitors of the Maki of Madagascar may also be looked for. “Or,” says Rtitimeyer, “does the hypothesis of a Polar land, once possessing an abundance of animal life, partly covered by the ocean and partly by a coat of ice, appear an unfounded assumption to us who now witness 238 THE DOCTRINE OF DESCENT the elevation of a similar frozen surface in the northern hemisphere, and are surrounded in the Alps by a still existing—in our glacial drift by a scarcely vanished— arctic scene? Or need the conjecture that the almost exclusively graminivorous and insectivorous Marsupials, sloths, armadilloes, ant-eaters, and ostriches, once pos- sessed an actual point of union in a southern continent, of which the present flora of Terra del Fuego, the Cape, and Australia, must be the remains,—need this conjec- ture raise difficulties at a moment when from their fossil remains Heer restores to our sight the ancient forests of Smith’s Sound and Spitzbergen? ” Having ventured to reconstruct the southern conti- nent, with its strange fauna, of which the remains are so widely dispersed, Riitimeyer casts about for more specific evidence in favour of the hypothesis to which the course of the world’s formation everywhere gives rise, that fresh-water animals and likewise terrestrial ani- mals came up from the sea. Hence the notably small division of sirenoid fish (Lepidosiren, Protopterus), which breathe air during the dry season of the year, must not be considered reptiles adapting themselves to aquatic life, but the reverse. The organ which in fish served as a hydrostatic apparatus, the swim bladders, becomes in them the lung. Thus we must go back from terres- trial to aquatic tortoises, and from them to those deni- zens of the sea which are allied to the Enaliosaurians, so frequent in the Jurassic strata. The evolutionary and biographical history of the land crabs shows us in the plainest manner how the inhabitant of the sea be- comes a terrestrial animal; a special problem which, as we have already mentioned, Fritz Miller has completely MODIFICATION OF FISHES. 239 solved and capitalized, in his essay, “for Darwin.” Of the sirens, commonly but erroneously reckoned among the Cetacea, and of which the majority prefer remain- ing at the mouths of large estuaries, one entire species has penetrated into the great inland lakes of Africa; and certain species of salmon as well as the sturgeons, which alternate periodically between salt and fresh water, are in the phase of gradually forsaking ocean life. From my special experience, I may add that the brackish water sponges are certainly dependent on the marine families, and that the fresh-water species unmistakably point to these brackish forms. If in all these cases we are dealing with gradual trans- formation, and more or less voluntary adaptation, there is no lack of conspicuous instances of forcible and almost sudden severance; of upheavals by which former sections of the ocean became inland seas. What were the modi- fications undergone by the fish and crabs secluded with them, is shown by the-fine observations of Lovén on the animals of Lakes Wener and Wetter, and of Malmgren on those of Ladoga. The latter brings evidence that the salmon-trout of the Alps (Salmo salvelinus) is derived from the Polar Sea, and is own brother to the Scan- dinavian Salmo alpinus. Riitimeyer pronounces the opinion that by more minutely tracing the relations of the fresh-water fauna to those of the denizens of the ocean, the cosmopoli- tanism of fresh-water animals will be explained, as well as the relation of antarctic to arctic life. For the present, however, these two great animal-groups, as regards the higher, warm-blooded classes, are somewhat sharply con- trasted. It is only from scanty remains that we know 240 THE DOCTRINE OF DESCENT. that so early as the Jurassic era, the northern hemisphere was peopled by Marsupials, but, it is evident, not densely. We must suppose that, retaining their character, the Mar- supials of the southern continent tested and proved their powers of adaptation, whereas on the other side of the equator a race of mammals of completely different cast proceeded from them. This is the race which still char- acterizes the whole surface of the earth from the north to the point of contact with the more stable remnants of antarctic life. While with reference to their origin we can appeal only to reason and inference, the historical connection between the mammalia now peopling the Old and the greater part of the New World, and their pred- ecessors up to the most ancient Tertiary periods, is manifest to our eyes. The remains of the earliest mammals here to be con- sidered, are found in the Eocene deposits of Switzerland, and in corresponding strata in France and the south of England. From the southern edge of the Jurassic plateau, neither the Alps nor any other land was visible, and the ocean which washed its shores has been traced as far as China. The mammalia of this period, as far as they are known, amount, according to the synopsis made by Ritimeyer in 1867, to at least 70 species. The ma- jority are ungulate, therefore Graminivora; of these, by far the greater number Pachydermata. Now, when the entire world scarcely maintains so many Pachyderms, the proportions are quite altered. In Europe, the pig alone represents this division, and Ruminants everywhere predominate. In its present animal population, Africa might be approximately compared to Eocene Europe. But as to these Ungulates must be added a large num- TERTIARY FAUNA. 241 ber of Carnivora, resembling the Viverrida (polecats, martens, &c.) and hyenas, and as viverride exist in Af rica as well as in Asia, and as, moreover, the musk ruminants represented in this primitive fauna are now likewise Asiatic and African, and, finally, as the French opossums of those ages still live in Central and South America, “we gain an impression that the most ancient Tertiary fauna of Europe is the source of a truly continental animal society now represented in the tropical zone of both worlds, but most emphatically in Africa.” Far more heterogeneous is the picture of the higher animal life of the middle and more recent Tertiary periods which we reconstruct from the numerous and in parts highly prolific repositories of these remains. To draw narrower limits within these periods is impractica- ble; from place to place, from stratum to stratum, there is coherence; nowhere does a species appear that might not be derived from another; and our authority says that anatomy, morphology, paleontology, and geographical distribution, seemed to impress no doctrine upon him with such energy and pertinacity as that separate species of a genus, species without any historical and therefore without any previous local link to any original stock, do not exist. The most celebrated repository of Ter- tiary mammals is Pikermi, a short distance from Athens, an accumulation of skeletons complete and in fragments, which pre-supposes a profusion of animals, of which at any rate the most densely inhabited regions of Africa may, according to Livingstone’s descriptions, give us an idea. Again the Carnivora give way to the Graminivora, 242 THE DOCTRINE OF DESCENT. though the feline beasts of prey make themselves con- spicuous; and among the great Tertiary beasts of prey are some which have a range as great as the tiger of the present age. The territory of the extinct sabre-toothed tiger (Machairodus) at that time extended over a great part of America and Europe. Let us also mention that the canine animals appear somewhat later, and that the bears are of still more recent origin. At this period the most abundant material still favours the ungulates. Cloven feet still preponderate. Pigs and musk-animals are the most constant. But the tapir, in shape like the older forms, is now joined by the rhinoceros, the true horses, and the elephants. If the origin of the rhinoceros is somewhat obscure, the extraction of the mastodon, the older form of the elephant, is hitherto quite unknown.” And yet though we search in vain through the known mammalian fauna of the Eocene period for the most nearly allied parent forms, there are numerous tokens that even in Europe and Asia, “the most of the Eocene must be regarded as the true root forms of the Miocene genera.” (R.) This is shown by the discoveries at Nebraska in North America, where important genera, which, like the Palzotherium, disappeared from the Old World in the Eocene period, took refuge in company with newer genera. We likewise find there, intermediate forms between the lama and the camel, which in this case alone gives its true significance to the once unmean- ing word, vicarious genera. At Nebraska we moreover find the triple-hoofed horse (Anchitherium), and we hence know the origin of the single-hoofed horse of the Old and New Worlds. What has happened in the Old World since that age DISTRIBUTION OF ANIMALS IN AMERICA. 243 is confined to the extinction of many Pachydermata, a displacement of the rhinoceros, elephant, tapir, and hip- popotamus, and an extremely abundant development of the true ruminants and the cattle which proceeded from them with an exaggerated form of head. Bears and canine species occupy the territory where viverride and hyenas once predominated; but as ‘“ numerous locally and historically limited species, a large number—among the smaller fauna a majority—of Miocene races remain in possession of the ancient and probably constantly increas- ing habitat.” (R.) “In this gradual change of things, no one will be able to discern aught but phenomena of the same order of which we are still the witnesses.” (R.) How circumstances occurred in America has been described in a masterly style by Riitimeyer as follows: “ America affords a basis for the distribution of animals completely different from that of the Old World. In the latter, ridges, open only in places, divide the entire continent into mountainous zones, and correspond to the distribution of temperature. Thus in a twofold manner they prescribe a definite range east and west to the extension of animals; while a migration from north to south is impeded less by the height of the mountains than that on their summit the north comes into contact with the scorching south. Behind this wall, moreover, in the expanse from the Caspian Sea to China, there is a zone of steppes and deserts which fences in the animals more effectually than the mountain chains. In America, not beasts of prey alone, but graminivora also, may advance without hindrance from the regions of the lichen on the Mackenzie River, through the pine forests of Lake Superior, to the land of the magnolia 244 THE DOCTRINE OF DESCENT. in Mexico; 40°—50° of latitude separate the extremes which meet in the Himalayas, and the vast plains and huge river systems seem almost to solicit immigra- tion. The accordance of the whole faunas of Mex- ico and Guiana, moreover, shows how little the isthmus of Panama checks the advance to South America, where again one mighty fluvial system trenches upon the other without any lofty partitions; nor is there any arid desert in the whole extent from the Canadian seas to Patagonia. “We shall probably not be wrong in ascribing the remarkable extension of fossil and present mammals of America in a great measure to this circumstance. As we have seen, the Miocene fauna of Nebraska is the offspring of the Eocene fauna of the Old World. The Pliocene animals of Niobrara, which are buried in the same district as Nebraska, but on more recent arenaceous strata, still further corroborate this statement: elephants, tapirs, and many species of horses, scarcely differ from those of the Old World; the pigs, judging by their dentition, are descendants of European miocene Paleochceride. The ruminants are represented by the same genera, and partially by the same species, as in the analogous strata of Europe, as deer, sheep, and buf- faloes; neither do the carnivora or the minute animal life offer an exception. Many genera of an entirely Old-World cast have in the lapse of time penetrated far into South America, and there died out shortly before the arrival of man, or perhaps by his co-operation, as was the case with the two species of mammoth of the Cordilleras and the South American horse, whose present successors reached this insular continent by a far shorter DISTRIBUTION OF ANIMALS IN AMERICA. 2,45 road. Even a species of antelope and two other horned ruminants (Leptotherium) found their way to Brazil. Two sorts of tapir, of which the dentition, even in Cuvier’s eyes, is scarcely distinguishable from the Indian species; two species of pigs, still bearing in their milk-teeth un- mistakable characters of their aboriginal form; and a number of deer, besides the lamas, a later and originally American offshoot of the Eocene Anoplotheria—are one and all living remnants of this ancient colony from the East, which did not reach its dwelling-place without copious losses on its long pilgrimage. It can scarcely be doubted that many of the beasts of prey which in the Diluvium of South America retained their family charac- ter more than they do now, must have arrived there in the same manner. Let us now remember that even the Eocene Czenopithecus of Egerkingen distinctly pointed to the present apes of America, and that the Didelphidze (Opossums) lie buried in the same European soils. It might almost appear that it was pre-eminently the di- vision of arboreal quadrumana which, with the opos- sums, domesticated itself in the vast forests, of their new abode, and, receiving a fresh impulse, gave rise to a multitude of special forms, without however having, even in the present times, reached the pitch of development attained by their cousins who had remained behind in the Old World. “We may now appropriately return to our previous remark that this migration of animals did not find the south of the New World destitute of mammals, but rather already occupied by the toothless representatives of ant- arctic, or at least of southern animal life. The diluvial fauna of South America collected by Lund, Castlenau, \ 246 THE DOCTRINE OF DESCENT. and Weddell, from the Brazilian caves, and the alluvium of the Pampas, among the 118 species cited, actualiy includes, in addition to those already mentioned, as be- ing of probably Old-World pedigree, no less than 35 species of Edentata, and these animals of consider- able bulk. Not reckoning the 36 rodents and bats, and the smaller fauna in general, they constitute nearly half of the larger diluvial animals of South America. The assemblage of Edentata previously settled in these regions thus held their own against the invasion from the north. “Tt is comprehensible that the same external causes which led the march of the children of the north con- stantly further, may likewise have invited the members of the antarctic fauna to extend themselves northwards. As we even now encounter the incongruous forms of the sloth, the armadillo, and the ant-eater in Guatemala and Mexico, in the midst of a fauna in great part con- sisting of races still represented in Europe, we also find, even in diluvial eras, gigantic sloths and armadillos rang- ing far into the north. Megalonyx Jeffersoni, and Mylo- don Harlemi, sentries of South American origin thrown out as far as Kentucky and Missouri, are a phenomenon as heterogeneous in the land of the bison and the deer, as is the mastodon in the Andes of New Granada and Bo- livia. Over the whole enormous extent of both portions of the New Continent, the mixture and interpenetration of two mammalian groups of completely diverse families, constitutes the most conspicuous feature of its fauna; and it is significant that each group increases in the abundance of its representatives and in the originality of their ap- pearance as we approach its point of derivation.” CAUSES OF GEOGRAPHICAL DISTRIBUTION. 2,47 Hence, on both sides of the ocean, north of the very sinuous boundary of the antarctic or southern fauna, we find ourselves still in the midst of the diluvial animal world, which extended itself, by a bridge in the vicinity of the North Pole, from the old continents to the mainland of America, and there for a longer period re- tained its ancient appearance in the mastodons and horses. There, as well as here, the present order of things— the cantonment of animals—has been in many ways de- termined and modified by mighty glacifications and pro- longed periods of refrigeration. Hence the accordance of so many plants of the extreme north with Alpine plants after the Eocene vegetation had made its entry from the east. Since that age, the reindeer has been forced back to the north, and the musk ox has been expelled and exterminated from the Old World. The elephants, fleeing before the ice, have not returned; and the mammoth, immigrating with a rhinoceros from the north-east, has been destroyed with his associate. Others of his comrades, such as the primeval ox, died out only a few centuries ago as wild cattle; others, like the buf- falo and the beaver, are nearly extinct as denizens of Europe; and others again, the deer and roe-deer, will perish with the forests and the game-laws. But of al- most all the species of which we search for the extrac- tion, Paleontology supplies us with the history and de- rivation; and in derivation we find the causes of geo- graphical distribution sketched in vivid outlines. 248 THE DOCTRINE OF DESCENT. XI. The Pedigree of Vertebrate Animals. THE final result towards which the doctrine of Descent directs its efforts, is the pedigree of organisms. To work it out is to collect the almost inconceivable pro- fusion of facts accumulated in the course of about a century by descriptive botany and zoology, including comparative anatomy and the history of development and to submit the existing special hypotheses to a minute scrutiny and renewed verification. We have therefore claimed in behalf of the doctrine of Derivation the priv- ilege on which the progress of science generally relies— that of investigating according to determined points of view, and accepting probabilities as truth in the garb of scientific conjecture or hypothesis. It is manifest that when the doctrine of Descent first made its appearance with the arguments proposed by Darwin, it was only possible to indicate the most general outlines of this great pedigree, which it was the special task of the new di- rection of science to demonstrate in all its details. But however and wherever specific research was attempted, either the results contributed the form of some part of the great pedigree, or there was, from the first, reason to pre-suppose certain kinships, and the conjecture was PEDIGREE OF VERTEBRATE ANIMALS. 249 tested. The further an inquirer has carried his sur- vey of the conditions of organization in any of the larger groups, the less will he be able to divest himself of the genealogical idea in his every act and thought. All this is so self-evident, that one would scarcely sup- pose that the use of this method could have been made a subject of reproach to the doctrine of Descent. Never- theless, it frequently occurs, and the champions of the doctrine of Descent are blamed for often speaking of mere probabilities, forgetting that even in cases in which the probability ultimately proves false, the refuted hy- pothesis has led to progress. Of this the science of lan- guage has recently borne testimony. It is well known that linguistic comparison within the family of Indo- Germanic tongues suggested the reconstruction of the primitive language which formed: their common basis. Johannes Schmidt ™* now proves that the fundamental forms disclosed may have originated at widely different periods, and hence that the primitive language, regarded as a whole, is a scientific fiction. Nevertheless, inquiry was essentially facilitated by this fiction, and with it was intimately connected the formation of a pedigree of the Indo-Germanic linguistic family, as a hypothesis supported by many indications. A bifurcation was as- sumed into a South European language, with Greek, Italian, and Celtic ramifications, and another language, from a second division of which proceeded the funda- mental language of North Europe and the Aryan fun- damental language. Although Johannes Schmidt has demonstrated that this pedigree is false, as the existence of Slavo-Lettish shows the impossibility of the first di- 250 THE DOCTRINE OF DESCENT. vision assumed, the value of the hypothesis is undimin- ished. It was the road to truth. In our science Haeckel has made the most_extensive use of the right of devising hypothetical pedigrees as landmarks for research. It matters nothing that he has repeatedly been obliged to correct himself, or that others have frequently corrected him; the influence of these pedigrees on the progress of the zoology of Descent is manifest to all who survey the field of science, not to mention that in the last ten years a series of researches have conclusively fixed their results in good pedigrees. As we propose to give merely an introduction to the doctrine of Descent, we shall content ourselves with showing how the system or the pedigree is constituted in its application to the single group of the Vertebrata. Mammals. Birds. Reptiles, k SS Amniota. ? Enaliosaurians. Amphibians, ae | Amphioxus. oa a Testacea, epee monet Primordial Vertebrata. Annulosa. As we have seen above, the most important indica- tions of the pedigree of the species are contained in the ' PEDIGREE OF VERTEBRATE ANIMALS. 