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The Descent of Man, by Charles Darwin, 2nd ed. [1874], at sacred-texts.com
Position of man in the animal series--The natural system genealogical-- Adaptive characters of slight value--Various small points of resemblance between man and the Quadrumana--Rank of man in the natural system-- Birthplace and antiquity of man--Absence of fossil connecting links--Lower stages in the genealogy of man, as inferred, firstly from his affinities and secondly from his structure--Early androgynous condition of the Vertebrata--Conclusion.
Even if it be granted that the difference between man and his nearest allies is as great in corporeal structure as some naturalists maintain, and although we must grant that the difference between them is immense in mental power, yet the facts given in the earlier chapters appear to declare, in the plainest manner, that man is descended from some lower form, notwithstanding that connecting-links have not hitherto been discovered.
Man is liable to numerous, slight, and diversified variations, which are induced by the same general causes, are governed and transmitted in accordance with the same general laws, as in the lower animals. Man has multiplied so rapidly, that he has necessarily been exposed to struggle for existence, and consequently to natural selection. He has given rise to many races, some of which differ so much from each other, that they have often been ranked by naturalists as distinct species. His body is constructed on the same homological plan as that of other mammals. He passes through the same phases of embryological development. He retains many rudimentary and useless structures, which no doubt were once serviceable. Characters occasionally make their re-appearance in him, which we have reason to believe were possessed by his early progenitors. If the origin of man had been wholly different from that of all other animals, these various appearances would be mere empty deceptions; but such an admission is incredible. These appearances, on the other hand, are intelligible, at least to a large extent, if man is the co-descendant with other mammals of some unknown and lower form.
Some naturalists, from being deeply impressed with the mental and spiritual powers of man, have divided the whole organic world into three kingdoms, the Human, the Animal, and the Vegetable, thus giving to man a separate kingdom. (1. Isidore Geoffroy St.-Hilaire gives a detailed account of the position assigned to man by various naturalists in their classifications: 'Hist. Nat. Gen.' tom. ii. 1859, pp. 170-189.) Spiritual powers cannot be compared or classed by the naturalist: but he may endeavour to shew, as I have done, that the mental faculties of man and the lower animals do not differ in kind, although immensely in degree. A difference in degree, however great, does not justify us in placing man in a distinct kingdom, as will perhaps be best illustrated by comparing the mental powers of two insects, namely, a coccus or scale-insect and an ant, which undoubtedly belong to the same class. The difference is here greater than, though of a somewhat different kind from, that between man and the highest mammal. The female coccus, whilst young, attaches itself by its proboscis to a plant; sucks the sap, but never moves again; is fertilised and lays eggs; and this is its whole history. On the other hand, to describe the habits and mental powers of worker-ants, would require, as Pierre Huber has shewn, a large volume; I may, however, briefly specify a few points. Ants certainly communicate information to each other, and several unite for the same work, or for games of play. They recognise their fellow-ants after months of absence, and feel sympathy for each other. They build great edifices, keep them clean, close the doors in the evening, and post sentries. They make roads as well as tunnels under rivers, and temporary bridges over them, by clinging together. They collect food for the community, and when an object, too large for entrance, is brought to the nest, they enlarge the door, and afterwards build it up again. They store up seeds, of which they prevent the germination, and which, if damp, are brought up to the surface to dry. They keep aphides and other insects as milch-cows. They go out to battle in regular bands, and freely sacrifice their lives for the common weal. They emigrate according to a preconcerted plan. They capture slaves. They move the eggs of their aphides, as well as their own eggs and cocoons, into warm parts of the nest, in order that they may be quickly hatched; and endless similar facts could be given. (2. Some of the most interesting facts ever published on the habits of ants are given by Mr. Belt, in his 'Naturalist in Nicaragua,' 1874. See also Mr. Moggridge's admirable work, 'Harvesting Ants,' etc., 1873, also 'L'Instinct chez les Insectes,' by M. George Pouchet, 'Revue des Deux Mondes,' Feb. 1870, p. 682.) On the whole, the difference in mental power between an ant and a coccus is immense; yet no one has ever dreamed of placing these insects in distinct classes, much less in distinct kingdoms. No doubt the difference is bridged over by other insects; and this is not the case with man and the higher apes. But we have every reason to believe that the breaks in the series are simply the results of many forms having become extinct.
Professor Owen, relying chiefly on the structure of the brain, has divided the mammalian series into four sub-classes. One of these he devotes to man; in another he places both the marsupials and the Monotremata; so that he makes man as distinct from all other mammals as are these two latter groups conjoined. This view has not been accepted, as far as I am aware, by any naturalist capable of forming an independent judgment, and therefore need not here be further considered.
We can understand why a classification founded on any single character or organ--even an organ so wonderfully complex and important as the brain--or on the high development of the mental faculties, is almost sure to prove unsatisfactory. This principle has indeed been tried with hymenopterous insects; but when thus classed by their habits or instincts, the arrangement proved thoroughly artificial. (3. Westwood, 'Modern Classification of Insects,' vol. ii. 1840, p. 87.) Classifications may, of course, be based on any character whatever, as on size, colour, or the element inhabited; but naturalists have long felt a profound conviction that there is a natural system. This system, it is now generally admitted, must be, as far as possible, genealogical in arrangement,--that is, the co- descendants of the same form must be kept together in one group, apart from the co-descendants of any other form; but if the parent-forms are related, so will be their descendants, and the two groups together will form a larger group. The amount of difference between the several groups--that is the amount of modification which each has undergone--is expressed by such terms as genera, families, orders, and classes. As we have no record of the lines of descent, the pedigree can be discovered only by observing the degrees of resemblance between the beings which are to be classed. For this object numerous points of resemblance are of much more importance than the amount of similarity or dissimilarity in a few points. If two languages were found to resemble each other in a multitude of words and points of construction, they would be universally recognised as having sprung from a common source, notwithstanding that they differed greatly in some few words or points of construction. But with organic beings the points of resemblance must not consist of adaptations to similar habits of life: two animals may, for instance, have had their whole frames modified for living in the water, and yet they will not be brought any nearer to each other in the natural system. Hence we can see how it is that resemblances in several unimportant structures, in useless and rudimentary organs, or not now functionally active, or in an embryological condition, are by far the most serviceable for classification; for they can hardly be due to adaptations within a late period; and thus they reveal the old lines of descent or of true affinity.