251 evolutionary history of the individual. But though all vertebrate animals testified their family connection by agreeing iter se in the distribution of the germ as well as in the fundamental important organs, the spinal cord and the veretebral column, every token of their descent from inferior animals, which is unconditionally demanded by the theory, seemed to be entirely wanting. In other words, it seemed that in all vertebrate animals the mem- ory of their original derivation had been obliterated by curtailed development (comp. p. 211). Thus the case remained until Kowalewsky a few years ago studied the development of the lancelet (Amphioxus), the lowest vertebrate animal known, and showed that in this crea- ture the typical phenomena of vertebrate development FIG. 22,—Larva of the Lancelet after Kowalewsky. are preceded by the phases required by the theory. We have already made acquaintance with this form of de- velopment (p. 51, &c.), and we here again. point out its profound significance. It is only when the Amphioxus has passed through the phase of the vibrating, sac-like 252 THE DOCTRINE OF DESCENT. gastrula larva that the future dorsal side becomes flat- tened, and the protuberances arise, which shortly after close into the sheath of the spinal marrow, while beneath it originates this important cellular column, the chorda dorsalis, or notochord. With this the lancelet becomes a vertebrate animal, and the preceding phases do not (according to the view at one time inculcated by C. E. v. Baer respecting such phenomena) recail the inferior and undeveloped in general by the absence of differ- entiation, but they agree in genesis and distribution, in the differentiation of their cellular layers, and in their totality, with the Gastrula phases of invertebrate ani- mals. We are therefore fully justified in regarding these first incidents in the evolution of the Amphioxus as a reminiscerice of the roots of the pedigree of the Verte- brata; and this direct indication of the descent of verte- brate from invertebrate animals is supported by a second and no less important discovery by the Russian natural- ist. It is, that during their development a number of the Tunicata of the division of the Ascidians temporarily possess a spinal cord, and the rudiments of a vertebral column. Kowalewsky’s researches have been ratified on all essential points and in many ways extended by Kupfer, and the facts which interest us may be explained by the diagram, Fig: 23, representing the forepart of the larva of an Ascidian in a somewhat advanced stage. The bulk of the Ascidian larva consists of a body of which our figure shows the whole, and a rudder-like tail. The appendages projecting from the body on the right are organs of adhesion, by means of which the larva fixes itself for its definitive transformation. At o the orifice PEDIGREE OF,VERTEBRATE ANIMALS. 253 of the mouth is formed; d developes into the branchial cavities and the intestinal canal, and we will incidentally remark that in the lancelet also, the anterior end of the 254 THE DOCTRINE OF. DESCENT. primitive intestine becomes the branchial cavity. But with reference to the vertebrate animals, the most im- portant parts of the Ascidian larva are the following. “It possesses a true spinal cord with a vesicularly ex- panded brain (ra). The distribution and position of this organ agrees accurately with the corresponding parts of the vertebrate animal, and Kupfer has even discerned the rudiments of nerves (sss), which, if the observation is confirmed, will still more incontroverti- bly establish the homology of the organ in question with the spinal cord of the Vertebrata and the nerves pro- ceeding from it in pairs. But we know that it is not the spinal cord alone, but its combination with the ver- tebral column which constitutes the characteristic fea- ture of the vertebrate animal. This vertebral column the Ascidian larva likewise possesses (c) in the form of the noto-chord, and, as in the vertebrate animal, this embryonic vertebral column lies between the intestine and the spinal cord. So far goes the accordance; henceforth, the development of this part, so important to the verte- brate animal, becomes retrogressive in the Ascidian. The rudder-like tail, with the spinal cord contained in it, and the noto-chrord, are cast off when the animal be- comes fixed; the larval brain which promised so well, shrink into an insignificant nervous ganglion, and the complete animal gives no cause for suspecting its analogy with the Vertebrata. These laborious observations prove that the Verte- brata are not the sole proprietors of the spinal cord and vertebral column, but received these organs as a heritage from lower grades of organization as their progenitors. It does not occur to the Darwinists to regard man as the LOWEST VERTEBRATE ANIMALS. 255 direct offspring of the present apes; neither do they infer from these observations on the Ascidian larva that vertebrate animals are descended from the Ascidians. Their accordance much rather forces us to assume an unknown primordial vertebrate family, springing from some branch of the heterogeneous division of the Annulosa. From these diverged on one side the Tes- tacea, who might perhaps be called mischanced vertebrata, and on the other the true vertebrate animals.” The Amphioxus which lives in the sand in shallow places on various coasts, and is daily caught by thou- sands at Messina for example, is five or six centimetres in length, and is compressed after the manner of a fish, pointed at both ends, and semi- transparent whilst alive. It pos- sesses no trace of limbs, at the posterior end only a pair of minute membranous margins, the indication of dorsal and caudal fins, and is so simple in its internal structure that it is usually, though inaccurately, termed a fish. Its skeleton is limited to the noto-chord, and some minute cartilaginous rods at the mouth and gills. It has no brain, and, except a small ciliated sac, perhaps to be in- terpreted as an olfactory organ, no sensory apparatus; the heart is tubular. And thus between the lancelet and other true fishes there exists so wide a difference that the possibility remains open that the fishes passed through some other course of development than phases like that of the Amphioxus. i FIG. 24.—Full-grown Ascidian, 256 THE DOCTRINE OF DESCENT. Our knowledge of the genealogy of the fishes may be laid down in the following diagram :— Dipnoi. Teleostei. Ganoids. Elasmobranchii. Marsipobranchii. The Marsipobranchii (Cyclostomi), it is true, exhibit important peculiarities, such as deficiency of limbs, en- tire absence of bony plates or scales on the integument; but the brain, heart, and vertebral column (which, al- though persistently cartilaginous, is far superior to that of the Amphioxus), show their direct connection with the fishes. Fossil remains of these animals, universally known in the genus lamprey (Petromyzon), are not forthcom- ing, and, at the most, only their horny teeth could have been preserved. After these manifest gaps in our knowledge, the suc- ceeding orders of fishes present themselves in a connec- tion all the more conspicuous. The starting-point is formed by the Elasmobranchii, to which belong the true chimeras, sharks, and rays. Brain and gills testify their kindred with the Marsipobranchii. In the construction of the cranium, facial bones, pectoral and pelvic arches, and the anterior extremities, heart and intestine, they exhibit forms to which, as Gegenbaur has shown in his PEDIGREE OF FISHES. 257 well-known observations, the homologous parts of the Ganoids are related either as progressive developments or as reductions. Huxley has also prepared the way for a correct apprehension of these relations. To be fully convinced of this, detailed study is certainly requisite; for in its absence it is impossible to imagine, how in the Elasmobranchii the true branchial apparatus is wanting, and how the cartilaginous arch, which, in them, replaces the gills, is applied in the Ganoids, partly as the palate, and partly as the attachment for the true lower jaw, while the internal gills of the former, become the external gills of the latter; how in the skeleton of the anterior ex- tremities, a gradual simplification may be exhibited, step by step, from the sharks and rays to the Ganoids, and especially the sturgeon,—a process of which the two ex- tremes are’ reached in the Teleostei on the one side and the higher Vertebrata on the other—in the latter in the multiform perfection of the arm and hand. Of the Ganoids only scattered remnants survive, the sturgeon family and some few American and African genera, of which, as Rutimeyer says, a flight into fresh water has been the salvation. They just suffice to ex- plain the relation of this once extraordinarily extensive group, to the Elasmobranchii as well as the Teleostei. In the Teleostei, the metamorphosis of the organ- ization of the Elasmobranchii initiated in the Ganoids, is carried yet further. It is only with great qualifica- tion that they can be termed ‘“ more highly developed,” in the skeleton perhaps, to which older zoologists attrib- uted too much importance. Brain, heart, the form of the extremities, and the reproductive system, are indeed distinct developments which, in combination with the 258 THE DOCTRINE OF DESCENT. external shape and integuments, have exhibited great powers of adaptation, but have not proved capable of any further development. Comparative anatomy has vainly spent much labour in attempting to trace the con- dition of the higher animals from the special organiza- tion of the Teleostei, or to explain the peculiarities of the Teleostei from above downwards. It was labour lost, for the solution is to be reached only by the method in- dicated in the derivation of the Teleostei, through the Ganoids, from the shark-like fishes. Hence, at the present period, a development is con- cluded with the Teleostei, and we must look to another grade for the transition from the fishes to the amphibians. We find one in the order of the mud-fishes (Dipnoi), scantily represented by only few species (Lepidosiren Protopterus). These fish-like animals, living in American and African rivers which dry up in the hot season of the year, are fish by right of their skeleton and scales, and some other characteristics; the skull, however, al- most resembles that of an amphibian, and they also pro- visionally use their swim-bladdets as lungs; and by thus breathing alternately water and air, they set before us the transition of the gill-breathing larve of the amphibians to the phase of air-breathing. Of the true fishes at the present time, they most nearly approach the family of the Crossopterygii, represented by the Af rican Polypterus; and the discovery of a very re markable Australian fish, the Ceratodus, confirms this affinity. Through forms thus resembling the Dipnoi, the ad- vance from the fishes to the amphibians was probably accomplished. But, as a scientific friend, profoundly TRANSITION TO THE AMPHIBIANS. 259 versed in the history of development, has pointed out to me,—supporting his remark on the comparison of the respiratory organs of the Marsipobranchii with those of the amphibians,—it is possible that frogs and sala- manders may be directly descended from beings closely analogous to the division of the Marsipobranchii termed Myxine. It is to be hoped that this highly interesting observation thay soon be made public. We gather from the general Ontogenesis of the amphibians, that the tailed forms are the most ancient. This is also the case with the oldest amphibian-like animals, the Labyrinthodonts. From their remains (Archegosaurus and others), chiefly contained in the Carboniferous formation, we have learnt that they had incomplete limbs or none, that their ven- tral side was partially provided with bony plates, the vertebral column fish-like, and that their skull, with some of the characters of the present amphibians, combined others which remind us partly of certain bony Ganoids, and partly of the reptiles which subsequently appeared. Now if in the singularly elongated snake-like Ccecilia, which is however without tail or limbs, some peculiarities ‘of the skull of the Labyrinthodont appear again, we must own our utter ignorance as to the actual progenitors of this, as well as of the two other living orders of the Ceecilia and the Batrachians. Here, therefore, we are, as we have said, thrown entirely on the evolutionary history of the individual. By what right we may frame a picture with great probability approaching the truth, the reader may have gathered from our previous chapters. Among the tailed amphibians, it is not only in Ontogenesis that we see the passage from gill to lung- 260 THE DOCTRINE OF DESCENT. breathing; the systematic series from the proteus to the triton and the salamander, likewise exhibits this physiological ascent, linked with various morphological transformations, which may similarly be shown between the ancient and modern specimens of the Labyrintho- donts. The Batrachians, indeed, rise higher in develop- ment than the Coecilia; but, as the friend above men- tioned informs me, they more nearly approach the Myxine in the construction of the internal gills of their larva. We shall obtain a general view of the reptiles by means of the appended diagram, in which we shall avoid any minute systematic designations. “8 : , g = ~ w #1 8/8 gla g| £18 ° 8 oO a > : Ss 2 g BY 8 e sis| #|% 8 |. 2/82! > Ele] BB] Ss] 8 je 2) 2) 2. eel 3 tele ie S| & A |} RFiln!| a |ma] oOo }o Present Age............0 Diluvium ......... lL Fit t iT Tertiary Period tT) t| t t halk... secsersc t t tft r Oolite.. +] + t t+ LF] PF yt Trias +t +/t t T t DYAS sscvarsinnreecedegseasecs T COAL teteeestcecens tesuconrs \ \ Mba He ae \ Se ms Le ( = ‘ \ i ' Pia ‘ Ss ' ‘ 4 \ \ * Sy : fo) vee XK ‘S *S ~ i / Pig . x 5 ! : 1 1 1 Pa ~ . Aww ewe tenn = ’ s ’ PEDIGREE OF REPTILES. 261 The class presents a very comprehensive picture, al- though only four orders now exist, of which two, the lizards and the snakes, are scarcely to be separated. That the snakes, which first appear in the Tertiary period, are a direct offshoot from the lizards, is reduced to a cer- tainty by comparative anatomy and the history of devel- opment. In the various families of lizards we see the absence of feet occurring in conjunction with the elonga- tion of the body and the multiplication of the vertebre; and the modifications peculiar to the skull of the “true” snakes are likewise represented in the systematic series in every gradation, beginning with the skull of the true lizard. We cannot specify the fossil genera in which the transformation was initiated; but in this case a doubt would be only a capricious denial. It is otherwise with the remaining orders, which in the beginnings, hitherto accessible to us, exhibit diversities so decidedly marked, that in none has it been possible to trace a direct descent from any known member of another. Prof. Huxley, a great authority on the anatomy of these animals, says on this subject as follows:— , “Tf we ask, in what manner the earliest representatives of these orders are distinguished from their living or latest known representatives, we shall find, in all cases, that the amount of difference in itself is remarkably small in comparison with the length of time during which the order has existed. So far as I know, there is no fact to show that the later Plesiosauria, or Ichthyosauria, exhibit an advance upon the earlier members of the group. It is not clear that the Dinosauria of the wealden and of the Cretaceous formations are more highly or- ganized than those of the Trias; and even where a dif- 262 THE DOCTRINE OF DESCENT. ferentiation of structure is to be observed, as in the Lacertilia, or Crocodilia, it goes no further than a modi- fication of the form of the articular surfaces of the verte- bra, or of the degree to which the internal nasal aper- tures are surrounded by bone. The osteological differ- ences, which alone are exhibited by fossil remains, have doubtless been accompanied by many changes in the organization of the destructible parts of the body; but everything tends to show that the amount of change in the organization of reptiles since their first known ap- pearance upon the earth, is not great in itself; and is wholly insignificant, if we take into consideration the lapse of time, and the changes of the surface of the globe, which are represented by the Mesozoic and Ter- tiary formations. “From the point of view of the evolution hypothesis, it is necessary to suppose that the Reptilia have all sprung from a common stock, and I see no justification for the supposition that the rapidity of their divergence from this stock was greater before the epoch of the Trias than it has been since. Consequently, seeing that the approx- imation of the oldest known representatives of the differ- ent orders is so slight, reptiles must have lived before the Trias for a length of time, compared with which that which has elapsed from the Triassic epoch until now is small—in other words, the commencement of the ex- istence of reptiles must be sought in a remote palzozoic epoch.” Comparison thus points us back to ages which afford no record of the actual derivation of this class. Even the Ichthyosauria and Plesiosauria, so frequently men- tioned in conjunction, deviate widely from one another PEDIGREE OF VERTEBRATE ANIMALS. 