We can further see why a great amount of modification in some one character ought not to lead us to separate widely any two organisms. A part which already differs much from the same part in other allied forms has already, according to the theory of evolution, varied much; consequently it would (as long as the organism remained exposed to the same exciting conditions) be liable to further variations of the same kind; and these, if beneficial, would be preserved, and thus be continually augmented. In many cases the continued development of a part, for instance, of the beak of a bird, or of the teeth of a mammal, would not aid the species in gaining its food, or for any other object; but with man we can see no definite limit to the continued development of the brain and mental faculties, as far as advantage is concerned. Therefore in determining the position of man in the natural or genealogical system, the extreme development of his brain ought not to outweigh a multitude of resemblances in other less important or quite unimportant points.
The greater number of naturalists who have taken into consideration the whole structure of man, including his mental faculties, have followed Blumenbach and Cuvier, and have placed man in a separate Order, under the title of the Bimana, and therefore on an equality with the orders of the Quadrumana, Carnivora, etc. Recently many of our best naturalists have recurred to the view first propounded by Linnaeus, so remarkable for his sagacity, and have placed man in the same Order with the Quadrumana, under the title of the Primates. The justice of this conclusion will be admitted: for in the first place, we must bear in mind the comparative insignificance for classification of the great development of the brain in man, and that the strongly-marked differences between the skulls of man and the Quadrumana (lately insisted upon by Bischoff, Aeby, and others) apparently follow from their differently developed brains. In the second place, we must remember that nearly all the other and more important differences between man and the Quadrumana are manifestly adaptive in their nature, and relate chiefly to the erect position of man; such as the structure of his hand, foot, and pelvis, the curvature of his spine, and the position of his head. The family of Seals offers a good illustration of the small importance of adaptive characters for classification. These animals differ from all other Carnivora in the form of their bodies and in the structure of their limbs, far more than does man from the higher apes; yet in most systems, from that of Cuvier to the most recent one by Mr. Flower (4. 'Proceedings Zoological Society,' 1863, p. 4.), seals are ranked as a mere family in the Order of the Carnivora. If man had not been his own classifier, he would never have thought of founding a separate order for his own reception.
It would be beyond my limits, and quite beyond my knowledge, even to name the innumerable points of structure in which man agrees with the other Primates. Our great anatomist and philosopher, Prof. Huxley, has fully discussed this subject (5. 'Evidence as to Man's Place in Nature,' 1863, p. 70, et passim.), and concludes that man in all parts of his organization differs less from the higher apes, than these do from the lower members of the same group. Consequently there "is no justification for placing man in a distinct order."
In an early part of this work I brought forward various facts, shewing how closely man agrees in constitution with the higher mammals; and this agreement must depend on our close similarity in minute structure and chemical composition. I gave, as instances, our liability to the same diseases, and to the attacks of allied parasites; our tastes in common for the same stimulants, and the similar effects produced by them, as well as by various drugs, and other such facts.
As small unimportant points of resemblance between man and the Quadrumana are not commonly noticed in systematic works, and as, when numerous, they clearly reveal our relationship, I will specify a few such points. The relative position of our features is manifestly the same; and the various emotions are displayed by nearly similar movements of the muscles and skin, chiefly above the eyebrows and round the mouth. Some few expressions are, indeed, almost the same, as in the weeping of certain kinds of monkeys and in the laughing noise made by others, during which the corners of the mouth are drawn backwards, and the lower eyelids wrinkled. The external ears are curiously alike. In man the nose is much more prominent than in most monkeys; but we may trace the commencement of an aquiline curvature in the nose of the Hoolock Gibbon; and this in the Semnopithecus nasica is carried to a ridiculous extreme.
The faces of many monkeys are ornamented with beards, whiskers, or moustaches. The hair on the head grows to a great length in some species of Semnopithecus (6. Isidore Geoffroy St.-Hilaire, 'Hist. Nat. Gen.' tom. ii. 1859, p. 217.); and in the Bonnet monkey (Macacus radiatus) it radiates from a point on the crown, with a parting down the middle. It is commonly said that the forehead gives to man his noble and intellectual appearance; but the thick hair on the head of the Bonnet monkey terminates downwards abruptly, and is succeeded by hair so short and fine that at a little distance the forehead, with the exception of the eyebrows, appears quite naked. It has been erroneously asserted that eyebrows are not present in any monkey. In the species just named the degree of nakedness of the forehead differs in different individuals; and Eschricht states (7. 'Uber die Richtung der Haare,' etc., Muller's 'Archiv fur Anat. und Phys.' 1837, s. 51.) that in our children the limit between the hairy scalp and the naked forehead is sometimes not well defined; so that here we seem to have a trifling case of reversion to a progenitor, in whom the forehead had not as yet become quite naked.
It is well known that the hair on our arms tends to converge from above and below to a point at the elbow. This curious arrangement, so unlike that in most of the lower mammals, is common to the gorilla, chimpanzee, orang, some species of Hylobates, and even to some few American monkeys. But in Hylobates agilis the hair on the fore-arm is directed downwards or towards the wrist in the ordinary manner; and in H. lar it is nearly erect, with only a very slight forward inclination; so that in this latter species it is in a transitional state. It can hardly be doubted that with most mammals the thickness of the hair on the back and its direction, is adapted to throw off the rain; even the transverse hairs on the fore-legs of a dog may serve for this end when he is coiled up asleep. Mr. Wallace, who has carefully studied the habits of the orang, remarks that the convergence of the hair towards the elbow on the arms of the orang may be explained as serving to throw off the rain, for this animal during rainy weather sits with its arms bent, and with the hands clasped round a branch or over its head. According to Livingstone, the gorilla also "sits in pelting rain with his hands over his head." (8. Quoted by Reade, 'The African Sketch Book,' vol i. 1873, p. 152.) If the above explanation is correct, as seems probable, the direction of the hair on our own arms offers a curious record of our former state; for no one supposes that it is now of any use in throwing off the rain; nor, in our present erect condition, is it properly directed for this purpose.
It would, however, be rash to trust too much to the principle of adaptation in regard to the direction of the hair in man or his early progenitors; for it is impossible to study the figures given by Eschricht of the arrangement of the hair on the human foetus (this being the same as in the adult) and not agree with this excellent observer that other and more complex causes have intervened. The points of convergence seem to stand in some relation to those points in the embryo which are last closed in during development. There appears, also, to exist some relation between the arrangement of the hair on the limbs, and the course of the medullary arteries. (9. On the hair in Hylobates, see 'Natural History of Mammals,' by C.L. Martin, 1841, p. 415. Also, Isidore Geoffroy on the American monkeys and other kinds, 'Hist. Nat. Gen.' vol. ii. 1859, pp. 216, 243. Eschricht, ibid. s. 46, 55, 61. Owen, 'Anatomy of Vertebrates,' vol. iii. p. 619. Wallace, 'Contributions to the Theory of Natural Selection,' 1870, p. 344.)
It must not be supposed that the resemblances between man and certain apes in the above and in many other points--such as in having a naked forehead, long tresses on the head, etc.,--are all necessarily the result of unbroken inheritance from a common progenitor, or of subsequent reversion. Many of these resemblances are more probably due to analogous variation, which follows, as I have elsewhere attempted to shew (10. 'Origin of Species,' 5th edit. 1869, p.194. 'The Variation of Animals and Plants under Domestication,' vol. ii. 1868, p. 348.), from co-descended organisms having a similar constitution, and having been acted on by like causes inducing similar modifications. With respect to the similar direction of the hair on the fore-arms of man and certain monkeys, as this character is common to almost all the anthropomorphous apes, it may probably be attributed to inheritance; but this is not certain, as some very distinct American monkeys are thus characterised.
Although, as we have now seen, man has no just right to form a separate Order for his own reception, he may perhaps claim a distinct Sub-order or Family. Prof. Huxley, in his last work (11. 'An Introduction to the Classification of Animals,' 1869, p. 99.), divides the primates into three Sub-orders; namely, the Anthropidae with man alone, the Simiadae including monkeys of all kinds, and the Lemuridae with the diversified genera of lemurs. As far as differences in certain important points of structure are concerned, man may no doubt rightly claim the rank of a Sub-order; and this rank is too low, if we look chiefly to his mental faculties. Nevertheless, from a genealogical point of view it appears that this rank is too high, and that man ought to form merely a Family, or possibly even only a Sub- family. If we imagine three lines of descent proceeding from a common stock, it is quite conceivable that two of them might after the lapse of ages be so slightly changed as still to remain as species of the same genus, whilst the third line might become so greatly modified as to deserve to rank as a distinct Sub-family, Family, or even Order. But in this case it is almost certain that the third line would still retain through inheritance numerous small points of resemblance with the other two. Here, then, would occur the difficulty, at present insoluble, how much weight we ought to assign in our classifications to strongly-marked differences in some few points,--that is, to the amount of modification undergone; and how much to close resemblance in numerous unimportant points, as indicating the lines of descent or genealogy. To attach much weight to the few but strong differences is the most obvious and perhaps the safest course, though it appears more correct to pay great attention to the many small resemblances, as giving a truly natural classification.
In forming a judgment on this head with reference to man, we must glance at the classification of the Simiadae. This family is divided by almost all naturalists into the Catarrhine group, or Old World monkeys, all of which are characterised (as their name expresses) by the peculiar structure of their nostrils, and by having four premolars in each jaw; and into the Platyrrhine group or New World monkeys (including two very distinct sub- groups), all of which are characterised by differently constructed nostrils, and by having six premolars in each jaw. Some other small differences might be mentioned. Now man unquestionably belongs in his dentition, in the structure of his nostrils, and some other respects, to the Catarrhine or Old World division; nor does he resemble the Platyrrhines more closely than the Catarrhines in any characters, excepting in a few of not much importance and apparently of an adaptive nature. It is therefore against all probability that some New World species should have formerly varied and produced a man-like creature, with all the distinctive characters proper to the Old World division; losing at the same time all its own distinctive characters. There can, consequently, hardly be a doubt that man is an off-shoot from the Old World Simian stem; and that under a genealogical point of view he must be classed with the Catarrhine division. (12. This is nearly the same classification as that provisionally adopted by Mr. St. George Mivart, ('Transactions, Philosophical Society," 1867, p. 300), who, after separating the Lemuridae, divides the remainder of the Primates into the Hominidae, the Simiadae which answer to the Catarrhines, the Cebidae, and the Hapalidae,--these two latter groups answering to the Platyrrhines. Mr. Mivart still abides by the same view; see 'Nature,' 1871, p. 481.)