263 in very essential characters, which refer their supposed common origin to a remote period. We will mention only the fin-like extremities of the former, which are of an obviously piscine type. We are thus thrown back vaguely on such mixed forms as may have been analogous to the Labyrinthodont; nay, the question arises whether the Ichthyosauria alone, or perhaps the Plesiosauria with them, did not diverge from the fishes independently of the other branches of the reptile family; an eventuality which is taken into account in the pedi- gree at p. 250. A certain resemblance with the skull of the tortoises (Chelonia) is exhibited by that of the Dicynodonta. In them also the jaws, as appears from their shape, were manifestly cased in horny sheaths; but at the same time the upper jaw contained two huge tusks, and it is scarcely possible to imagine a direct transition from the Dicynodonta, appearing in the Trias, to the more recent tortoise. In some particulars of the skall, as well as in the situation of the posterior nasal apertures, the forms of older crocodiles exhibit an affinity with the lizards, from the older and unknown forms of which they probably branched off. The winged saurians, or Pterodactyles, may also be a branch of the lizards. They have gained by adaptation several characters, such as the shape and lightness of head, the length, slenderness, and pneumatic character of the tubular bones, which they share with the birds. But it is not in them, but in the division comprising several families which Hux- ley terms Ornithoscelide, or reptiles with the legs of a bird, that we must look for the actual progenitors of the birds. For among them one of the most im- portant characters of the birds is, in some genera, in 18 264 THE DOCTRINE OF DESCENT. course of preparation, so that in the full-grown animal its origin may still be recognized; in others, as the genus Campsognathus, it is accomplished. We allude to the . peculiarity already discussed in p. 10, that the upper portion of the tarsus is anchylosed with the tibia, the lower with the metatarsus, and that the ankle-joint is hence inserted into the tarsus. All existing reptiles are sharply distinguished from the Amphibians and Fishes by several phenomena ac- companying their development. They possess two or- gans enveloping the embryo; the amnion, which is es- sentially a protecting sheath, and the allantois, by which the foetal circulation, nutrition, and respiration is regu- lated and carried on. In the Batrachians we find indi- cations at least of the allantois, and must suppose that the greater part of the fossil reptiles had already adopted this advance in general organization. It implies an ad- vance, inasmuch as animals developed by the aid of the amnion and allantois make further progress during the embryonic phase than is the case with the inferior Ver- tebrata, and that they hence leave the egg with greater powers of resistance. We must ascribe the adoption of the amnion and allantois to remote periods of amphibian and reptile development, for the additional reason that the possession of their embryonic sheaths and organs is shared by the birds which are descended from true reptiles, and by the mammals which cannot be descended from true reptiles. The birds are, anatomically, so closely allied to the reptiles, that Huxley, who has carried out the com- parison most rigorously, has joined the two classes into a greater systematic unit, under the name of Sauropsida, PEDIGREE OF BIRDS. 265 or lizard-like animals. The scale of a lizard and a feather seem to be totally different things; but in their first rudiments they are completely identical, and the feather has a far greater analogy with the scale, than with the hair. The plumage, which seems to impress a spe- cific character upon the bird, is therefore to be traced from the formation of scales. Of the internal soft organs, we will only remark upon the heart and lungs. All the older geologists placed the heart of the bird on the same level with that of the mammal and of man; in its specific arrangements, however, it is only to be inerpreted by the heart of the reptile, and the wind-pipe is not ramified as in the mammal. That the reptiles exhibit a gradual transition to the leg of the bird, has been repeatedly pointed out. The pelvis of the bird, which is remarkable for the length of the pubis and ischium, and is open in front, likewise represents only a slight advance in devel- opment upon the pelvic structure already shown in sev- eral of the Ornithoscelide. Thus Huxley says with ref- erence to the ischium of the Hypsilophode, that ‘ the remarkable slenderness and prolongation of the ischium give it a wonderfully ornithic character.” Finally, in the skull, peculiarities possessed by the bird in contrast with the mammal, such as the simple condyle of the occiput, the quadrate bone, the cochlea of the auditory labyrinth, the composition of the lower jaw, its articulation with the skull by the intervention of the quadrate bone, &c., are not specific characters of the bird alone, but of reptiles in general. This similarity of type in reptiles and in birds is perfectly manifest from the comparison of living birds with living reptiles. But the proof that the bird is de- rived from the reptile is rendered unimpeachable by 266 THE DOCTRINE OF DESCENT. the discoveries, scanty as they are, of fossil intermediate forms. The pelvis and leg of the Ornithoscelide have Oy already been discussed. But in the slates of Solnhofen we have more- \ over become acquainted with the A \\ Archzopteryx, a bird unfortu- A nately mutilated and in many 25, impression of the tail of the Archzopteryx Macrurus, Ow), but exhibiting a very valuable and . interesting intermediate stage be- n , tween the tail of a reptile and a I { | j i AN ways damaged by pressure (Fig. Ay i . | bird. Among existing birds, the Nandu,orAmerican ostrich(Rhea), aN \\\ | alone possesses numerous separate : caudal vertebree; but the tail of NNN this bird projects so little, that it ; \ in no way recalls the tail of a lizard. lf Now the Archeopteryx exhibits a long tail, bordered by two rows of stiff feathers, of which the impression remains in extraordinary preservation. The skull of this valuable specimen, now in the British Museuin, is so much injured, that no idea can be framed of its construction. It is impossible to decide whether the jaws bore teeth. The example of the tortoises shows that within the reptile type the formation of teeth was replaced by horny sheaths, without a correlated de- velopment of the power of flight; the Pterodactyles, on the other hand, combine with the power of flight a light head, provided nevertheless with numerous teeth. FIG, 25. PEDIGREE OF BIRDS. 267 The obscurity which surrounded these parts of the old antediluvian birds has been cleared up by a discovery by the American naturalist, Marsh. He found in the upper Chalk of Kansas the remains of two genera of birds, which by their bi-concave vertebra remind us of the characteristics of the ancient reptiles, and by this alone present extremely valuable intermediate stages, but which, moreover, bore teeth in both jaws. These teeth are small and sharp, and were so numerous that in the lower jaw of the animal named Ichthyornis dispar, twenty might be counted on each side. Thus we are now quite clear as to the kinship of the bird. It is a reptile adapted to aerial life, and those birds which we see more estranged from flight have ac- quired the characters correlated with more or less in- capacity for flight only by means of retrogression. It fares the worse with the internal arrangement of this class of animals. Partly from their geographical distri- bution, partly from anatomical indications, especially of the skull, it may be inferred that the ostrich-like birds are not, in virtue of their strength of leg and adeptness in running, the youngest members of their class and the most nearly allied to the mammals, but that they are the oldest of those now living. The nature of the imperfection of their wings shows, as we have said, that : they are in a state of arrest or retrogression. Beyond this general experience it is impossible to go. If we contemplate the bird as a flying animal, those of course rank highest which have learnt to fly the best. This palm avowedly accrues to the birds of prey as a whole, although other orders are not deficient in pre-eminent flyers. Brehm and others hold the parrots, because 268 THE DOCTRINE OF DESCENT. of their docility, to be the highest birds. But all this is arbitrary, and can only accidentally correspond with the true and unknown ramification of the ornithic branch in the pedigree of the Vertebrata. The most ancient known remains of the Mammalia are found in the Trias. They occur somewhat more frequently in the central Mesozoic strata, and they all belong to Marsupial animals. Now as Marsupials, in comparison with the inferior classes of vertebrate ani- mals from which they must be derived, are very highly developed, and as in the Monotremata (Duck-mole, Ornithorhyncus, and Porcupine ant-eater, Echidna,) we possess mammals which are manifestly far beneath the Marsupials, we are referred entirely to conjecture and inference for the origin of the mammals. These point to amphibian-like beings, in which certain peculiarities of the mammalian skull, as the double condyle of the occi- put, were prefigured, and which by the formation of the amnios and allantois approached the true reptiles. These progenitors of the Mammalia are not, however, represented in any order of reptiles or amphibians now extant. The pedigree (p. 269) in which we have grouped the more accurately known fossil Mammalia with those now living, contains considerable gaps, and rests in a great measure on hypothesis, but it gives, never- theless, with approximate probability a correct rep- resentation of the consanguinity of the orders, and in comparison with the system as it was constructed in the school-books prior to the revival of the doctrine of Descent, it must be esteemed a great and suggestive advance. As regards the structure of their skull, the constitu- 260 PEDIGREE OF MAMMALS. “eyEWaTJOUO J sretdnsivyy 1 (srewueyy teqs0"Tg) a vyeyuapa (ferg go sisvag jetpiounsg) (sayenSuq jerprounsg) sinwyy < soyeynsun SeeuM = steag: fay d JO siseag Syeg BIOAI}IOSUT \ skayuop, sjuvydaq = XB YY sual sjuapoy we “aad O1dad 270 THE DOCTRINE OF DESCENT. tion of the pectoral arch, and their persistence in the phase (embryonic in other mammals) in which the rectum and the urinary and genital ducts open into a single cloaca, the Monotremata (Ornithorhynchus, Echidna), limited to Australia and Tasmania, are the lowest mem- bers of their class, and must be considered as remnants of a division reaching from indeterminable past ages down to the present time. It may be presumed that the Marsupials were developed from an analogous grade. Their powers of adaptation have been chiefly testified in Australia, where the subdivisions of the order, usually designated as families, are, in dentition and habits of life, developed in a manner analogous to several of those orders which appear on the second great scene of mam- malian development, namely, the Northern hemisphere. Far advanced beyond the Monotremata as to skeleton, they remain on a low grade with respect to the repro- ductive system, and are implacental, like the Monotre- mata. That is to say, the embryonic blood-vessels do not enter into those close relations with the blood-vessels of the maternal ovary, by which the more perfect develop- ment of other mammals within the mother’s womb is effected. This character and the correlative formation of the pouch in which to carry the immaturely born off- spring, bind together the various families of Marsupials, which deviate from one another like other orders. With the exception, therefore, of the two orders named, in all mammals the embrvo is attached to the ma- ternal organism by the so-called placenta. The blood- vessels of the developing offspring which reach the wall of the uterus by the intervention of the allantois, form coils and loops, between which grow similar offshoots PEDIGREE OF MAMMALS. 271 and appendages of the blood-vessels of the ovary, so that through the walls of the contiguous blood-vessels an abundant exchange of fluids takes place between the two, and therewith a prolonged nutrition and a further and more complete development of the fcetus. The higher character of the placental mammals, usually plainly evinced by their anatomical relations, is thus based on the existence of the placental mass. All inter- mediate grades are, however, wanting which would en- title us to infer with certainty the direct transition from implacental to placental mammals. The Edentata, (Bruta), manifestly the lowest of placental mammals, are so devoid of any nearer morphological relations with the Marsupials, that we must needs be content to as- sume generally, on these indications, supported by geographical distribution and geology, that they repre- sent a very ancient branch of the placental mammals. As we saw in the tenth chapter, they are scattered remnants which can only by compulsion be united into a single order. Sloths, armadilloes, ant-eaters, differ from one another at least as much as rodents, insectivora, and bats. The doctrine of Descent is not discredited because it is unable to account for these fragments of bygone animal life, but in the absence of data it is for the time in presence of an impossi- bility. To ascertain the relationships of other orders, the modern systematizers, and also the supporters of the system of Descent, have thought fit to lay great stress on the presence or absence of the so-called decidua. This requires a short explanation. In many orders of mammals, the vascular processes and villi of the wall 272 TITE DOCTRINE OF DESCENT. of the ovary become so closely connected with the fcetal portion of the placenta, that at birth the entire mem- branous coating of the ovary is detached and thrown out with it. In others, the vascular villi are not so closely adherent; they yield without important lacerations, and hence no deciduous membrane (Membrana decidua) is ejected. Now, as it appears to me, the specific conditions of the formation of the decidua have been far too little compared to justify our inferring any close affinity from the mere fact that portions of the coating of the ovary are lost in parturition. Much rather it must be unre- servedly admitted that the formation of decidua might be occasioned by subordinate circumstances of the most varied kinds, and hence in orders only remotely allied, or allied merely as placental mammals. We therefore consider the decidua to be a subordinate systematic fea- ture where anatomical and morphological reasons are opposed to it. We go yet further. In the modern system the form of the placenta is likewise employed in the grouping of organisms. If among the Deciduata, lemurs, rodents, in- sectivora, bats and monkeys, are classed together as or- ders with discoidal placenta, this combination is cer- tainly supported by a series of other reasons, and it is quite probable that within this group of orders the form of the placenta is due to homology, that is to Descent. But when beasts of prey, elephants and the Daman (Hyrax) are further cited as orders with zonary placenta, we find ourselves in the same position as when the de- cidua was reckoned decisive as to the closer affinity; and we are of opinion that the subordinate form of the placenta might similarly arise in different ways, just as it PEDIGREE OF MAMMALS. 273 has been variously developed in the well-substantiated division of the Ungulata. To corroborate our view by example, we are certainly unable to make any positive statements as to the derivation of the Proboscide. It is, however, none the less certain that nothing positive is implied by the customary classification by reason of their zonary placenta. But we shall more nearly approach the truth if we place this branch of unknown origin typically nearer to the Ungulata than to the beasts of prey. If, moreover, as non-deciduate mammals, the Cetacea are held to be more closely allied to the Ungulata than to the Carnivora, which are -deciduate,—in our eyes, this circumstance is not decisive, as more important reasons argue that the Cetacea were first developed from carniv- orous genera. In our exposition of the geographical distribution of animals, we derived instruction from Riitimeyer with Rhinoceroses. Tapirs. Horses. ~ Hipparion. SQ | Macrauchenidee. S Anchitherium. Paleeotheridze. reference to the relationships of the Ungulata in particular. In no other division do we possess such abundant fossil material. In the older Tertiary strata 274 THE DOCTRINE OF DESCENT. we encounter the remains of two Ungulate families, the Palzotheride and the Anoplotheridz, essentially distin- guished from one another by their dentition, and form- ing the starting-points of the groups of Ungulates of which some now appear so greatly isolated. The root to which these two families lead back is unknown; on the other hand, partly from the direct comparison of these genera with the present Ungulata, partly from numerous intermediate links found in the Miocene, Pliocene, and Diluvium, it appears that, in the lapse of time, the separa- tion which characterizes the present age was initiated, and the seeming isolation was produced by the extinc- tion of the intermediate links. It was this isolation which induced the older systematizers to institute three orders of Ungulata. The special pedigree emanating from thePalxotheride includes, among the present Ungulata, the horse, tapir, and rhinoceros. The transi- tion from the Palzotherium | ff te the horse may be directly traced, and this, moreover, i in the two most important characters, the dentition and ) the feet. In the Anchithe- rium and Hipparion, the transformation from the tri- dactyle to the unidactyle P v7] E Ungulate is accomplished; FIG. 26.—Skeleton of the foot. (P) and Riitimeyer’s brilliant re- Anchitherium. (H) Hipparion. x (B) Hors, searches have shown how, in the milk dentition of each genus, the definitive dentition of the aboriginal genus PEDIGREE OF MAMMALS. 275 is repeated, and Phylogenesis is unequivocally expressed in Ontogenesis. The Anchitherium is a three-toed horse, in which, however, the middle toe has already under- taken the chief task. But in the Hipparion the two side toes are entirely raised from the ground, and by disuse are brought to the condition of arrest which is completed in the horse. In the constitution of the molar teeth the tapirs have remained most faithful to the ancestral type. The cir- cumstance that the tapir has four toes in front, whereas the Paleotheridz known to us, have three shows, how- ever, that the genus Palzotherium cannot have been the ancestral stock of the tapirs. For the supposition that the tapir acquired the fourth toe is contrary to all experience respecting the formation of the extremities. Rhinoceroses are also four-toed in front, and their close kindred with the tapirs is testified by the structure of their toes and a series of details in the skeleton. Hippopotami. Pigs. Tragulidze. Deer. Antelopes. Oxen, Anoplotheridee. An isolated branch of the Palzotheride seems to be the fossil genus Macrauchenide, which combines the 276 THE DOCTRINE OF DESCENT. characteristics of the horse and rhinoceros with those of the camel. How far the latter, as ruminants, are directly connected with the Macrauchenidz, or whether the form of their skull, approaching that of the horse, points to actual homology, it is for the present impossible to say. The Anoplotheridz are likewise distinguished by a sort of undifferentiated dentition, from which a number of specific forms might deviate in different directions. The Tragulide are descended from them in a direct line; they form a small group not unlike the musk ani- mals, and are confined to South Africa and Southern Asia. As chewing the cud, they are more nearly allied to the other typical ruminants with which we are ac- quainted; but, on the other hand, they occupy an inter- mediate position towards the other non-ruminants of the division, of which the whole was united in the pre-historic world through the Anoplotheride. The Suide, or pig- like animals, were very profusely represented in the Eocene and Miocene periods. From a side branch of their predecessors, reaching up to the Anoplotheridz, are descended the river-horses, or hippopotami. The func- tion of ruminating is, as we know, correlated with a com- plex structure of the stomach as well as a peculiar mech- anism of the cesophagal groove. It is naturally im- possible to determine in which fossil animals these arrangements originated; yet it seems to have occurred at a very early period. Perhaps the more highly in- tegrated structure of some non-ruminating genera, such as the hippopotamus and the peccary, may have been transmitted from the age of the Anoplotheridez, and the very conspicuous accordance of the ruminating Tragulide PEDIGREE OF MAMMALS. 279, with the Anoplotheridz stamps the latter with tolerable certainty as ruminants. Disregarding the camels, already mentioned as of doubtful origin, the typical ruminants separate into the deer-like and the horned. Through the hornless musk animals, the deer are connected with the Tragulide and the older genera. The giraffes form a side branch. But although the Helladotherium, nearly allied to the giraffe and at one time inhabiting the Athenian territory in herds, and the colossal Sivatherium, found in the spurs of the Himalayas, afford some clue to the position of the giraffes, so entirely isolated in the present world, the details of their derivation still remain very obscure. From the antelopes to the closely allied—in fact, scarcely separable—genera of goat and sheep, and similarly to the oxen, the systematic as well as the pale- ontological series, and likewise the ontogenetic phases, present transitions undeniably evincing family relation- ship. Besides the relations of the milk dentition of the filial to the ancestral genera, which Rititimeyer has also followed minutely, great interest attaches to the gradual transformation of the skull, which reaches its extreme in the oxen, and advances from the antelope and sheep, through Ovibos, Bubalus (buffalo), Bison, to Bos (ox). In the latter, the erect position of the frontal bone attains its utmost grade, and this transformation of the skull of the antelope is repeated in the calf. The usual classification of the Sirenia, or sea-cows, with the Cetacea, was decidedly a systematic misconcep- tion, arising from one-sided and, moreover, merely su- perficial consideration of the locomotive organs. All other characteristic indications—above all, the structure 278 THE DOCTRINE OF DESCENT. of the skull and the nature of the teeth—remove them from the Cetacea as much as they approximate them to the Ungulata. In the hippopotamus we have a mem- ber of this order nearly converted into an aquatic animal. We must think of the Sirenia as originally emanating from some unknown genera, which probably branched off at a very early period. A very uncertain position is occupied by the Hyra- coidz, now represented only by a few species of the genus Hyrax. To say that their characteristics recall at once the Ungulates, the Rodents, and the Insectivora, affords no explanation. Considering the great impor- tance of the molar teeth in deciding derivation, the chief stress should perhaps be laid on their similar- ity in the hyrax and the rhinoceros, and we hence regard the hyrax as an offshoot of an old Ungulate family. With respect to the progenitors of the Proboscidz, we refrain from any conjecture. Later than the Graminivora, the Carnivora, and espe- cially the beasts of prey, seem to have appeared on the scene of arctic animal life. ‘Granting the possibility (and it is scarcely possible to do otherwise) that placental for- mations may have originated in various ways, the pos- sibility likewise exists that the Carnivora, and indeed other orders too, such as the Rodents especially, may be direct descendants of carnivorous Marsupials. The oldest beasts of prey known are feline, or resemble the Viverride and hyenas. Then come the Canide, and latest of all the Ursidz. In skull, dentition, and extremi- ties, the seals and walruses (Pinnipedea) constitute a side branch. Although there can be no idea of any special PEDIGREE OF MAMMALS. 279 affinity between the otter and the seal, the comparison of the two will aid us in imagining how from true beasts of prey and terrestrial animals the strange figure of seals and walruses must have proceeded. If the conjecture already propounded should be con- firmed, that the detachments and ejections of the placenta, which constitute the phenomena of the decidua, assume very heterogeneous forms in groups belonging to the same family, and may be alike in others no more nearly related, the Cetacea would he installed in our pedigree in the vicinity of the beasts of prey. Between a lion and a whale an angle is enclosed, containing a countless multitude of intermediate forms. But we must always bear in mind that our business is, not to bridge over the chasms between the present peripheral ends of the series of development representing the extreme forms, but to discover the points of derivation and attach- ment. Fossil whale-like animals are known in the Ter- tiary period, such as the Zeuglodon and Squalodon. The remains of the former colossal genus are kept in good preservation at Berlin, where Johannes Miiller dis- covered their relations to both seals and whales. The dentition is seal-like; in the skeleton there is much similarity with the whales; and although the Zeuglodons must have been preceded by a great series of species, and followed by another of considerable, if not equal, length, before the present Cetacea proceeded from them, a development of this sort seems, nevertheless, extremely probable and natural. By their still perfect dentition and the still proportionate dimensions of the skull, the Delphinoidz are the oldest members of the true Cetacea. They were joined by the sperm whales or cachelots I, 280 THE DOCTRINE OF DESCENT. (Physeteridz), and the last members are the right whales (Balzenidz). This is evinced by the fact that the ‘whale- bone or baleen plates are developed only after the ru- dimentary teeth have made their appearance in the jaws of the embryo, a heritage from the profusely and per- sistently toothed ancestors. In the Lemuridz, the system unites the heterogeneous remains of a collection of animals which, by reason of their prehensile hind feet with their opposable hallux, were regarded as fellow-members of the order of “true apes.” The connecting link is not their anatomical con- stitution—they diverge widely in the form of the skull and in dentition—but rather their geographical distribu- tion, restricted to Madagascar and a few advanced posts of Asia. Undue influence has also been allowed, cer- tainly very unscientifically, to a certain peculiar outlandish impression which they make upon the observer. The constitution of their skull refers them to a very low grade in the scale of the mammalia. If we view them as a whole, they exhibit no general relations with any particular order of mammals, but, according to the in- dividual genera, point to those orders which, like them- selves, possess discoidal placenta; the majority of rea- sons favour the hypothesis that the Lemuridz now living are the last and little modified offshoots of a division of "mammals at one time far more richly developed, and that Rodents, Insectivora, Cheiroptera, and Apes, are twigs of this great branch. The Rodents are particularly interesting, because, in conjunction with stubborn persistency in the very characteristically constituted dentition, accompanied by several peculiarities of skull, they manifest the most ex- PEDIGREE OF MAMMALS, 261 traordinary power of adaptation to arboreal and steppe- life, to land and water. The Insectivora, although not nearly so rich in species, offer a similar spectacle of adaptations by which their genera have become almost repetitions of the Rodents; and the Cheiroptera (bats), in their most numerously represented division, may be regarded as a side branch of the Insectivora, if they have not proceeded directly from animals resembling the Lemuridz. In what geological period the monkeys were evolved from lemur-like forms we do not know. The few fossil monkeys with which we are acquainted belong to the higher families of apes, and pre-suppose a long series of ancestors. The same conjecture is forced upon us by the geographical isolation of the American monkeys from those of the Old World, which is also combined with considerable anatomical differences, although it could not occur to zoologists or comparative anatomists to deny their close systematic affinity. The relation of the lower to the higher apes requires further discussion, which we shall combine with our dis- quisition on the relation of man with the monkeys. 282 THE DOCTRINE OF DESCENT. XII. Man. WHEN Goethe declares, “ We are eternally in contact with problems. Man is an obscure being; he knows little of the world, and of himself least of all,” 7?—he almost repeats what J. J. Rousseau says in Emile,” “We have no measure for this huge machine (the world); we cannot calculate its relations; we know neither its primary laws nor its final cause; we do not know ourselves; we know neither our nature nor our active principle.” Such and such-like quotations are wont to be made to us as justifying and confirming assertions of the nar- rowness of our powers of understanding, and of the limits of science. But in Anthropology we cannot possibly at- tribute any greater authority to the worthy J. J. Rous- seau than to a Father of the Church; and to the Goethe, whose casual utterances are transmitted to posterity by Eckermann, we oppose the other Goethe, who in the fulness of youthful vigour, exclaims— Joy, supreme Creation of Nature, feeling the power . All sublimest thoughts, which lifted her as she made thee, In thyself to re-echo——* and who conceives the most beautiful organization, as he * Freue dich, héchstes Geschépf der Natur, du fiihlest dich fahig Ihr den héchsten Gedanken, zu dem sie schaffend sich aufschwang, Nachzudenken——™ THE DOCTRINE OF DESCENT. 283 designates man, to be in perfect harmony with these sublimest thoughts. Our previous reflections and deductions would lack their conclusion were man to be excluded,—could not and must not all that is said of the genesis and connec- tion of animal being, be directly applicable to the knowl- edge of his nature also. The repugnance to the doctrine of Descent, the doubt with regard to it, the indignation lavished upon it, are all concentrated on its applicability and application to man; and if the body be perforce abandoned to us, the mental sphere of man is at least to remain inscrutable, a molt tangere to the investigation of nature. A few years ago, it was a consolation to the opponents of the doctrine of Descent that Darwin had not directly pronounced himself with respect to man. Anger was vented on his adherents, who had outdar- wined Darwin. To this was added the unfortunate mis- apprehension that the champions of the doctrine of Descent made the human race proceed from the ennoble- ment of the orang, chimpanzee, or gorilla—in short, from extant apes. But from the first appearance of the Darwinian doc- trine, every moderately logical thinker must have re- garded man as similarly modifiable, and as the result of the mutability of species; and Darwin has now told us, in his work on the “ Descent of Man,” why he did not enunciate this self-evident inference in his first book; he did not wish thereby to strengthen and provoke preju- dice against his view. Knowing human weakness, he. withheld the conclusion. ‘It seemed to me sufficient,” he says, “in the first edition of my ‘ Origin of Species,’ that by this work ‘light would be thrown on the origin 284 THE DOCTRINE OF DESCENT. of man and his history,’ and this implies that man must be included with other organic beings in any general con- clusion respecting the manner of his appearance on this earth.” Nay, Darwin himself has now gone further, and, to the terror of all who can scarce imagine man except as created shaven and armed with a book on etiquette, he has sketched a certainly not flattering, and perhaps in many points not correct, portrait of our presumptive ancestors in the phase of dawning humanity. Before we seriously discuss this serious subject, we will take leave to quote a more superficial verdict given by a clever essayist.*4 “ Let us suppose, merely as a joke, that Nature, which we see everywhere advancing from the most simple to the compiex, from the lower to the higher, had not suddenly waived this law in the presence of man; that she had not suddenly given up her evolution for his sake; that she had not suddenly begun in him a new creation; but that here, as elsewhere, she had proceeded quietly, gradually, naturally, and that man was thus nothing more than the last link of the in- terminable series of animals, nothing more than a ‘ de- veloped ape.’ The first thought that would then ob- trude itself upon us, would be that the facts were not altered in the slightest degree; that man would remain as he is, with the same shape, the same face, the same gait, the same gestures, the same dispositions, powers, feelings, thoughts, and with the same dominion over the apes as heretofore. This is very simple, very self-evident, but also very important. For it confers on him—on man—the powerful sensation that, as he now is, he is a being of a quite peculiar kind, very different from SUPERFICIAL OBJECTIONS. 285 even the most kindred creature; and, moreover, that this peculiar nature is his most peculiar property, whether he received it as a ready-made gift, or worked it out laboriously from a lower condition in tens of thousands of years. But if his present constitution is not in the slightest degree injured by his (assumed) animal origin, neither can his aims and tasks, his endeavours and voca- tions—in short, his whole future—be any other than, from his entire nature, he must imagine and believe it to be. Or must the cultivated portion of mankind be really so profoundly dismayed by the idea of descending from apes, that in despair at the impossibility of maintaining and improving the civilization, which by no means fell into their lap like ripe fruit, but which was painfuliy acquired, they would abandon their business and pur- suits, their forms of law and government, their arts and sciences, and sink to the level of the Australian bush- men—that they would let go that by which they had raised themselves so far above the apes, and by which they are constantly raising themselves still higher, merely because it was once difficult to raise themselves above these apes even by a hair’s breadth? But what man destined by nature for a ruler, would have refused to grasp the crown because his father was a hind? Or what born Raphael would have forsworn palette and pencil, because his parent had been a sign-painter? Mankind, like each individual, will use and improve its powers be- cause it has them, not because it has obtained them from hither or thither.” We give these transient fireworks their due, but we require more profound arguments whence to derive the final verdict. To the votaries of the doctrine of Descent, ' 286 THE DOCTRINE OF DESCENT. its application to man is a simple deduction from a gen- eral law, gained by the method of induction. As Goethe postulated the inter-maxillary bone in man even before he had seen or proved it, so must the doctrine of Descent extend to man all its results and more or less plainly demonstrated laws. The deduction is effected by the accumulated observations of comparative anatomy, evo- lutionary history, and paleontology, checking and con- firming one another. Thus, for all who are not satis- fied with belief in miracle and subjection to the hy- pothesis of a revelation, nothing remains but the doctrine of Descent. To apply it to man is not more hazardous, ‘but, on the contrary, as inherently necessary, as it is for us zoologists to make use of it in judging some polype hitherto unknown, a star-fish or a mouse. This our adversaries deny. Man, they say, has qualities which separate him absolutely from the animal, and, assum- ing the doctrine of Descent generally, preclude its ap- plicability in this one case. To this assertion, so fre- quently to be heard, we will, in the first instance, oppose a general remark as to the apprehension of human nature.. It is commonly overlooked that, quite regardless of the validity of the doctrine of Descent or even of its existence, there is a notable inconsistency in the idea of humanity. The philosophy of history has regarded mutability, which is, in fact, capability of progress, as the essence of human nature. But if any sort of in- separable dependence of the mind upon the body be admitted, as is the case with all but an extreme spirit- ualistic party, the progress of mental power in mankind was inconceivable without some parallel transformation THE BODY OF MAN. 287 of the bodily substratum extending beyond the limits of mere variability. Even on the assumption that the mind forms its own organ, the brain, the specific idea of man would necessarily have consisted in bodily im- provement, as contrasted with the supposed rigidity of the animal organism. For, in principle, it is the same whether changes take place perceptibly in arms and legs, or imperceptibly to the eye, in the molecules of the brain. We are, therefore, only retrieving the shortcom- ings of philosophy when we attribute to the bodily muta- bility of man the extension which accrues to it from the applicability of the doctrine of Descent to the particular case. The bodily accordance betwixt man and animal leaves the doctrine of Descent so little to desire, that the ap- prehension of Mephistopheles lest grovelling humanity should finally be alarmed at his likeness to the Deity, might far rather be applied to his likeness to the animal. The human body, like the body of every animal, points in its evolution to an elaboration from the undifferen- tiated to the specialized form. The general distribution of the body and the development of the several organs is common to man and all mammals, and in the earlier stages of the embryonic state to all vertebrate animals, and indicates this general kinship. The existence there- fore of a discoidal placenta (unless we prefer a special reiterated new creation of this organ of development, in which the Creator adhered to the pattern of the placenta of the lemurs, rodents, insectivora, bats, and apes) reduces us to the alternative that in the natural and to us un- known development of man, chance, or some quite different chain of causes, led in this case, as in the other, 288 THE DOCTRINE OF DESCENT. to the discoidal placenta, or that the accordance is based on consanguinity with the discoido-placental mammals. We have already (p. 272) objected to the inference that all mammalian orders are akin, should be drawn with certainty from the superficial accordance of the placenta, and we must therefore justify ourselves now, when we lay a stress on the accordance of the placenta of man and apes. The orders mentioned above all possess a placenta of small extent and discoidal form. In the shape of this disc, and in the number and distribution of the blood-vessels in the umbilical cord by which the foetal respiration and nutrition are carried on, sundry varieties occur. Thus in the family of the Pithecoid apes, the placenta falls into two discs, whereas the um- bilical cord agrees with that of man; in the American apes, on the contrary, the placenta is simple and the blood-vessels are different. In the orang and gorilla we know nothing of these organs, but the chimpanzee agrees with man, in that it has a simple discoidal placenta with two conducting (arteriz umbilicales) and one reconduct- ing vessel (vena umbilicalis). With a general similarity of the human placenta with that of the discoido-placental mammals, man is especifi- cally nearer to one at least of the so-called Anthropoid apes, than this one is to the other apes. And thus the constitution of the placenta is certainly of great impor- tance in discriminating the systematic position of man. Enormously improbable as is the chance contemplated above, equally probable and solely credible is consan- guinity; and with regard to general organization, in any specific comparison of man with the mammalia, the apes must occupy the foreground. MAN AND APES. 289 This comparison has been admirably conducted by Huxley and Broca.