The anthropomorphous apes, namely the gorilla, chimpanzee, orang, and hylobates, are by most naturalists separated from the other Old World monkeys, as a distinct sub-group. I am aware that Gratiolet, relying on the structure of the brain, does not admit the existence of this sub-group, and no doubt it is a broken one. Thus the orang, as Mr. St. G. Mivart remarks, "is one of the most peculiar and aberrant forms to be found in the Order." (13. 'Transactions, Zoolog. Soc.' vol. vi. 1867, p. 214.) The remaining non-anthropomorphous Old World monkeys, are again divided by some naturalists into two or three smaller sub-groups; the genus Semnopithecus, with its peculiar sacculated stomach, being the type of one sub-group. But it appears from M. Gaudry's wonderful discoveries in Attica, that during the Miocene period a form existed there, which connected Semnopithecus and Macacus; and this probably illustrates the manner in which the other and higher groups were once blended together.
If the anthropomorphous apes be admitted to form a natural sub-group, then as man agrees with them, not only in all those characters which he possesses in common with the whole Catarrhine group, but in other peculiar characters, such as the absence of a tail and of callosities, and in general appearance, we may infer that some ancient member of the anthropomorphous sub-group gave birth to man. It is not probable that, through the law of analogous variation, a member of one of the other lower sub-groups should have given rise to a man-like creature, resembling the higher anthropomorphous apes in so many respects. No doubt man, in comparison with most of his allies, has undergone an extraordinary amount of modification, chiefly in consequence of the great development of his brain and his erect position; nevertheless, we should bear in mind that he "is but one of several exceptional forms of Primates." (14. Mr. St. G. Mivart, 'Transactions of the Philosophical Society,' 1867, p. 410.)
Every naturalist, who believes in the principle of evolution, will grant that the two main divisions of the Simiadae, namely the Catarrhine and Platyrrhine monkeys, with their sub-groups, have all proceeded from some one extremely ancient progenitor. The early descendants of this progenitor, before they had diverged to any considerable extent from each other, would still have formed a single natural group; but some of the species or incipient genera would have already begun to indicate by their diverging characters the future distinctive marks of the Catarrhine and Platyrrhine divisions. Hence the members of this supposed ancient group would not have been so uniform in their dentition, or in the structure of their nostrils, as are the existing Catarrhine monkeys in one way and the Platyrrhines in another way, but would have resembled in this respect the allied Lemuridae, which differ greatly from each other in the form of their muzzles (15. Messrs. Murie and Mivart on the Lemuroidea, 'Transactions, Zoological Society,' vol. vii, 1869, p. 5.), and to an extraordinary degree in their dentition.
The Catarrhine and Platyrrhine monkeys agree in a multitude of characters, as is shewn by their unquestionably belonging to one and the same Order. The many characters which they possess in common can hardly have been independently acquired by so many distinct species; so that these characters must have been inherited. But a naturalist would undoubtedly have ranked as an ape or a monkey, an ancient form which possessed many characters common to the Catarrhine and Platyrrhine monkeys, other characters in an intermediate condition, and some few, perhaps, distinct from those now found in either group. And as man from a genealogical point of view belongs to the Catarrhine or Old World stock, we must conclude, however much the conclusion may revolt our pride, that our early progenitors would have been properly thus designated. (16. Haeckel has come to this same conclusion. See 'Uber die Entstehung des Menschengeschlechts,' in Virchow's 'Sammlung. gemein. wissen. Vortrage,' 1868, s. 61. Also his 'Naturliche Schopfungsgeschicte,' 1868, in which he gives in detail his views on the genealogy of man.) But we must not fall into the error of supposing that the early progenitor of the whole Simian stock, including man, was identical with, or even closely resembled, any existing ape or monkey.
ON THE BIRTHPLACE AND ANTIQUITY OF MAN.
We are naturally led to enquire, where was the birthplace of man at that stage of descent when our progenitors diverged from the Catarrhine stock? The fact that they belonged to this stock clearly shews that they inhabited the Old World; but not Australia nor any oceanic island, as we may infer from the laws of geographical distribution. In each great region of the world the living mammals are closely related to the extinct species of the same region. It is therefore probable that Africa was formerly inhabited by extinct apes closely allied to the gorilla and chimpanzee; and as these two species are now man's nearest allies, it is somewhat more probable that our early progenitors lived on the African continent than elsewhere. But it is useless to speculate on this subject; for two or three anthropomorphous apes, one the Dryopithecus (17. Dr. C. Forsyth Major, 'Sur les Singes fossiles trouves en Italie:' 'Soc. Ital. des Sc. Nat.' tom. xv. 1872.) of Lartet, nearly as large as a man, and closely allied to Hylobates, existed in Europe during the Miocene age; and since so remote a period the earth has certainly undergone many great revolutions, and there has been ample time for migration on the largest scale.
At the period and place, whenever and wherever it was, when man first lost his hairy covering, he probably inhabited a hot country; a circumstance favourable for the frugiferous diet on which, judging from analogy, he subsisted. We are far from knowing how long ago it was when man first diverged from the Catarrhine stock; but it may have occurred at an epoch as remote as the Eocene period; for that the higher apes had diverged from the lower apes as early as the Upper Miocene period is shewn by the existence of the Dryopithecus. We are also quite ignorant at how rapid a rate organisms, whether high or low in the scale, may be modified under favourable circumstances; we know, however, that some have retained the same form during an enormous lapse of time. From what we see going on under domestication, we learn that some of the co-descendants of the same species may be not at all, some a little, and some greatly changed, all within the same period. Thus it may have been with man, who has undergone a great amount of modification in certain characters in comparison with the higher apes.
The great break in the organic chain between man and his nearest allies, which cannot be bridged over by any extinct or living species, has often been advanced as a grave objection to the belief that man is descended from some lower form; but this objection will not appear of much weight to those who, from general reasons, believe in the general principle of evolution. Breaks often occur in all parts of the series, some being wide, sharp and defined, others less so in various degrees; as between the orang and its nearest allies--between the Tarsius and the other Lemuridae--between the elephant, and in a more striking manner between the Ornithorhynchus or Echidna, and all other mammals. But these breaks depend merely on the number of related forms which have become extinct. At some future period, not very distant as measured by centuries, the civilised races of man will almost certainly exterminate, and replace, the savage races throughout the world. At the same time the anthropomorphous apes, as Professor Schaaffhausen has remarked (18. 'Anthropological Review,' April 1867, p. 236.), will no doubt be exterminated. The break between man and his nearest allies will then be wider, for it will intervene between man in a more civilised state, as we may hope, even than the Caucasian, and some ape as low as a baboon, instead of as now between the negro or Australian and the gorilla.
With respect to the absence of fossil remains, serving to connect man with his ape-like progenitors, no one will lay much stress on this fact who reads Sir C. Lyell's discussion (19. 'Elements of Geology,' 1865, pp. 583- 585. 'Antiquity of Man,' 1863, p. 145.), where he shews that in all the vertebrate classes the discovery of fossil remains has been a very slow and fortuitous process. Nor should it be forgotten that those regions which are the most likely to afford remains connecting man with some extinct ape- like creature, have not as yet been searched by geologists.
LOWER STAGES IN THE GENEALOGY OF MAN.
We have seen that man appears to have diverged from the Catarrhine or Old World division of the Simiadae, after these had diverged from the New World division. We will now endeavour to follow the remote traces of his genealogy, trusting principally to the mutual affinities between the various classes and orders, with some slight reference to the periods, as far as ascertained, of their successive appearance on the earth. The Lemuridae stand below and near to the Simiadae, and constitute a very distinct family of the primates, or, according to Haeckel and others, a distinct Order. This group is diversified and broken to an extraordinary degree, and includes many aberrant forms. It has, therefore, probably suffered much extinction. Most of the remnants survive on islands, such as Madagascar and the Malayan archipelago, where they have not been exposed to so severe a competition as they would have been on well-stocked continents. This group likewise presents many gradations, leading, as Huxley remarks (20. 'Man's Place in Nature,' p. 105.), "insensibly from the crown and summit of the animal creation down to creatures from which there is but a step, as it seems, to the lowest, smallest, and least intelligent of the placental mammalia." From these various considerations it is probable that the Simiadae were originally developed from the progenitors of the existing Lemuridae; and these in their turn from forms standing very low in the mammalian series.
The Marsupials stand in many important characters below the placental mammals. They appeared at an earlier geological period, and their range was formerly much more extensive than at present. Hence the Placentata are generally supposed to have been derived from the Implacentata or Marsupials; not, however, from forms closely resembling the existing Marsupials, but from their early progenitors. The Monotremata are plainly allied to the Marsupials, forming a third and still lower division in the great mammalian series. They are represented at the present day solely by the Ornithorhynchus and Echidna; and these two forms may be safely considered as relics of a much larger group, representatives of which have been preserved in Australia through some favourable concurrence of circumstances. The Monotremata are eminently interesting, as leading in several important points of structure towards the class of reptiles.
In attempting to trace the genealogy of the Mammalia, and therefore of man, lower down in the series, we become involved in greater and greater obscurity; but as a most capable judge, Mr. Parker, has remarked, we have good reason to believe, that no true bird or reptile intervenes in the direct line of descent. He who wishes to see what ingenuity and knowledge can effect, may consult Prof. Haeckel's works. (21. Elaborate tables are given in his 'Generelle Morphologie' (B. ii. s. cliii. and s. 425); and with more especial reference to man in his 'Naturliche Schopfungsgeschichte,' 1868. Prof. Huxley, in reviewing this latter work ('The Academy,' 1869, p. 42) says, that he considers the phylum or lines of descent of the Vertebrata to be admirably discussed by Haeckel, although he differs on some points. He expresses, also, his high estimate of the general tenor and spirit of the whole work.) I will content myself with a few general remarks. Every evolutionist will admit that the five great vertebrate classes, namely, mammals, birds, reptiles, amphibians, and fishes, are descended from some one prototype; for they have much in common, especially during their embryonic state. As the class of fishes is the most lowly organised, and appeared before the others, we may conclude that all the members of the vertebrate kingdom are derived from some fishlike animal. The belief that animals so distinct as a monkey, an elephant, a humming-bird, a snake, a frog, and a fish, etc., could all have sprung from the same parents, will appear monstrous to those who have not attended to the recent progress of natural history. For this belief implies the former existence of links binding closely together all these forms, now so utterly unlike.