*? The latter has set himself the task of investigating, solely as a descriptive anatomist and zoologist, regardless of all dispute as to principle, and undisturbed by the doctrine of Descent, whether the anatomical constitution of man, as compared with that of the ape, justifies, on general zoological principles, the union of the two in a single order—Primates. Huxley proves that the anthropomorphous apes (gibbon, chim- panzee, orang, gorilla) differ from the lower apes much more than from man; and that if we are obliged to as- sume the reciprocal consanguinity of the apes, the com- mon derivation of the anthropomorphous apes and man is at least equally natural. Between the peripheral members of the systematic groups of monkeys—for instance, between the American Sahuis and the Old-World Pavians and Anthropomorpha —notable differences exist in the constitution of the limbs and other parts of the skeleton, together with the soft parts belonging to them, in the muscles especially, as well as in dentition and the structure of the brain. It is false to call apes quadrumana, for within the order of the apes the contrast between hand and foot makes its appearance in its essential anatomical attributes, and in the anthropomorphous apes, in the gorilla especially, it is almost as distinct as in man. Luca, the anatomist renowned for his careful meas- urements of the cranium, imagines that he has discerned a highly important demarcation between man and the ape. In the ape, the three bones forming the axis of the skull, the basi-occipital bone, and the two sphenoid bones, lie almost in a line, whereas in man there is a 290 THE DOCTRINE OF DESCENT. double flexure of this axis; moreover, in the apes the angles increase with age, which in man decrease, and vice versé. Likewise in man the occipital foramen be- comes more horizontal with age, more vertical in the ape. But all this shows only, what the doctrine of Descent asserts, that the two series, ape and man, diverge from one another, and that the youthful individuals are more alike than the older ones,—that the ape as he grows be- comes more bestial; man, as the riddle of the sphinx already intimated, more human. The flexure of the basal bone and the horizontal position of the occipital foramen occasions the upright gait, wherewith the dif- ferentiation between hands and feet is completed. This flexure of the cranial axis may therefore still be em- phasized as a human character, in contradistinction to the apes; the peculiar characteristic of an order can scarcely be elicited from it; and especially as to the question of Descent, this circumstance seems in no way decisive. Not only as regards hand and foot, but also in denti- tion and brain, the anthropomorphous apes approach man much more nearly than they do the inferior wide- nosed monkeys of the New World. These, have six molar teeth, and their brain displays the imperfections of the brain of the lemurs and rodents. Like the mon- keys of the Old World, on the contrary, the anthro- pomorphous apes possess five molar teeth, and every portion of the human brain, even to the hippocampus minor, is likewise present.. The dispute as to this insig- nificant portion of the brain, which R. Owen claimed as an exclusively human characteristic, possesses a merely historic interest, since, in conjunction with the posterior MAN AND APES. 2g1 corner of the lateral ventricle, it has been exhibited by a number of distinguished anatomists in the orang and chimpanzee. Thus, for those who will not relinquish their hope of finding specific distinctions between the brain of man and ape, there remain only the furrows and ridges on the surface of the cerebrum, the so-called convolutions of the brain. But here, again, it is in vain to look for funda- mental differences, unless the chief stress is to be laid upon the circumstance that in the human embryo the folds commence in the frontal, in the apes in the supra- orbital lobes. The constant convolutions common to all human brains are seen in the orang and chimpanzee. These convolutions are lost, or rather exist in less per- fection, in the apes approaching the Anthropomorpha; they are totally absent in the Ouistitis. But so great is the resemblance of the brain of the two apes mentioned, with that of man, that, as Broca says, “it requires the eye of an experienced anatomist to discriminate, in drawings reduced to the same dimensions, their brain from the human brain, especially if the object of com- parison selected, be the brain of negroes or Hottentots, which are more simple than those of white men.” A desperate attempt to rescue a specifically human cerebral character was made by the lamented Gratiolet, the anat- omist, of Paris. Man was to be distinguished by the so-called transitional or bridging convolutions. These transitional folds are convolutions, by which the pos- terior lobes of the cerebrum are joined to the anterior and lateral portions. But Broca has lucidly demon- strated that it is the same with this as with other char- acteristics, and that the transitional folds in the orang, 292 THE DOCTRINE OF DESCENT. for instance, are far more like those of man than of the chimpanzee, and that the differences which exist can at the most have the value of specific or generic char- acters, , The distance between the lower and higher apes is far greater than between the latter and man; and if the consanguinity of the entire apedom is decisive in favour of Darwinistic views, there can be the less doubt of the kindred connection of the Old-World apes to mankind. But the form of the mature skull and of the dentition (to lay a stress upon these organs), preclude the idea that the direct ancestors of man are to be found among the apes now living. The cheap jest, produced with so much glee, of inquiring why we do not behold the in- teresting spectacle of the transformation of a chimpanzee into a man, or conversely, of a man by retrogression into an orang, merely testifies the crudest ignorance of the doctrine of Descent. Not one of these apes can revert to the state of his primordial ancestors, because, except by retrogression—by which a primordial condition is by no means attained—he cannot divest himself of his acquired characters fixed by heredity; nor can he ex- ceed himself and become man; for man does not stand in the direct line of development from the ape. The development of the anthropoid apes has taken a lateral course from the nearest human progenitors, and man can as little be transformed into a gorilla as a squirrel can be changed into a rat. Man’s kinship with the apes is, therefore, not impugned by the bestial strength of the teeth of a male orang or gorilla, or by the crests and protuberances on the skulls of these animals. A renowned zoologist, one of the few who adhere to the MAN AND APES. 293 old belief, has taken the useless trouble of proving that the skull of the orang could not possibly be transformed into the human head. As if the doctrine of Descent had ever asserted such nonsense! The bony skull of these apes has reached an extreme, comparable to that of our domestic cattle. But this extreme appears only grad- ually in the course of growth, and the calf knows little of it, but possesses, as we have already mentioned, the cranial form of its antelope-like ancestors. In the present antelopes, and likewise in goats and sheep, this form, transitory in the calf, has remained stable. Now, as the youthful skull of the anthropomorphous apes exhibits, with undeniable distinctness, a descent from progenitors with a well-formed and still plastic cranium, and a den- tition approximating to that of man, the transformation of these parts in conjunction with the brain, the latter by reason of its persistently small volume, has, as it were, struck out a disastrous path, while in the human branch, selection has effected a higher conservation of these cranial characters. With this falls also the objection recently raised by the venerable Karl Ernest v. Baer, that it is inconceivable how, from.the monkey’s feet, arranged for climbing and grasping, the human foot, adapted for flat treading and walking, should be evolved in the struggle for existence. The tendency to oppose the big toe to the others, that is, to a prehensile foot, is known to be a human attribute, and this tendency is certainly inherited. How far the capacity for climbing may have been developed in the primordial ancestors, is as much unknown as these pri- mordial ancestors themselves. Thus the aptitude in climbing shown by most of the present monkeys is only 294 THE DOCTRINE OF DESCENT. remotely connected with the inaptitude of man, and as a criterion of consanguinity, can hardly be taken into con- sideration. While requiring by logical deduction, a common origin for man and the anthropomorphous apes, the doc- trine of Descent, as it is almost superfluous to say, repu- diates the senseless demand for intermediate forms be- tween man and the gorilla. What future times may perhaps discover, are intermediate forms which go back to the common point of derivation of the present apes and of man. And thus, notwithstanding the very close relations already discussed, there remains the chasm which is approximately expressed by the comparative weights of the lowest human brain yet measured and the brain of the gorilla. The brain of a bushwoman, nor- mally efficient after the manner of her tribe, amounted to 2 lbs. 4 ozs. (Cuvier’s brain weighed 4 Ibs. 4 ozs.), that of a gorilla may be estimated, from the capacity of the cranium, at about 1 lb. 6 ozs., which gives the approx- imate ratio of 3:2. But exalted above the animal as man may feel himself in his bodily nature, in this again he forms no exception, as many animal forms occupy an equally isolated position with reference to their unmis- takably nearest kindred. ' Need we imagine a twofold creation of vertebrate animals, because the lancelet is now separated from the fishes by a whole scale of intermediate forms no longer extant? The example of the horse is, among others, highly instructive in this case. Let us bear in mind that, in the nature of the limbs and teeth, this genus differs far more from all other extant graminivora than man differs from the ape. Had not the fossil ungulates been DEFICIENCY OF INTERMEDIATE FORMS. 295 found which demonstrate the common origin of the horse with the didactyles and the multidactyles, we should still not deem the horse a special miraculous creation, but incontrovertibly deduce his true kinship with the other ungulata. This pure deduction is not requisite, as the progenitors of the horse are present in conspicuous re- mains; and, as we have already seen, elicited in R. Owen, half a century ago, the conviction of a direct metamor- phosis of the tridactyle genera into the unidactyle. Our acquaintance with the tridactyle horses was a lucky chance; they were indigenous in those parts of Europe which have been most diligently laid bare and explored in behalf of Palzeontology. But that our museums are still destitute of the fossil progenitors of man, is not more strange than the defi- ciency, hitherto existing, of intermediate forms, which, for example, would conclusively decide the position of the Dinotherium in the system. We will also refer again to the elephants, who, with their nearest ally, the mas- todon, occupy towards the other Pachydermata a position elucidated by no fossils, and far more isolated than that of man to the apes. We hope herewith to have shown that the argument that, by peculiarities not bridged over, —by upright gait, comparative hairlessness, chin, prepon- derance of brain, &c.,—man betrays a position absolutely apart, cannot be admitted by comparative anatomy and paleontology. The demand, therefore, that the ad- herents of the doctrine of Descent should produce the intermediate forms which at one time necessarily existed, can be made only by dilettantes to whom the province of life, as a whole, has remained a sealed book. As we observed before, the bodily nature of man is 20 296 THE DOCTRINE OF DESCENT. sometimes ceded to natural inquiry as a means of more certainly rescuing the other side of the dualism. But here, too, we will not be defrauded of our say and our own opinion. The mental powers of man, in their origin, growth, and effects, are likewise susceptible of investiga- tion, and psychology only too long thought it possible to elude physiology. Let us, therefore, proceed in good heart to a short examination. It is universally admitted that a certain relationship, or analogy, exists in the psychical capacity of the higher animals and man. Reason alone, it is said,—the essence of psychical agencies by which man attains self-con- sciousness, and rises to abstract conceptions, combines ideas, especially religious ideas, and lives in art and sci- ence,—this the animal does not possess. We reply that animals certainly do not possess this degree of mental development, but neither does man possess it in lower phases of evolution. The soul of the new-born infant is, in its manifesta- tions, in no way different from that of the young ani- mal; these manifestations are the functions of the in- fantine nervous system; with this they grow and are de- veloped together with speech. The grade to which this development rises is generally dependent on the preced- ing generations. The psychical capacities of each indi- vidual bear the family type, and are determined by the laws of heredity. For it is simply untrue that, inde- pendently of colour and descent, each man, under con- ditions otherwise alike, may attain a like pitch of mental development. As a proof of this primary equality of mankind, single instances of gifted negroes and Indians are held up to us. But these have behind them un- MENTAL PROGRESS. 297 numbered generations practised in. multifarious employ- ments, skilled in human intercourse, even if it be one- sided; and if these rare phenomena are thoroughly in- vestigated, they still remain behind the average individ- uals of advanced races. Now it is certain that in each race, each individual passes through grades in the scale of mental development, which, in perfect analogy with the laws of anatomical development, are uni- versally valid; whereas, as we have seen, the psycho- logical peculiarities of the race become valid. But it is in mankind as in the individual; in the lapse of time, it has acquired those higher powers of mind which we call Reason. History shows, and no one denies, a mental advance, but only in nations which have taken part in history, and only so long as this part and the exercise of the mental organs has been continued. But inferior human races exist—we may also call them human species— which are related to the others, as are lower animals to higher. It might even be given as the characteristic of the genus man, that its species occupy such extraor- dinarily different grades of mental condition. We are not misled by the contrary statements of missionaries and other philanthropists; by the talk of human dignity and divine resemblance; nor do we seek for consolation in the development still to be expected in all nations which have hitherto lagged behind. It is indeed self- evident from the theory of descent and selection, that many of the races now standing far behind in a mental point of view will in future have made a great advance. But for others, if we contemplate the ethnology and anthropology of savages, not from the standpoint of 298 THE DOCTRINE OF DESCENT. philanthropists and missionaries, but as cool and sober naturalists, destruction in the struggle for existence as a consequence of their retardation (itself regulated by the universal conditions of development), is the natural course of things. If we examine the mental condition of mankind, and compare it with the psychical capacities of animals, we must not take the average Indian or European as our standard, but the Australian and Papuan races, which in body also have remained at a grade which the other more favoured races outgrew in pre-historic times long past. Many, it is true, overcome all difficulties, inas- much as, assured of an equalizing human dignity as of a dogma needing no further foundation, they are ready with the assertion that it is impossible to doubt that they have retrograded from a higher mental develop- ment and sunk into barbarism. But if this possibility might be admitted as regards some few races, such as the Fuegians; in others, in the Australians for instance, any real evidence of this previous more elevated condi- tion is wanting. The superior mental prerogatives which are supposed to separate man from the animal, hinge more or less on the following points. Man alone, it is said, is capable of development or progress. Specifically, human is all progress regulated and effected by human speech, for many animals like- wise possess the gift of communication. But if we are not to imagine man as having advanced from all eternity, the question is, how was this advance initiated, and the whole concern is fundamentally reduced to the problem of the origin of language. We will return to this subject. FREE WILL. 299 Progress in general is not however to be denied to the animal. Who can question that some canine races, of which the descent from stupid jackals and wolves is as good as certain, have raised themselves mentally far above their ancestry? Who, that has read the comprehensive investigations of H. Miller, the brother of our Fritz Miller, can doubt that the honey-bee, as it gradually attained its bodily advantages and peculiarities, devel- oped likewise the higher mental powers, corresponding with the more minute and complex organism of her brain? Man—such is the thesis we propound, reserving the question of language—differs from many animals only in the degree and means of progress. It is therefore unscientific to contrast humanity and animality in the abstract. Man alone, it is further maintained, has a free will. In so far as the more highly developed man acts in accordance with philosophical, moral and religious prin- ciples, for which he is indebted to education and in- struction—in so far as he is able to apprehend ideals, and strive after them with his own mental and bodily power, this command of will may be readily admitted, although we know that this “ freedom” is likewise the collective result of natural causes. But the more simple and uniform the conditions of life, the more do the deal- ings of men lose the semblance and character of free- dom, and the more does the individual act after the will of the tribe—I might say, of the herd—that is to say, instinctively. In this case actions are not performed even with the astounding premeditation with which some few happily organized indlvidual animals of some few species turn the circumstances to account with apparent- _300 THE DOCTRINE OF DESCENT. ly complete free will. The free will of the morally ele- vated man, is no common property of all mankind. Man alone, and all men, are supposed to have a con- science. We consider, on the contrary, that conscience, which is known to be utterly lost in many individuals of even the most civilized nations, is, like moral will, a result of education in some few races and tribes. Fear of detection after a bad action, is not conscience; and that well-trained dogs have sensations of conscientious shame far superior to the animal terror of savage can- nibals after they have wrought the murder of their fel- . low-men, it is impossible to deny. Of this, evidence in profusion is accumulated in the anthropological compila- tions of Waitz. , That a consciousness of the Divine existence is a fundamental property of all men, we likewise hold in question. It is, again, an established phrase that the most barbarous nations are guided by emotions and cravings, however obscure, towards the unknown God. This as- sumption is as old as the well-known attempt to prove the existence of God, “ De quo omnium natura consentit, id verum esse necesse est” (That in which all intuitively agree, must necessarily be true). How often has this saying of Cicero been thoughtlessly repeated? This idea of God is, however, as little intuitive as the discrimina- tion of good and evil by the conscience. Others main- tain the contrary. Thus Gerland says of the Australians: 8° “The statement that Australian civiliza- tion indicates a higher grade is nowhere more clearly proved than here (in the province of religion), where everything resounds like the expiring voices of a pre- vious and richer age; but we in no way receive the im- AUSTRALIAN IDEA OF GOD. 301 pression that we are dealing with stagnation or incom- plete development. Thus the idea that the Australians have no trace of religion or mythology is thoroughly false. But this religion is certainly quite deteriorated, and has degenerated into a wild, disjointed, and often incredibly absurd demonology, into a superstitious fear of apparitions.” But when a few lines later in the work quoted, we are informed that the natives to the west of the Liverpool range, ascribe everything in nature which they cannot ex- plain to the Devil-Devil, and that this is manifestly only a name, derived from the English Devil, for a Deity of whom they: have not preserved any distinct conception, the shallowness of this evidence in favour of the hypothe- sis of a previous standpoint, now sunk into oblivion, en- ables us to infer the value of the other instances. We have far more reason to believe this low state of mental development in harmony with the bodily condition, when we hear that the natives of the Gulf of St. Vincent and the neighbourhood of Adelaide are extremely hairy, and that even the brown-coloured down of the children is so abundant and so long, that the skin of boys of five or six years of age assumes a furry appearance. But, contrary to all experience and history, we are required to believe ** that the inhabitants of the northern parts of Australia are the most aboriginal, for ‘they are the most civilized, as well as the best developed, in mind and body; they only are fixed in one dwelling-place; and in any case the sup- position is easier and more natural that the other natives should have degenerated, with their eternal wanderings, than that the former, fixed by the more convenient terri- tory, should have raised themselves.” 