Nevertheless, it is certain that groups of animals have existed, or do now exist, which serve to connect several of the great vertebrate classes more or less closely. We have seen that the Ornithorhynchus graduates towards reptiles; and Prof. Huxley has discovered, and is confirmed by Mr. Cope and others, that the Dinosaurians are in many important characters intermediate between certain reptiles and certain birds--the birds referred to being the ostrich-tribe (itself evidently a widely-diffused remnant of a larger group) and the Archeopteryx, that strange Secondary bird, with a long lizard-like tail. Again, according to Prof. Owen (22. 'Palaeontology' 1860, p. 199.), the Ichthyosaurians--great sea-lizards furnished with paddles--present many affinities with fishes, or rather, according to Huxley, with amphibians; a class which, including in its highest division frogs and toads, is plainly allied to the Ganoid fishes. These latter fishes swarmed during the earlier geological periods, and were constructed on what is called a generalised type, that is, they presented diversified affinities with other groups of organisms. The Lepidosiren is also so closely allied to amphibians and fishes, that naturalists long disputed in which of these two classes to rank it; it, and also some few Ganoid fishes, have been preserved from utter extinction by inhabiting rivers, which are harbours of refuge, and are related to the great waters of the ocean in the same way that islands are to continents.
Lastly, one single member of the immense and diversified class of fishes, namely, the lancelet or amphioxus, is so different from all other fishes, that Haeckel maintains that it ought to form a distinct class in the vertebrate kingdom. This fish is remarkable for its negative characters; it can hardly be said to possess a brain, vertebral column, or heart, etc.; so that it was classed by the older naturalists amongst the worms. Many years ago Prof. Goodsir perceived that the lancelet presented some affinities with the Ascidians, which are invertebrate, hermaphrodite, marine creatures permanently attached to a support. They hardly appear like animals, and consist of a simple, tough, leathery sack, with two small projecting orifices. They belong to the Mulluscoida of Huxley--a lower division of the great kingdom of the Mollusca; but they have recently been placed by some naturalists amongst the Vermes or worms. Their larvae somewhat resemble tadpoles in shape (23. At the Falkland Islands I had the satisfaction of seeing, in April, 1833, and therefore some years before any other naturalist, the locomotive larvae of a compound Ascidian, closely allied to Synoicum, but apparently generically distinct from it. The tail was about five times as long as the oblong head, and terminated in a very fine filament. It was, as sketched by me under a simple microscope, plainly divided by transverse opaque partitions, which I presume represent the great cells figured by Kovalevsky. At an early stage of development the tail was closely coiled round the head of the larva.), and have the power of swimming freely about. Mr. Kovalevsky (24. 'Memoires de l'Acad. des Sciences de St. Petersbourg,' tom. x. No. 15, 1866.) has lately observed that the larvae of Ascidians are related to the Vertebrata, in their manner of development, in the relative position of the nervous system, and in possessing a structure closely like the chorda dorsalis of vertebrate animals; and in this he has been since confirmed by Prof. Kupffer. M. Kovalevsky writes to me from Naples, that he has now carried these observations yet further, and should his results be well established, the whole will form a discovery of the very greatest value. Thus, if we may rely on embryology, ever the safest guide in classification, it seems that we have at last gained a clue to the source whence the Vertebrata were derived. (25. But I am bound to add that some competent judges dispute this conclusion; for instance, M. Giard, in a series of papers in the 'Archives de Zoologie Experimentale,' for 1872. Nevertheless, this naturalist remarks, p. 281, "L'organisation de la larve ascidienne en dehors de toute hypothese et de toute theorie, nous montre comment la nature peut produire la disposition fondamentale du type vertebre (l'existence d'une corde dorsale) chez un invertebre par la seule condition vitale de l'adaptation, et cette simple possibilite du passage supprime l'abime entre les deux sous-regnes, encore bien qu'en ignore par ou le passage s'est fait en realite.") We should then be justified in believing that at an extremely remote period a group of animals existed, resembling in many respects the larvae of our present Ascidians, which diverged into two great branches--the one retrograding in development and producing the present class of Ascidians, the other rising to the crown and summit of the animal kingdom by giving birth to the Vertebrata.
We have thus far endeavoured rudely to trace the genealogy of the Vertebrata by the aid of their mutual affinities. We will now look to man as he exists; and we shall, I think, be able partially to restore the structure of our early progenitors, during successive periods, but not in due order of time. This, can be effected by means of the rudiments which man still retains, by the characters which occasionally make their appearance in him through reversion, and by the aid of the principles of morphology and embryology. The various facts, to which I shall here allude, have been given in the previous chapters.
The early progenitors of man must have been once covered with hair, both sexes having beards; their ears were probably pointed, and capable of movement; and their bodies were provided with a tail, having the proper muscles. Their limbs and bodies were also acted on by many muscles which now only occasionally reappear, but are normally present in the Quadrumana. At this or some earlier period, the great artery and nerve of the humerus ran through a supra-condyloid foramen. The intestine gave forth a much larger diverticulum or caecum than that now existing. The foot was then prehensile, judging from the condition of the great toe in the foetus; and our progenitors, no doubt, were arboreal in their habits, and frequented some warm, forest-clad land. The males had great canine teeth, which served them as formidable weapons. At a much earlier period the uterus was double; the excreta were voided through a cloaca; and the eye was protected by a third eyelid or nictitating membrane. At a still earlier period the progenitors of man must have been aquatic in their habits; for morphology plainly tells us that our lungs consist of a modified swim-bladder, which once served as a float. The clefts on the neck in the embryo of man shew where the branchiae once existed. In the lunar or weekly recurrent periods of some of our functions we apparently still retain traces of our primordial birthplace, a shore washed by the tides. At about this same early period the true kidneys were replaced by the corpora wolffiana. The heart existed as a simple pulsating vessel; and the chorda dorsalis took the place of a vertebral column. These early ancestors of man, thus seen in the dim recesses of time, must have been as simply, or even still more simply organised than the lancelet or amphioxus.