302 THE DOCIRINE OF DESCENT. This inverts all that has hitherto been called anthro- pology. Moreover, there are even very advanced na- tions without any consciousness of God. Schweinfurth relates that the Niam-Niam, that highly interesting dwarf people of Central Africa, have no word for God, and therefore it must be supposed, not the idea; and Moritz Wagner has given a whole selection of reports on the absence of religious consciousness in inferior nations. When, in spite of all these corroborations, it is always re- torted afresh that even among the lowest savages some sort of feeling of superior powers is manifested, the dis- pute finally results in mere verbal criticism, which has no farther interest for the doctrine of Descent. And yet we cannot leave this subject without alluding to a fact, universally known, but, strange to say, not as yet employed in this connection, and which, as it would seem, is by itself sufficient to invalidate the assertion that the idea of God is immanent in human nature. We mean the fact that many millions in the most cultivated nations, and among them the most eminent and lucid thinkers, have not the consciousness of a personal God; those millions of whom the heroic David Strauss became the spokesman when he adopted for his own the motto of his favourite, Ulrich von Hutten: I have dared it— Jacta est alea! And now as to Language? All modern philologists agree that languages are developed, and that most prob- ably all linguistic families pass through three stages. In the stage of the radical languages all words are roots, and are merely placed side by side. In the second stage, that of the agglutinated languages, one root defines the other, and the defining root ultimately becomes merely DEVELOPMENT OF LANGUAGE. 303 a determinative element. Finally, in the inflected lan- guages, the determinating element, of which the deter- minating significance has long vanished from the national consciousness, unites into a whole with the formative ele- ment. As we have said, this development, in which retrogression takes an extensive share, is universally ad- mitted. Opinions differ only as to the origin of the lin- guistic material, which the acuteness of the philosophers extracts in the guise of “roots.” A great authority, Max Miiller,®* discerns in the existence of the roots evidence of the absolute separation of man from the ani- mal. While Locke says that man is distinguished from the animal by the power of forming general ideas, the philologist ought to say that human language is distin- guished from the animal capacity of communication by the power of forming roots. To trace up all words to imitation and exclamatory sounds is inadmissible, as we most frequently come upon roots of fixed form and gen- eral meaning which are inexplicable in themselves. He deems the existence of these ready-made roots, before which linguistic science stands helpless, an insurmounta- ble impediment to the apprehension of man as a link in the general evolution of organisms. This point excepted, this excellent scholar naturally admits all those phenomena of heredity, acquisition, and degeneration, which are manifested in the laws of lan- guage, and find the?r most perfect analogies in our doc- trine of Descent. If, for instance, we compare Zend with Sanscrit, and hear several of its words explained, we are at once reminded of the rudimentary organs and their significance. A host of anomalies are, like the isolated organisms of present times, primeval and pe- 304 THE DOCTRINE OF DESCENT. culiarly normal remnants and witnesses of bygone lin- guistic periods. In short, down to the minutest details, linguistic research stumbles on accordance and analo- gies with the doctrine of the derivation of organisms. And, forsooth, we are to halt before the origin of lan- guage as before a something incomprehensible and in- scrutable! This is not done, however, by the majority of com- parative linguists in the present day. Though Max Miller calls the roots “ phonetical fundamental types produced by a power inherent in human nature,” though, according to him, man in a more perfect state possessed the power of giving to the reasonable conceptions of his mind a better and more subtle expression, the talented Lazarus Geiger ®’ terms the hypothesis of a now extinct power of forming languages, and the other hypothesis connected with it, of a primordial state of higher perfec- tion, a recourse to the incomprehensible and a return to a standpoint of mysticism. For that which is not un- derstood is not necessarily incomprehensible. It is not our business to side with Geiger, who attributes an essen- tial share in the ejaculation of words to the visual per- ceptions, or with Bleek, G. Curtius, Schleicher, Steinthal, and many others, who assign to the imitation of sounds the first place in the evocation of language. This much is, however, certain, that although those who are not critical, find Max Miiller’s standpoint highly convenient, in science, it is unique. In this province, interwoven as it is with the investigation of nature, the greater number of authorities, on linguistic grounds, comparative and philosophical, have been forced to the conclusion that, from an irrational primordial state, man-like beings grad- LANGUAGE AND MIND. 305 ually became human, while with language, the work of many years, reason made its appearance. As early as 1851, when the doctrine of Descent was still unheard of, Steinthal 8° says: “As language arises, mind originates.” Ten years after Darwin, Geiger writes: “Language created reason; before language, man was irrational.” To him, and to all who have aban- doned the standpoint of mysticism, “man is a genus springing from an animal condition by means of the origin and unfolding of his idiosyncrasy.” And this con- clusion is not, as orthodoxy and reaction are anxious to impress upon the multitude, borrowed from Darwinism, but deduced from linguistic inquiry in its own way, only by a scientific method. It need only be indicated that, as Geiger has historically proved in so many instances, “slow development, the emergence of contrast from im- perceptible deviations, is the cause that the same word ac- quires various meanings;” that the creation of language therefore rests upon this process, and nowhere makes its appearance suddenly and abruptly; that the so-called laws of sound are habits of sound; that the special meaning which a sound has acquired in lapse of time is always the result of mere chance, or, in other words, of development. This deduction of linguistic inquiry most fully con- firms the result of natural inquiry. And any one who takes the trouble to follow the course of linguistic science will be convinced that its champions, except, perhaps, Bleek, Schleicher, and Friedrich Miller, are labouring rather to discredit, than to acknowledge, the influence of the doctrine of Descent. All the higher is our estimate of it, and therewith the most powerful objection to the in- clusion of man in the great law of derivation is set aside. 306 THE DOCTRINE OF DESCENT. The rest is incidental and a matter of detail. The question so often ventilated, and now thoroughly worn out, whether mankind is descended from one or more pairs, is solved by the inference that the stock in which language first arose, separated itself gradually from its animal progenitors, and that the selection which led to language and reason necessarily took place among large communities of individuals. The scriptural conception of the unity of the human race would be more nearly approached if all linguistic families pointed to a single source. But if it could be shown that certain linguistic families lead to utterly discordant roots, the investiga- tion of nature might furnish the inevitable corollary that language originated in various parts of the world,— in other words, that a separation into species took place before selection had reached the point of forming lan- guage. The latter case is by far the most probable, and is, in fact, received as the only one possible by most of the linguists occupied with this question, and is most especially defended by Friedrich Miiller.8® “At the time,” he says, “ when there were races and no nations, man was a speechless animal, as yet, entirely destitute of the mental development which rests upon the agency of language. Independently of the premisses unfolded by natural history, this hypothesis is forced upon us by the contemplation of the languages themselves. The various families of languages, which linguistic science is able to discriminate, not only presuppose, by their diver- sity of form and material, several independent origins, but, within one and the same race, they point to several mutually independent points of origin.” In order to afford the reader some notion of the con- 307 nection of the families of nations, we give the subjoined pedigree, in which Friedrich Miiller closely adheres to Haeckel’s sketch. RACES OF MAN. *SUOIJEU JO UISLIO pue asenSur] Jo uoreuLoy ay} Jo Suruulseg aovl uveu “BVPI. “SURIQNN « “eplaviq, ae We ‘20ur -osuoy, ‘sAReAY = ueoNeWYy ‘only ‘suel[Rysny "Sd0139N “$}0} uvolyy ‘suendeg -ua}Jopy “OHBISY ulajseq ‘o1UBIDO “sIye yy ‘aoey SALW | \ / “Ayorea “yar “AJQUIVA - *Kyayea pesey-Ayng parrey-yus paitey-2929],J parrey-ynL Sr Zs Qe ‘sotoads porrey-1y sens ‘saroads pairey-A[[OO MA, UR, DATTIWG “GNINNVYW dO SHOVY AHL AO AAUDIGAG SMaTInW GNV TawMoavH It makes mention of species and races of mankind, the species being regarded as no longer existing, while 308 THE DOCTRINE OF DESCENT, the present forms of man are distinguished only as races. On this subject, we shall not lavish many words, since, examined in the light, it is an affair of words only. In the order of Primates, man constitutes a single family, and represents it by a single genus. Whether Negroes, Caucasians, Papuans, American-Indians, &c., be called species or races, matters little. The facility of intercross- ing the different nations would favour their characteri- zation as races; but as the crossing of species does not differ in principle from the crossing of races, and as to the bodily varieties displayed in colour, hair, skull, limbs, and other characters are added the profound dif- ferences of language, the division of the genus homo into species, diverging into many races, seems after all more natural. But ultimately, as in the question of spe- cies in general, the individual feeling of each person proves decisive. Whether it was a lucky hit to found the division of mankind on the position of the hairs, in tufts or equally distributed upon the scalp, and further- more on the section of the hair, whether it be more flat and oval or circular in form, and finally on the in- clination to curl or to lie stiff and smooth, the future must decide. The twelve races cited in the table given above, may be characterized by the aid of natural history; and as within the limits of the best known races, languages and families of languages may be found, which preclude any common origin, it follows that the formation of language began only after the still speechless primordial man had diverged into races. In geological periods and primordial history, all chronology is extremely decep- tive: we may, nevertheless, acquiesce in an estimate made ANTIQUITY OF MAN, 309 by Friedrich Miiller as to the development of the lan- guages of the Mediterranean races. The linguistic fam- ilies of the nations dwelling chiefly in the basin of the Mediterranean are Basque, Caucasian, Hamito-Semitic and Indo-Germanic languages. “The languages of these four families,” says Friedrich Miller, “are, as is generally accepted by the most competent linguists, not mutually related. If we therefore see that the Mediter- ranean race includes four families of people in no way related to one another, the inference is obvious that, as each language must be. traceable to a society, the single race must have gradually fallen into four societies, of which each independently created its own language. A further inference is, that the race, as such, doe; not acquire a language; for, were this the case, race and language would now be co-extensive, which is not the case.” We must therefore assume that at the time when the various nations of the Mediterranean race were one,— the time when man belonged to no nation, but merely to a race,—mankind was destitute of language. Miiller considers 3000 years approximately sufficient for the period elapsing between the divergence of the race into still speechless societies, and the epoch at which they formed nations, separated and characterized by lan- guages; a period which might seem to many, estimated as far too short. Ifthe ancient civilized people of Egypt be now added on, and the period of its conjectured mi- gration from Asia computed, “the year 6,500 before the commencement of our chronology seems to be the earliest epoch at which we may speak of a Hamito- Semitic primzeval people in the north of Europe.” There- 310 THE DOCTRINE OF DESCENT. fore a Mediterranean race already existed 12,000 years ago. But what space of time was requisite to enable primitive man to separate into races, is entirely beyond computation, and the more so as not the slightest trace of him has hitherto been found. With the invariable testimony of Geology that the periods of the terrestrial strata imperceptibly merged into one another, and that, especially from the Tertiary, through the Diluvial period, to the present age, conti- nuity has been only locally interrupted, the question of the ‘fossil man,” formerly looked upon as cardinal, has assumed another aspect. In Europe, man lived with the mammoth and the rhinoceros with a bony nasal par- tition (Elephas primogenius, Rhinocerus tichorhinus). It has been asserted that European man existed as early as the upper Tertiary age, but the evidence is disputa- ble. Such remains as we have of this oldest man known to us, display a high grade of development, and cer- tainly belong to the period at which man had already found in language the implement wherewith gradually to free himself from the dross of his lowly origin. Whether the primitive man be found or not, his origin is certain. REFERENCES AND QUOTATIONS. ‘ Luthardt, Apologetische Vortrage. 7 Vortrag. P. 129. ? Philosophia quaerit, theologia invenit, religio possidet veri- tatem. ° Tageblatt der Naturforscher-Versammlung in Leipzig, 1872. P. 12. The discourse was also printed separately. “A. Fick, Physiologie, 1860. ® Any one who wishes to be more deeply instructed in the problem of sensation, as an universal primary characteristic of the ‘constituent elements of matter, may be referred to the very lucid and interesting work, “Das Unbewuste vom Standpunkt der Physiologie und Descendenztheorie” (Berlin, 1872). Published anonymously. *L. Geiger, Ueber den Ursprung der Sprache. (Stuttgart, 1869.) P. 207. . 7 Rollet, Ueber Elementartheile und Gewebe und deren Unter- scheidung. Rollet, Untersuchungen, etc. 1871. ® Karl Ernst v. Bar, Ueber Entwickelungsgeschichte der Thiere, Beobachtung und Reflexion, 1828. ° Ib. I. 223. Ib. I. 230, &c. 4 Credner, Elemente der Geologie, 1872. P. 253. 1 Agassiz, Essay on Classification, 1858. ‘It exhibits every- where the working of the same creative Mind, through all times, and upon the whole surface of the globe.” 18 Riitimeyer, Beitrage zur Kenntniss der fossilen Pferde. Verhandlung der naturforschenden Gesellschaft in Basel, 1863. III. 642. 21 311 312 THE DOCTRINE OF DESCENT. “The passages are from an occasional address—Oratio de ellure halvitabili—contained in the Amoenitates academice. “ Initio rerum ex omni specie viventium unicum sexus par fuisse creatum, suadet ratio.” * Ib. Non multum a veritate me aberratum confido, si dixerim, omnen continentem terram fuisse in infantia mundi aquis sub- mersam et vasto oceano obtectam, preter unicam in immenso hoc pelago insulam, in qua commode habitaverint animalia omnia et vegetabilia late germinaverint.” 46 Tot numeramus species, quot ab initio creavit infinitum ens.”’ ™ Geoffroy St. Hilaire wrote to Cuvier: ‘‘ Venez jouer parmi nous le réle de Linné, d’un autre légisiateur de l'histoire naturelle.” *® Ossements fossiles. #” L. Agassiz, An Essay on Classification, 1859. P. 253 :— “As representatives of Speczes, individual animals bear the closest relations to one another; they exhibit definite relations also to the surrounding element, and their existence is limited within a definite period. “As representatives of Genera these same individuals have a definite and specific ultimate structure, identical with that of the representatives of other species,”’ etc. See also P. 261 :— “ Branches or types are characterized by the plan of their struc- ture; “‘ Classes, by the manner in which that plan is executed, as far as ways and means are concerned ; “ Orders, by the degrees of complication of that structure ; “ Families, by their form, as far as determined by structure ; “ Genera, by the details of the execution in special parts; and, “« Spectes, by the relations of individuals to one another,-and to the world in which they live, as well as by the proportions-of their parts, their ornamentation,” etc. 2° Haeckel, Generelle Morphologie der Organismen (Berlin, 1866). IT. 323, &c. 4 L’espéce est—‘‘la réunion des individus descendant l'un de l'autre et des parents communs, et de ceux qui leur ressemblent autant qu’ils se ressemblent entr’eux.” Cuvier, Le Régne Ani- mal. REFERENCES AND QUOTATIONS. 313 * OQ. Schmidt, Die Spongien der Kiiste von Algier, 1868, and Versuch einer Spongienfauna des atlantischen Gebietes, 1870. ** Haeckel, Die Kalkschwamme. Eine Monographie in Zwei Banden, Text und einem Atlas mit 60 Tafeln Abbildungen (Berlin, 1872). *4 Hilgendorf, Ueber Planorbis multiformis in Steinheimer Siiss- wasserkalk. Monatsbericht des Berliner Akademie aus dem Jahre 1866. P. 474, &c. *® Waagen, Die Formenreihe des Ammonites subradiatus. Beneke’s Beitrige, 1869. Vol. 2. Zittel, Die Fauna der dltern Cephalopoden fiihrenden Tithonbil- dungen. Palaontologische Mittheilungen,.1870. Neumayr, Jurastudien, Jahrbuch det geologischen Reichsanstalt, 1871. L. Wiirtenberger, Neuer Beitrag zum geologischen Beweise der Darwin’ schen Theorie, 1873. 6 Darwin, The Variation of Plants and Animals under Domesti- cation, 1868. “7 L. Oken, Die Zeugung, 1805. Lehrbuch der Naturphiloso- phie, 1809-11, Pt. 3. °8 T have borrowed the following account from my essay: ‘“‘ Wae Goethe ein Darwinianer?”” (Was Goethe a Darwinist?) Gratz, Leuschner and Lubinsky, 1871. Also another small work of mine: “ Goethe’s Verhaltniss zu den organischen Naturwissenschaften”’ (Berlin, 1852). To the pas- sages given in the text, which might make Goethe appear as a Darwinist, I may add the following from Eckermann’s ‘‘ Gesprache mit Goethe” (3 Ed. p. 191). ‘Thus man has in his skull ‘two empty cavities. The question why? would not go far, whereas the question how? teaches me that these cavities are remains of the animal skull, which in those inferior organisms exist to a greater degree, and are not entirely lost even in man, notwith- standing his higher elevation.” 29 A somewhat depreciative opinion of Goethe’s importance in this sphere is pronounced by V. Carus in his “Geschichte der Zoologie”” (Miinchen, 1872). The reader, may compare: ‘“ How little, notwithstanding his repeated study of anatomy, he had 314 THE DOCTRINE OF DESCENT. gained a true insight into the structure of animals, as determined by law, is testified by his Introduction to Comparative Anatomy. He finds no other means of harmonizing the dry details of descrip- tive anatomy, and the morphology which vaguely hovered before him, but by indicating the idea of a primitive type for animals, which he is, however, unable to define or to render in any way palpable by more general indications. His whole idiosyncrasy made such a type a necessity to him, not scientifically, but zstheti- cally,” etc. P. 590. 5° R. Owen has declared his attitude towards the doctrine of descent in the concluding chapter of his ‘‘ Manual of the Com- parative Anatomy of the Vertebrata.’ It is published sepa- rately under the title of ‘Derivative Hypothesis of Life and Species,” 1868. SE TDs sca “‘such cause being the servant of predetermining intelligent will,”’ 32 « No one can enter the saddling-ground at Epsom before the start for the Derby, without feeling that the glossy-coated, proudly- stepping creatures led out before him are the most perfect and beautiful of quadrupeds. As such, I believe the horse to have been predestined and prepared for man.”” Ib. P. 11, 83 «T deem an innate tendency to deviate from parental type, operating through periods of adequate duration, to be the most probable nature or way of operation of the secondary law whereby species have been derived one from the other.” Ib. P. 22. 34 Lamarck, Philosophie Zoologique (Paris, 1809). In the text allusion is made to the following passages: « Ainsil’on peut assurer que, parmi ses productions, la nature n’a réellement formé ni classes, ni ordres, ni familles, ni espéces con- stantes, mais seulement des individus qui se succédent les uns aux autres, et qui ressemblent 4 ceux qui les ont produits. Or ces individus appartiennent a des races infiniment diversifiées, qui se nuancent sous toutes les formes et dans tous les degrés d’organisation, et qui chacune se conservent sans mutation tant qu’aucune cause de changement n’agit sur elles.” I. 22. “La supposition presque généralment admise, que les corps vivans constituent des espéces constamment distinctes par des REFERENCES AND QUOTATIONS. 315 caractéres invariables, et que l’existence de ces espéces est aussi ancienne que celle de la nature méme, fut établie dans un temps ot l’on n’avait pas suffisament observé, et ot les sciences naturelles étaient 4 peu prés nulles. Elle est tous les jours démentie aux yeux de ceux qui ont beaucoup vu et qui ont longtemps suivi la nature.” I. 54. “Les espéces n’ont réellement qu’une constance relative a la durée des circonstances dans lesquelles se sont trouvés les indi- vidus qui les représentent.” I. 55. “«—les considérations, et nous font voir— “rt, Que tous les corps organisés de notre globe sont de vérita- bles productions de la nature, qu’elle a successivement executées a la suite de beaucoup de temps. “2, Que dans sa marche la nature a commencé et recommence encore tous les jours, par former les corps organisés les plus simples, et qu’elle ne forme directement que ceux-la, c’est 4 dire que ses premiéres ébauches de l’organisation, qu'on a désignées par l'expression de générations spontanées. «3, Que les premiéres ébauches de l’animal et du vegétal étant formées dans les lieux et les circonstances convenables, les facultés d'une vie commengante et d'un mouvement organique établi ont nécessairement developpé peu a peu les organes, et qu’avec le temps elles les ont diversifiés ainsi qui les parties. “4. Que la faculté d’accroissement dans chaque portion du corps organisé était inhérente aux premiers effets de la vie; elle a donné lieu aux différens modes de la multiplication et de régénérations des individus; et que par la, les progrés acquis dans la composi- tion de Vorganisation et dans la forme et la diversité des parties ont été conservés. “5, Qu’a l’aide d’un temps suffisant, des circonstances qui ont été nécessairement favorables, des changemens que tous les points de la surface du globe ont successivement subis dans leur état, en un mot, du pouvoir qu'ont les nouvelles situations et les nouvelles habitudes pour modifier les organes des corps doués de la vie, tous ceux qui existent maintenant ont été insensiblement formés tels que nous les voyons. “6, Enfin, que d’aprés un ordre semblable de choses, les corps 316 THE DOCTRINE OF DESCENT. vivants ayant éprouvé chacun des changemens plus ou moins grands dans l'état de leur organisation et de leurs parties, ce qu’on nomme espéce parmi eux a été insensiblement et successivement ainsi formé, n’a qu’une constance relative dans son état, et ne peut ainsi étre aussi ancien que la nature.” I. 65, &c. “La progession dans la composition de l’organisation subit. ¢a et la, dans la série générale des animaux, des anomalies opérées par l'influence des circonstances d’habitation et par celle des habitudes contractées.”” I. 135. “Dans tout animal qui n’a point dépassé la terme de ses developpemens, l’emploi plus fréquent et routiné d’un organe quelconque fortifie peu 4 peu cet organe, le developpe, l’agrandit, et lui donne une puissance proportionnée a la durée de cet emploi; tandis quel e défaut constant d’usage de tel organe l'affaiblit insensiblement, le détériore, diminue progressivement ses facultés, et finit par le faire disparaitre. “ Tout ce que la nature a fait acquerir ou perdre aux individus par l’influence des circonstances ot leur race se trouve depuis longtemps exposée, et par conséquent, par l'influence de l'emploi prédominant de tel organe ou par celle d’un défaut constant d'usage de telle partie, elle le conserve par génération aux nou- veaux individus qui en proviennent.” I. 235. “La volonté dépendant toujours d’un jugement quelconque n'est jamais véritablement libre ; car le jugement qui y donne heu est, comme le quotient d’une opération arithmétique, un ré- sultat nécessaire de l'ensemble des éléments qui l’ont formé.” I. 342. “Les animaux contractent, pour satisfaire 4 ces besoins, diverses sortes d’habitudes, qui se transforment en eux en autant de penchans, auxquels ils ne peuvent resister et qu'ils ne peuvent changer eux mémes. De 1a l’origine de leurs actions habituelles et de leurs inclinations particuliéres, auxquelles on a donné le nom d'instinct. Ce penchant des animaux a la conservation des habi- tudes et au renouvellement des actions qui en proviennent, étant une fois acquis, se propage ensuite dans les individus, par la voie de la reproduction ou de la génération, qui conserve ]’organisation et la disposition des parties dans leur état obtenu, en sorte que ce REFERENCES AND QUOTATIONS. 317 méme penchant existe déja dans les nouveaux individus, avant méme quills l’aient exercé.”” I. 325. 8° The acute author of the book, ‘‘Das Unbewusste’’ defines instinct in essentially the same manner as Lamarck. “In this sense it may be said that every instinct is in the last instance by its origin an acquired habit, and the proverb .that ‘habit is second nature’ thus receives the unexpected supplement that habit is also the beginning and origin of the first nature, z. ¢., of instinct. For it is always habit, ¢.¢., the frequent repetition of the same function, which so firmly impresses the mode of action, however acquired, upon the central organs of the nervous system that the predisposition thus originated becomes transmissible,”’ p. 182. %° ‘The highly important doctrine which Lyell has substantiated with his rich experience is also distinctly and concisely enunciated by Lamarck in the Philosophie Zoologique :— “Sil’on considére, d’une part, que dans tout ce que la nature opére elle ne fait rien brusquement, et que partout elle agit avec lenteur et par dégrés successifs, et de l'autre part que les causes particu- liéres ou locales des désordres, des bouleversemens, des déplace- mens, etc., peuvent rendre raison de tout ce que l’on observe a la surface de notre globe, et sont néanmoins assujetties a. ses lois et a sa marche générale, on reconnaftra qu’il n’est nullement nécessaire de supposer qu’une catastrophe universelle est venue culbuter et détruire une grande partie des opérations mémes de la nature.” I. 80. 5" Principles of Geology. 88 In 1870, as well as in 1872, the majority in the French Academy bore this testimony to Darwin. The reiterated proposal of electing him a member was certainly not rejected until such men as Milne-Edwards and Quatrefages .had made the stand- point clear to the scientific judges. ® Origin of Species. Fifth Ed. 1872. The other works cited are, “The Variation of Animals and Plants under Domestication,” 1868; ‘“‘ The Descent of Man and Sexual Selection,”’ 2nd ed., 1871; ‘‘ Expression of the Emotions in Man and Animals,” 1872. *° Malthus (1798) investigated the saaduions of the increase and decrease of human population. He finds that the rise in popula- 318 THE DOCTRINE OF DESCENT. tion is necessarily limited by the means of subsistence, and that the growth increases in proportion to the means of subsistence, setting aside some special impediments easily discovered. These . impediments, which always keep the population below the amount warranted by the means of subsistence, are moral restraint, crime, and misfortune. Malthus depicts the struggle for existence without pronouncing the word; he demonstrates that the dreams of a future blissful equality of all mankind on the, earth transformed into a vast garden, are based upon delusions. Each individual must much rather labour indefatigably to ameliorate his position. By the experience of breeders and gardeners he knows that animals and plants may be improved and ennobled. No organic ennoblement of the human race as a whole is perceptible, nor can the human race be ennobled save by condemning the less perfect individuals to celibacy. These, and similar thoughts in the work of Malthus, first sug~ gested Darwin’s theory, as he has informed us. “ Variation of Animals and Plants. I. 100. “ Two treatises by A. Kerner are also very instructive with regard to the question of species: ‘‘Gute and Schlechte Arten.” (Innsbruck, 1866.) And “ Die Abhangigkeit der Pflanzenwelt von Klima und Boden. Ein Beitrag zur Lehre von der Enstehung und Verbreitung der Arten, gestiitzt auf die Verwandtschaftsverhaltnisse, geographische Verbreitung und Geschichte der Cytisusarten aus dem Stamme Tubocytisus D.C.’ 1869. Kerner’s latest work, ‘Die Schutzmittel des Pollens” (Innsbruck, 1873), is likewise an ad- mirable investigation of the variability, adaptation, and formation of species. . “8 Origin of Species. 13th ed. p. 84. “4 Origin of Species. 13th ed. p. 96. “* Origin of Species. “P.7, The following pages contain an epitome of the objec- tions offered to the inadequacy of the theory of selection. “" Moritz Wagner, Die Darwin’sche Theorie und das Migrations- gesetz der Organismen, 1848. “ Nageli, Enstehung und Begriff der naturhistorischen Art. (Sitzungsberichte der bairischen Akademie der Wissenschaften), REFERENCES AND QUOTATIONS. 319 1865. Nageli’s later investigations (Sitzungsberichte der mathe- matisch-physikalischen Klasse der Miinchner Akademie, 1872, Pp- 305) contirm the doctrine of descent. He shows that the grega~ riousness of merely allied species and their varieties proves more favourable to the formation of species than isolation. ‘The asso- ciated forms—of certain Alpine plants—have, as it were, recipro- cally modified one another; they exhibit, to express myself thus, « specific social type, which zs different in each assemblage, and therefore in every neighbourhood. This fact incontro- vertibly shows that the forms have altered since they were asso- ctated, “The specific social type consists in their showing a notable accordance in certain characteristics, while in others they repre- sent extremes, and in these sometimes exceed all their congeners in other districts. “From these facts it follows undoubtedly that the movement in the cenobitic forms (z. e. living together) is divergent. For extreme characteristics are developed in them, whereas the eremitical forms exhibit a medium in their characteristics. “Nageli proves that since the glacial period an alteration has taken place in Alpine plants, and the manner in which it occurred.” 5 J. Broca, L’Ordre des Primates. Paralléle anatomique de l'homme and des singes, 1870. *\ Descent of Man, p. 367. 52 At the time at which we write, we have before us, unfortunately, only the incomplete reports of the daily papers, and the syllabus of Professor Max Miiller’s “Three Lectures on Mr. Darwin's Philosophy of Language.” ‘ 53 Z6llner, ‘‘ Ueber die Natur der Kometen ” (1 ed. p. 305). 5 For the further instruction of the reader, we will allow another Philosopher and Naturalist to speak respecting the primordial com- mencement of life, to’ our apprehension so simply accountable. The hypothesis of origin is under discussion. In the critical examination of the ‘“ Philosophie des Unbewussten” (7) it runs thus, p. 22. The “ Philosophie des Unbewussten”’ says, p. 558: “It is probable that before the origin of the first organisms, 320 THE DOCTRINE OF DESCENT. organic combinations existed which (p. 556) were under the influ- ence of a damp atmosphere, abounding in carbonic acid, and of a higher temperature, light, and stronger electric influences. If these presuppositions are adopted, and the consideration added that if conditions thus favourable to primordial generation once existed. which they must have done—they probably endured during con- siderable geological periods—the inference is in truth inevitable that in lapse of time and with change of circumstances, these organic substances aggregated into innumerable combinations. Among these innumerable modes of arrangement, groupings and combinations, by far the greater portion must remain at the grade of inorganic form, because it has not attained the needful chemical composition and physical properties ; a very much small- er portion of the results produced by these combinations of or- - ganic materials might perhaps transitorily approach the organic form or even actually assume it, yet without possessing the con- stitution necessary to maintain it permanently; a third and yet smaller portion might perhaps maintain this form for itself in the exchange of mater:al, about as long as the approximate du- ration of life of one of the most primitive of the present Pro- tists, yet lacked those properties which preserve the species by division and reproduction after the natural extinction of the individual; a fourth portion might possess the properties requi- site for self-preservation as well as.for the preservation of the genus, yet lacked that peculiar tendency to vary (Philosophie des Unbewussten, p. 591), or at least that tendency to vary in the particular direction which was alone capable of leading to de- velopment into higher forms; and finally a fifth portion possessed this property in addition to the others. It is the progeny of the fourth and fifth classes of our division which still populates the ocean and the earth.* From which species of Monera_ pro- ceeded the advanced development of the Infusoria ; whether from one still living or from an extinct species we do not know *It is a simpler and more probable explanation that these low organisms continue to exist because there is room for them. They remain in spite of differentiation and in consequence of differentia- tion. REFERENCES AND QUOTATIONS. 321 as yet; but this much we may accept as certain, that the ma- jority of the Protists that we still know, belong to that fourth class which is incapable of development. The persistence of the ephemeral creations of our second and third classes would natu- rally be secured only so long as circumstances continued favour- able to their renewed primordial generation, but from the telev- logical standpoint the first class must be described as that of the completely abortive attempts at creation.” These, and similar more or less interesting fancies to which we attribute no great importance, are all derived from Haeckel’s hypothesis of Autogony (“Generelle Morphologie der Organis- men,” 179 seq.), which he set up after his beautiful discoveries on the simplest organisms now existing—the Monera and the Protists. From this work we select the following passage :—‘‘ Doubtless we must imagine the act of autogony, the first spontaneous origin of the simplest organisms, to be quite similar to the act of crystal- lization. In a fluid, holding in solution the chemical elements composing the organism, in consequence of certain movements of the various elements among themselves, certain points of attrac- tion are formed, at which the atoms of the organogenetic ele- ments (carbon, oxygen, hydrogen, nitrogen) enter into such close contact with one another that they unite in the formation of a complex ternary or quaternary molecule. This primary group of atoms—perhaps a molecule of albumen — now acts like the analogous crystalline molecule, attracting the homogeneous atoms dissolved in the mother wather; and they now likewise coalesce in the formation of similar molecules, The albuminous granule thus grows and transforms itself into a homogeneous organic in- dividual, a structureless moner or mass of plasma, like a Pro- tameba, &c. Owing to the easy divisibility of its substance, this moner constantly tends towards the dissolution of its recenily consolidated individuality, but when the constantly prepondera- ting absorption of new substance outweighs the tendency to dis- integration, it is able to preserve life by the exchange of material. The homogeneous organic individual, or moner, grows by means of imbibition (nutrition) only until the attractive. power of the centre no longer suffices to hold the whole mass together. In 322 THE DOCTRINE OF DESCENT.. consequence of the preponderating divergent movements of the molecules in different directions, two or more centres of attrac- tion are now formed in the homogeneous plasma, which hence- forth act attractively on the individual substance of the simple mould, and thereby induce its fission, or partition, into two or more portions (reproduction). Each part forthwith rounds itself again into an albuminous individual, or mass of plasma, and the eternal process begins again, of attraction and disruption of the molecules, producing the phenomena of exchange of substance, or nutrition, and reproduction.” Relying on the known peculiarities of the combinations of carbon, Haeckel has .attributed to this substance the most im- portant part in his representation of the first development of life and the physiological phenomena of the lowest organisms. This is the “carbon theory ” so strongly deprecated by his antagonists. Minds would be less heated on the subject were it remembered that a refutation of this “adventurous attempt,” as Haeckel terms it, to assist the idea of genesis, would not change a hair in the compulsory logical necessity of acknowledging the evocation of life by natural means. The arguments against the carbon theory have been developed, among others, by Preyer, ‘ Ueber die Er- forschung des Lebens (Jena, 1873). It is shown that carbon, in its present terrestrial conditions, points almost exclusively to organic origin, and, as yet, no source of carbon has been de- monstrated adequate for the first formation of living bodies on the earth. : °° A. R, Wallace, The Malay Archipelago (3rd ed.: London, 1872), and Contributions to the Theory of Natural Selection (2nd ed.: 1871). 56 «The hypothesis of Pangenesis, as applied to the several great classes of facts just discussed, no doubt is extremely com- plex, but so assuredly are the facts. The assumptions, how- ever, on which the hypothesis rests cannot be considered as complex in any extreme degree; namely, that all organic units, besides having the power, as is generally admitted, of growing by self-division, throw off free and minute atoms of their con- tents, that is, gemmules. These multiply, and aggregate them- REFERENCES AND QUOTATIONS. 