There is one other point deserving a fuller notice. It has long been known that in the vertebrate kingdom one sex bears rudiments of various accessory parts, appertaining to the reproductive system, which properly belong to the opposite sex; and it has now been ascertained that at a very early embryonic period both sexes possess true male and female glands. Hence some remote progenitor of the whole vertebrate kingdom appears to have been hermaphrodite or androgynous. (26. This is the conclusion of Prof. Gegenbaur, one of the highest authorities in comparative anatomy: see 'Grundzuge der vergleich. Anat.' 1870, s. 876. The result has been arrived at chiefly from the study of the Amphibia; but it appears from the researches of Waldeyer (as quoted in 'Journal of Anat. and Phys.' 1869, p. 161), that the sexual organs of even "the higher vertebrata are, in their early condition, hermaphrodite." Similar views have long been held by some authors, though until recently without a firm basis.) But here we encounter a singular difficulty. In the mammalian class the males possess rudiments of a uterus with the adjacent passage, in their vesiculae prostaticae; they bear also rudiments of mammae, and some male Marsupials have traces of a marsupial sack. (27. The male Thylacinus offers the best instance. Owen, 'Anatomy of Vertebrates,' vol. iii. p. 771.) Other analogous facts could be added. Are we, then, to suppose that some extremely ancient mammal continued androgynous, after it had acquired the chief distinctions of its class, and therefore after it had diverged from the lower classes of the vertebrate kingdom? This seems very improbable, for we have to look to fishes, the lowest of all the classes, to find any still existent androgynous forms. (28. Hermaphroditism has been observed in several species of Serranus, as well as in some other fishes, where it is either normal and symmetrical, or abnormal and unilateral. Dr. Zouteveen has given me references on this subject, more especially to a paper by Prof. Halbertsma, in the 'Transact. of the Dutch Acad. of Sciences,' vol. xvi. Dr. Gunther doubts the fact, but it has now been recorded by too many good observers to be any longer disputed. Dr. M. Lessona writes to me, that he has verified the observations made by Cavolini on Serranus. Prof. Ercolani has recently shewn ('Accad. delle Scienze,' Bologna, Dec. 28, 1871) that eels are androgynous.) That various accessory parts, proper to each sex, are found in a rudimentary condition in the opposite sex, may be explained by such organs having been gradually acquired by the one sex, and then transmitted in a more or less imperfect state to the other. When we treat of sexual selection, we shall meet with innumerable instances of this form of transmission,--as in the case of the spurs, plumes, and brilliant colours, acquired for battle or ornament by male birds, and inherited by the females in an imperfect or rudimentary condition.
The possession by male mammals of functionally imperfect mammary organs is, in some respects, especially curious. The Monotremata have the proper milk-secreting glands with orifices, but no nipples; and as these animals stand at the very base of the mammalian series, it is probable that the progenitors of the class also had milk-secreting glands, but no nipples. This conclusion is supported by what is known of their manner of development; for Professor Turner informs me, on the authority of Kolliker and Langer, that in the embryo the mammary glands can be distinctly traced before the nipples are in the least visible; and the development of successive parts in the individual generally represents and accords with the development of successive beings in the same line of descent. The Marsupials differ from the Monotremata by possessing nipples; so that probably these organs were first acquired by the Marsupials, after they had diverged from, and risen above, the Monotremata, and were then transmitted to the placental mammals. (29. Prof. Gegenbaur has shewn ('Jenaische Zeitschrift,' Bd. vii. p. 212) that two distinct types of nipples prevail throughout the several mammalian orders, but that it is quite intelligible how both could have been derived from the nipples of the Marsupials, and the latter from those of the Monotremata. See, also, a memoir by Dr. Max Huss, on the mammary glands, ibid. B. viii. p. 176.) No one will suppose that the marsupials still remained androgynous, after they had approximately acquired their present structure. How then are we to account for male mammals possessing mammae? It is possible that they were first developed in the females and then transferred to the males, but from what follows this is hardly probable.