323 selves into buds and the sexual elements; their development de- pends on their union with other nascent cells, or units, and they are capable of transmission in a dormant state to successive gen- erations. “In a highly organised and complex animal, the gemmules thrown off from each different cell, or unit, throughout the body must be inconceivably numerous and minute. Each unit of each part, as it changes during development —and we know that some insects undergo, at least, twenty metamorphoses— must throw off its gemmules. All organic beings, moreover, include many dormant gemmules derived from their grand-parents and more remote progenitors. These almost infinitely numerous and minute gemmules must be included in each bud, ovule, spermao- zoon, and pollen grain. Such an admission will be declared im- possible, but, as previously remarked, number and size are only relative difficulties, and the eggs or seeds produced by certain animals or plants are so numerous that they cannot be grasped by the intellect.” Darwin, Variations of Animals and Plants, Il. 526. 37 A. Rollet, Ueber die Erscheinungsformen des Lebens und den beharrlichen Zeugen ihres Zusammenhanges. Almanach der kais. Akademie der Wissenschaften (Wien, 1872). ® Darwin, Variations of Animals and Plants, I. 200. 8° VY. Graber, Ueber den Tonapparat der Locustiden, ein Beitrag zum Darwinismus. Zeitschrift fiir wissenschaftliche Zoologie. Vol. 22. ! ® Hermann v. Nathusius, Vorstudien fiir Geschichte und Zucht der Hausthiere zunachst am Schweineschadel, 1864. % Ib., p. 108. 82 Descent of Man, I. 412. * Origin of Species. 13th ed., p. 171. *4 |. amarck also constructed a pedigree at the end of his ‘ Philo- sophie Zoologique,” in which he disposes of the greater number of classes, while he attributes to the remainder another point of derivation. He thus assumes in the animal kingdom two primordial forms derived from primordial generation. His scheme is as follows :— 324 THE DOCTRINE OF DESCENT. TABLEAU Servant & montrer l'origine des differents animaux. Vers, Infusoires, ee eash cn) Polypes. Heleiae tas Mh eee, Radiares, _yhguueeer SE Insectes. res Arachnides, Annelides. Crustacées. Cirripedes. Mollusques. eh Poissons. Reptiles. ‘ Oiseaux. = 5 Mammales amphibiens. Monotrémes. M. cetacées. “ML ongulés. - A comparison of this pedigree with the one which we now set up is extremely interesting, and shows the progress of our knowledge. * Zum Streit iiber den Darwinismus, ‘“‘ Augsburger Allgemeine Zeitung,” 1873, No. 130. ®° A short preliminary communication in the “ Revue Scienti- fique”’ (Paris, 1873). No. 37. *" Braun, Ueber die Bedeutung der Entwickelung in der Naturgeschichte (Berlin, 1872). “The vegetal kingdom shows us— “T. Plants, which in their vegetative development of the germ, Mammales onguiculés. REFERENCES AND QUOTATIONS. 325 exhibit a sexual generation, mostly in a thallus-like form. (Thallo- gens, Bryophytes, the Thallophytes of the authors, and Charas and Mosses.) “II. Plants in which the first generation is transitory, and only the second develops into the vegetative, leaf-forming stem, with- out, however, advancing to the stage of phenogams. (Acrogens Cormophytes, the ferns, &c.) “III, Plants in which metamorphosis advances as far‘as the formation of a blossom, yet without reaching the final formation, that of the formation of the carpel. (Phenogams without real fruit, gymnospermic Anthophytes.) “TV. Plants which reach the final and highest conclusion of vegetable development, that of true fructification. (Angiospermic Anthophytes; Monocotyledons and Dicotyledons as secondary gradations.) °° As we have discussed in this chapter individual development with reterence to historical development, we must also notice the strange opposition to the doctrine of descent offered by KOlliker. He has laid down his views in his ‘‘ Monographie der Penna- tuliden,” and, in a separate pamphlet, bearing the title of “ Mor- phologie und Entwickelungsgeschichte des Pennatulidenstammes, nebst allgemeine Betrachtungen zur Descendenzlehre”’ (Frank- furt, 1872). Whereas Darwinism derives the continuity and harmony of the organic world from variability, natural selection, heredity, and adaptation—in short, from palpable, visibly efficacious causes—Kélliker is of opinion “that the same general formative laws which govern inorganic nature hold good also in the organic kingdom, and hence a common pedigree and a slow transformation of one form into another are entirely unnecessary for the explana- tion and comprehension of the accordance of the forms and series of forms of the animate world” (p. 3). Except decided dualists, no one disputes the first part of Kdlliker’s thesis. But the identifica- tion of the development of the organic individual, excluding the law of heredity, with the simple process of crystallization, or any other operation of chemical combination repeating itself under given conditions, scarcely needs a detailed refutation. Kolliker says, and tries to prove, that the so-called monophyletic hypo- 326 THE DOCTRINE OF DESCENT. thesis, according to which the different families of organisms are derived from a single primordial form, has to struggle with insur- mountable difficulties ; that the hypothesis of descent from many families (polyphyletic) possesses more probability. If this be admitted, then—and here comes a bold leap of the imagination— the adherent of the polyphyletic hypothesis finds himself in a position to attribute different pedigrees and primordial forms not only to the higher divisions, but even to their genera, and to assume their independent origin. Nay, it even seems credible that the self-same species may appear in different pedigrees; as by the incontrovertible supposition of general laws of formation, it cannot be seen why like primary shapes should not, under certain circum- stances, be able to lead to like final forms (see p. 21). Nay, this hypothesis does more, for “‘ even if individuals of the same species occupy remote localities, as, for instance, Pennatula phosphorea, Funiculina quadrangularis, Renilla reniformis, &c., it is surely more fitting to assume their independent origin.” Kolliker’s polyphyletic hypothesis put an end to all difficulties, and, among others, it ex- plains the so-called “‘ representative forms” to be mentioned in our tenth chapter ; for, from ‘this standpoint, it is credible that these forms are not genetically connected, but belong to different pedi- grees’ (p. 23). And all this, and much more, is supposed to be conceivable, because the world of organisms, in its consecutive development, follows intrinsic causes or definite laws of formation, “laws which, in a perfectly definite manner, urge on the organisms to constantly higher development.” At the same time, Kolliker deliberates (p. 38) whether, just as here germs and buds, so also free existing youthful forms of animals did not possess the power of striking out a development different from the typical one, which freedom must be severely mulcted by the law of development, which can and must create individuals of the same species at the opposite poles, Kolliker (p. 44) thus sums up his fundamental view—‘‘that in and with the first origin of organic matter and of organisms, the whole plan of development, the collective series of possibilities, were also potentially given, but that various external impulses operated determinatively on individual developments, and impressed a definite stamp upon them.” Notwithstanding the REFERENCES AND QUOTATIONS. 327 scientific dress, dualism is here complete; whereas, Physics and Chemistry make their laws, applying to inorganic as well as to organic nature, comprehensible in their form, purport, and effects. Kdlliker knows nothing of the constitution of his laws. The doctrine of natural selection allows us to recognize the causes and effects of heredity and adaptation, and establishes the phenomenal series un- der the form of laws. But laws which are founded only on a plan which is to be carried out prospectively and in subservience to this dower of imperfect organisms, are ignored by natural science. °° Ueber die Herkunft unserer Thierwelt. Einezoo geographische Skizze von L. Riitimeyer (Basel, 1867). We have made* copious use in our text of this extremely instructive writing. 7 A, R. Wallace, Malay Archipelago. P. 10, &c. See also his remarkable and comprehensive work on the Geographical Distri- bution of Animals. ™ G. Koch, Die indo-australische Lepidopteren-Fauna in ihren Zusamnenhang mit den drei Hauptfaunen der Erde. (1 Ed., Ber- lin, 1873.) 72 Peschl, Neue Probleme der vergleichende Erdkunde, 1870. ™ Unless we connect the Dinotherium with Mastodon and Ele- phas, Between the pliocene Mastodon Borsoni and the Elephas primigenius, twenty species are interposed, among which are our still living species, the Indian and African elephants. The limits of the two genera are hereby entirely obliterated. According to other statements, the Elephas primigenius (the mammoth) falls into at least four geographical varieties, which join on to the American species. A dwarf species of elephant is found in the caves of Malta, which in dentition attaches itself to the African species. ™ Joh. Schmidt, The Relationships of the Indo-Germanic Lan- guages, 1872. ™ Various antagonists of the doctrine of descent have vented their moral dismay in the most poignant expressions, precluding any scientific discussion, on finding that the pedigree of the Verte- brata, and therewith of man, is actually traced beyond the verte- brated animals to so low a being as the Ascidians. It is otherwise with the critics of Kowalewsky’s and Kupffer’s observations, who acknowledge the facts, but think themselves obliged to differ in 22 328 THE DOCTRINE OF DESCENT. their interpretation. One of these is A. Giard, in his work on the “‘Embryogénie des Ascidiens.” (Archive de Zoologie expérimen- tale, Paris, 1872.) The pupil of Lacaze Duthiers says :—“ La chorde et l’appendice caudale sont chez la larve Ascidienne des organes de locomotion d'un importance assez secondaire malgré leur généralité, pour gu’on les vote disparaitre presque entzérement dans le genre Molgula, ot ils sont devenus inutiles par suite des mceurs de |’ani- mal adulte; l’homologie entre cette chorde dorsale et celle des ver- tébrés n’est donc qu’une homologze d'adaptation déterminée a remplir l’iodentité des fonctions, et n’indique pas de rapports de parente immediate entre les vertébrés et les Ascidiens.”” The author thus denies the consanguinity of the vertebrate animals and Ascid- jians, and traces back to adaptation the resemblance approaching identity occurring in the organs of the two. The inferences in these few sentences appear to us utterly at fault. The circumstance that in Molgula, and many other Testacea, development takes a nar- rower course, makes as little alteration in the importance of the facts as, for instance, the Nauplius development of the Peneus ob- served by Fritz Miiller, or the Navicula of the Molluscs, is prejudiced by the fact that the other Decapods have forfeited the Nauplius phase, or the Landsnails the navicula phase. But it is simply in- comprehensible in what the identity of functions is to consist which in the Vertebrata was capable of producing the notochord, with, it is particularly to be remarked, the spinal cord (which M. Giard entirely forgets) ; and, in the other case, the ‘‘homologie d’adapta- tion.’’ We, on the contrary, see these organs performing different functions, because in the one they remain of fundamental impor- tance through life, and not in the other. Thus we conversely lay the stress on the morphological identity accompanying functional difference. M. Giard adduces no facts. 7°T. H. Huxley, Manual of the Anatomy of the Vertebrated Animals. German Ed. ™ March, American Journal of Sciences and Arts, February, 1873. 7 Eckermann, Gespriche mit Goethe. II. 152. 7 Rousseau, Emile (Euvres, Paris, 1820, IX. 17). ‘‘ Nous n’avons point la mesure de cette machine immense; nous n’en pouvons REFERENCES AND QUOTATIONS. 329 calculer les rapports; nous n’en connaissons ni les premiéres lois, ni la cause finale ; nous nous ignorons nous-mémes ; nous ne con- naissons ni notre nature, ni notre principe actif.’’ ®° Metamorphose der Thiere. * R, Valdek in the “ Presse,” 1865, No. 327. * Huxley, Man’s Place in Nature, 1863. Manual of the Anatomy of the Vertebrated Animals. 8° Broca, L’Ordre des Primates. Paralléle anatomique des l'Homme et des Singes. (Paris, 1870.) 84 Waitz, Anthropologie der Dances 6 thl., p. 796. Bear- beitet von Gerland. 8 Do. p. 708, 8 « Augsburger Allgemeine Zeitung,’’ 1873, Nos. 92-94. Beilage. *' 1 heard the lectures delivered at Strasburg by this scholar, “Ueber die Resultate der Sprachswissenschaft,”’ with great interest and advantage. . * L. Geiger, Der Ursprung der Sprache, 1869, p. 37. 8° Steinthal, Der Ursprung der Sprache, 1851. °° Fr. Miiller, Allgemeine Ethnographie. Wen., 1873. Agassiz, L. 11, 80, 86, 223. Alca impennis, 186. Amblystoma, 210. Ammonites, 71, 74, 97. Amniotes, 250. Amphibians, 209, 258, 264. Amphioxus. See Lancelet. Anchitherium, 242, 273. Ancyloceras, 216. Anguis, 185. Annelids, 32, 198. Annulosa, 250. Anoplotheride, 274. Anteater, 246. : Antelopes, 273, 277. Apes, 280, 288, Arachnida, 33. Archzopteryx, 75, 266. Archzosauros, 73. Archegosauros, 259. Armadillos, 228, 246. Articulata, 32, 53, 72, 201, 219. Ascarides, 206, Ascidians, 35, 219, 252. Axolotl, 209. Baer, C. E. von, 48, 191, 197, 219, 293- Balenide, 280. Barrande, 11. INDEX. Bathybius, 26. Batrachians, 72, 259, 264. Bats, 228, 246, 269. Bavey, 212. Bears, 243. Beaumont, Elie de, 130. Beavers, 247. Bees, 28, 47. Beetles, 185. Belemnites, 74. Beneden, Van, 206. Birds, 227, 232, 250, 265, Bison, 277. Blastoids, 77. Bleek, 304. Bos, 277. Bourguignat, 224. Brachiopoda, 7o. Braun, 193, 220. Brehm, 182, 267. Broca, Prof. 160, 289. Briicke, 27. Bruta, 237, 271. Bubalos, 277. Buffaloes, 244, 247. Buffon, 5. 201, | Butterflies, 227. Cachelots, 279. Cznopithecus, 245. 331 332 Camels, 223. Camper, P. 101, 119, Campsognathus, 264, Canidae, 278. Carnivora, 241, 273, 278. Carpenter, Dr. 64, 93. Cassowaries, 237. Cassidulida, 77. Cats, 100, 170, 234. Cecidomyia, 47. Cephalopoda, 70, 154, 201. Ceratoda, 258. Cercaria, 46. Ceroxylus laceratus, 181. Cestoda, 206, Cetacea, 239, 273, 279. Chalina, 152. Chalinula, 152. Cheiroptera, 280. Chelonia, 263. Chimpanzee, 283. Chirotes, 185. Cidaridez, 77. Cirripedes, 207. Civets, 234. Cladonema Radiatum, 42, 202. Clymenia, 71. Clypeastra, 80. Coccoliths, 26. Ccckleshell, 199. Ceecilia, 259. Coelenterata, 32, 218. Coleoptera, 184. Comatula, 80, 197. Conchifera, 71. Copepoda, 207. Corals, 42. Crabs, 150, 153, 207, 210, 238. Credner, 65. Crinoid, 57, 77- Crocodiles, 75, 260, 262. Crossopterygii, 258. Ctenomys, 184. INDEX.. Curtius, G. 304. Cuscus, 234. Cuttlefish, 33. Cuvier, 30, 85. Cyclostomi, 256. Cytisus, 146. Darwin, 131, 183, 248. Deciduata, 272. Deer, 234, 246, 275. , Delphinoidz, 279. Dentalium Teredo, zoo. Desor, 76. Dicynodonta, 260. Didelphidz, 245. Dinosauria, 261. Dinotherium, 295. Dipnoi, 37, 256, 258. Distoma, 46, 206. Dodo, 186. Dogs, 99, 138. Dubois-Reymond, 15, 20. Duck-mole, 237. Dujardin, 42. Dumeril, A. 209. Earl, G. Windsor, 230. Echidna, 270. Echinidz, 76, 80. Echinoconide, 77. Echinodermata, 32, 76. Edentata, 79, 237, 246, 269, 271. Elasmobranchii, 256. Elephants, 80, 231, 242, 247, 265, 310 Enaliosaurians, 74, 238. Endocyclica, 77. Eozoon, 68. Ephippigera vitium, 172. Eucladia Johnsoni, 76, Feather stars, 76. Fick, A. 18, Fish, 258, 264. INDEX. Foraminifera, 93. Foster, G. 101, Fowls, 135. Frogs, 259. Firbringer, 185. Ganoids, 71, 256, Gargol, 44. Gasteropoda, 70, Gastrula, 51, 218, 221. Gegenbauer, 74, 256. Gerland, 300. Gibbon, 289. Giraffes, 277. Goats, 277. Goethe, 106. Gorilla, 288. Graber, Von, 171. Graminivora, 240, Graptolites, 69. Gratiolet, 291. Guinea-pig, 100. Heckel, 40, 89, 177, 198, 218, 250. Heer, 238. Heliconidz, 179. Helladotherium, 277. Herder, 4. Hilgendorf, 96. Hipparion, 275. Hippopotamus, 243, 276. Holothuria, 78, 197, 219, Honeysuckers, 233. Horses, 81, 170, 225, 242, 273, 295. Humboldt, W. von, 4, 222. Huxley, 257, 264, 289. Hydractinea carnea, 203. Hydra tuba, 178. Hydrophilus piceus, 53. Hyenas, 243, 278. Hylodon Martinicensis, 212. Hypsilophodez, 265. Hyrax, 269, 278. 333 Ichthyornis dispar, 267. Ichthyosauria, 74, 260. Inflata, 216, Insecta, 33. Insectivora, 269, 280. Kangaroos, 233. Kerner, 146. Korte, Dr. 117. Kowalewsky, 199, 219, 251. Labyrinthodonta, 72. Lacertilia, 262. Lama, 223, 245. Lamarck, 117, 124, 147. Lamellibranchiata, 1¢9. Lancelet, 36, 150, 219, 251. Laplace, 15. Leibnitz, 3." Lemurs, 269, 280. Lepidosirens, 37, 238, 258. Leptalidz, 180. Leptotherium, 245. Linnzus, 5, 84, Lions, 223. Lizards, 185, 261. Locke, 303. Lories, 233. Lucca, 286. Luthardt, 12. Lyell, Sir C. 128. Machairodus, 242. Macrauchenide, 275. Madrepores, 42. Mammals, 73, 240, 250, 264, 269, 277. Mammoths, 79, 244, 247. Man, 111, 201, 269, 288. Mantida, 181. Marsh, 267. Marsipobranchii, 256. Marsupial frog, 212, Marsupials, 73, 75, 228, 238, 240, 269. 334 Martens, 241. Mastodon, 80. Maupertuis, 4. Maury, 227. Mayer, Ernst, 118, «+Medusz, 31, 203, 210, 218. Megalonyx Jeffersoni, 246. Mollusca, 33, 199. Monera, 27. Monkeys, 79, 234, 269.—See Apes. Monotremata, 2€9. Miller, Friedrich, 305. Miller, H. 299. Miller, Johannes, 279. Miller, Max, 161, 303. Miller, Fritz, 207, 238. Musk animals, 242. Mylodon Harlemi, 245. Myriapoda, 33. Myxine, 259. Nageli, 160, 193. Nathusius, H. von, 175, 178. Naumayr, 97. Nauplius, 207, 210. Navicula, 199. Nematoids, 205. Oken, 1c5. Ophiura, 198. Opossums, 233, 245. Orang, 288. Orniscelidz, 260. Ornithorhyncus, 37, 237, 270. Orthoptera, 171. Ostrich, 237. Quistitis, 291. Ovibos, 277. Owen, R. 121, 193, 290. Oxen, 247, 275. Oysters, 202, Pachyderms, 79, 240, 243. Palzochceride, 244. INDEX. Paleoniscus, 71. Palzotheride, 273.. Paludina, 200. Pander, 219. Pavians, 289. .Peneus, 208. Petromyzon, 256. Phasmide, 181. Physeteridz, 280. Pigs, 175, 178, 242, 244, 275. Pigeons, 133, 170, 177. Pinnipedea, 278. Placoids, 71. Planorbis multiformis, ¢6. Platelmintha Suctoria, 206, Platypus, 233. Plesiosaurians, 74, 269. Pleuronectidz, 181. Polecats, 241. Polistes Gallica, 47. Polypes, 28, 30, 174, 203, 218. Polypterus, 258. Primates, 308. Proboscide, 273. Protamceba, 26, 40. Proterosaurus, 73. Proteus, 259. Protopterus, 238, 258. Pseudopus, 185. Pterodactyls, 75, 250, 263, 256. Pterotrachea, 199. Pulmo-gasteropoda, 224. Puma, 223. Quadrumana, 245. Radiata, 31. Rathke, 55. Regulara, 77. Reniera, 94, 153. Reptiles, 250, 264. Rhabdoliths, 26. Rhinoceros, 231, 242, 247, 273, 310. INDEX. Rhizopoda, 207. Rodents, 246, 269, 278, 280. Rollet, 26, 168, Rousseau, J. J. 4. Ruminants, 79, 240, 277. Ritimeyer, 81, 223, 227, 236, 257, 273, 277+ Sagitta, 37, 219. Sahuis, 289. Saint Hilaire, E.G. 85. Salamanders, 209, 211, 259. Salmon, 239. Sauria, 260. Sauropsida, 264. Scaphites, 75. Schleicher, 304. Schmidt, Johannes, 249. Schulze, Max, 154. Sea-cows, 277. Sea-cucumbers, 78, 197. Seals, 269. Sea-snails, 199. Sea-urchins, 76, 197. Semper, 42. Serpents, 260. Sheep, 136, 244, 277. Shrimp, 207. Siebold, Von, 47. Sirens, 239, 269, 277. Sivatherium, 277. Sloths, 228, 246. Snails, 224. Snakes, 185. Spatange, 77, 80. Spongiade, 30, 93, 218. Squalodon, 279. Starfish, 76, 197. Stauridium, 42, 202. Stein, 41. Steinthal, 304. Stone-lilies, 76, 80. Strauss, D. F. 302. 335 Sturgeons, 238. Suctoria, 206, Suida, 276. Surinam toad, 212. Siissmilch, 4. Tapeworm, 43, 206. Tapirs, 231, 242, 273. Tedania, 94. Teleostei, 256. Termites, 178. Tesselle, 77. Testacea, 250, 255. Tetrabranchiata, 1¢9. Thompson, W. 64. ‘Threadworms, 200. Thrushes, 234. Tiger, 242. Toenia solium, 44. Tortoises, 75, 238, 260, 263. Tragulidz, 275. Trematoda, 206. Trilobites, 69. Tritons, 209, 260. Trogons, 234. Tuco-tuco, 184. Turbellaria, 37, 45, 206. Turrilites, 75. Unger, F. 72. Ungulates, 240, 242, 259, 273, 278. Ursidez, 278. Verany, 182. Vermes, 32, 201. Vertebrata, 33, 154, 186, 250, 264. Viverrida, 241, 278. Wagner, M. 158, 302. Waitz, 300. Wallace, 164, 230. Walruses, 279. 330 INDEX. Watson, H. C. 151. Worms, 205. , Werner, 129. Wirtenberger, 96, 213. Whales, 269, 280. Wolves, 100. Zeuglodon, 279. ’ Wombats, 233. Zoea, 208. Woodpeckers, 234. Zoliner, 21, 162. THE END. arV1696 Cornell University Ubrary The doc! | | iii iii 924 031 191 004 __olin,anx . 3 1 RN ATI9 yy “<, VES.