It may be suggested, as another view, that long after the progenitors of the whole mammalian class had ceased to be androgynous, both sexes yielded milk, and thus nourished their young; and in the case of the Marsupials, that both sexes carried their young in marsupial sacks. This will not appear altogether improbable, if we reflect that the males of existing syngnathous fishes receive the eggs of the females in their abdominal pouches, hatch them, and afterwards, as some believe, nourish the young (30. Mr. Lockwood believes (as quoted in 'Quart. Journal of Science,' April 1868, p. 269), from what he has observed of the development of Hippocampus, that the walls of the abdominal pouch of the male in some way afford nourishment. On male fishes hatching the ova in their mouths, see a very interesting paper by Prof. Wyman, in 'Proc. Boston Soc. of Nat. Hist.' Sept. 15, 1857; also Prof. Turner, in 'Journal of Anatomy and Physiology,' Nov. 1, 1866, p. 78. Dr. Gunther has likewise described similar cases.);-- that certain other male fishes hatch the eggs within their mouths or branchial cavities;--that certain male toads take the chaplets of eggs from the females, and wind them round their own thighs, keeping them there until the tadpoles are born;--that certain male birds undertake the whole duty of incubation, and that male pigeons, as well as the females, feed their nestlings with a secretion from their crops. But the above suggestion first occurred to me from mammary glands of male mammals being so much more perfectly developed than the rudiments of the other accessory reproductive parts, which are found in the one sex though proper to the other. The mammary glands and nipples, as they exist in male mammals, can indeed hardly be called rudimentary; they are merely not fully developed, and not functionally active. They are sympathetically affected under the influence of certain diseases, like the same organs in the female. They often secrete a few drops of milk at birth and at puberty: this latter fact occurred in the curious case, before referred to, where a young man possessed two pairs of mammae. In man and some other male mammals these organs have been known occasionally to become so well developed during maturity as to yield a fair supply of milk. Now if we suppose that during a former prolonged period male mammals aided the females in nursing their offspring (31. Mlle. C. Royer has suggested a similar view in her 'Origine de l'homme,' etc., 1870.), and that afterwards from some cause (as from the production of a smaller number of young) the males ceased to give this aid, disuse of the organs during maturity would lead to their becoming inactive; and from two well-known principles of inheritance, this state of inactivity would probably be transmitted to the males at the corresponding age of maturity. But at an earlier age these organs would be left unaffected, so that they would be almost equally well developed in the young of both sexes.
CONCLUSION.
Von Baer has defined advancement or progress in the organic scale better than any one else, as resting on the amount of differentiation and specialisation of the several parts of a being,--when arrived at maturity, as I should be inclined to add. Now as organisms have become slowly adapted to diversified lines of life by means of natural selection, their parts will have become more and more differentiated and specialised for various functions from the advantage gained by the division of physiological labour. The same part appears often to have been modified first for one purpose, and then long afterwards for some other and quite distinct purpose; and thus all the parts are rendered more and more complex. But each organism still retains the general type of structure of the progenitor from which it was aboriginally derived. In accordance with this view it seems, if we turn to geological evidence, that organisation on the whole has advanced throughout the world by slow and interrupted steps. In the great kingdom of the Vertebrata it has culminated in man. It must not, however, be supposed that groups of organic beings are always supplanted, and disappear as soon as they have given birth to other and more perfect groups. The latter, though victorious over their predecessors, may not have become better adapted for all places in the economy of nature. Some old forms appear to have survived from inhabiting protected sites, where they have not been exposed to very severe competition; and these often aid us in constructing our genealogies, by giving us a fair idea of former and lost populations. But we must not fall into the error of looking at the existing members of any lowly-organised group as perfect representatives of their ancient predecessors.
The most ancient progenitors in the kingdom of the Vertebrata, at which we are able to obtain an obscure glance, apparently consisted of a group of marine animals (32. The inhabitants of the seashore must be greatly affected by the tides; animals living either about the MEAN high-water mark, or about the MEAN low-water mark, pass through a complete cycle of tidal changes in a fortnight. Consequently, their food supply will undergo marked changes week by week. The vital functions of such animals, living under these conditions for many generations, can hardly fail to run their course in regular weekly periods. Now it is a mysterious fact that in the higher and now terrestrial Vertebrata, as well as in other classes, many normal and abnormal processes have one or more whole weeks as their periods; this would be rendered intelligible if the Vertebrata are descended from an animal allied to the existing tidal Ascidians. Many instances of such periodic processes might be given, as the gestation of mammals, the duration of fevers, etc. The hatching of eggs affords also a good example, for, according to Mr. Bartlett ('Land and Water,' Jan. 7, 1871), the eggs of the pigeon are hatched in two weeks; those of the fowl in three; those of the duck in four; those of the goose in five; and those of the ostrich in seven weeks. As far as we can judge, a recurrent period, if approximately of the right duration for any process or function, would not, when once gained, be liable to change; consequently it might be thus transmitted through almost any number of generations. But if the function changed, the period would have to change, and would be apt to change almost abruptly by a whole week. This conclusion, if sound, is highly remarkable; for the period of gestation in each mammal, and the hatching of each bird's eggs, and many other vital processes, thus betray to us the primordial birthplace of these animals.), resembling the larvae of existing Ascidians. These animals probably gave rise to a group of fishes, as lowly organised as the lancelet; and from these the Ganoids, and other fishes like the Lepidosiren, must have been developed. From such fish a very small advance would carry us on to the Amphibians. We have seen that birds and reptiles were once intimately connected together; and the Monotremata now connect mammals with reptiles in a slight degree. But no one can at present say by what line of descent the three higher and related classes, namely, mammals, birds, and reptiles, were derived from the two lower vertebrate classes, namely, amphibians and fishes. In the class of mammals the steps are not difficult to conceive which led from the ancient Monotremata to the ancient Marsupials; and from these to the early progenitors of the placental mammals. We may thus ascend to the Lemuridae; and the interval is not very wide from these to the Simiadae. The Simiadae then branched off into two great stems, the New World and Old World monkeys; and from the latter, at a remote period, Man, the wonder and glory of the Universe, proceeded.
Thus we have given to man a pedigree of prodigious length, but not, it may be said, of noble quality. The world, it has often been remarked, appears as if it had long been preparing for the advent of man: and this, in one sense is strictly true, for he owes his birth to a long line of progenitors. If any single link in this chain had never existed, man would not have been exactly what he now is. Unless we wilfully close our eyes, we may, with our present knowledge, approximately recognise our parentage; nor need we feel ashamed of it. The most humble organism is something much higher than the inorganic dust under our feet; and no one with an unbiassed mind can study any living creature, however humble, without being struck with enthusiasm at its marvellous structure and properties.