In the adult male anthropomorphous apes, as Rütimeyer,[207] and others have insisted, it is precisely the effect which the jaw-muscles by their great development have produced on the skull, that causes it to differ so greatly in many respects from that of man, and has given to it “a truly frightful physiognomy.” Therefore as the jaws and teeth in the progenitors of man gradually become reduced in size, the adult skull would have presented nearly the same characters which it offers in the young of the anthropomorphous apes, and would thus have come to resemble more nearly that of existing man. A great reduction of the canine teeth in the males would almost certainly, as we shall hereafter see, have affected through inheritance the teeth of the females.
As the various mental faculties were gradually developed, the brain would almost certainly have become larger. No one, I presume, doubts that the large size of the brain in man, relatively to his body, in comparison with that of the gorilla or orang, is closely connected with his higher mental powers. We meet with closely analogous facts with insects, in which the cerebral ganglia are of extraordinary dimensions in ants; these ganglia in all the Hymenoptera being many times larger than in the less intelligent orders, such as beetles.[208] On the other hand, no one supposes that the intellect of any two animals or of any two men can be accurately gauged by the cubic contents of their skulls. It is certain that there may be extraordinary mental activity with an extremely small absolute mass of nervous matter: thus the wonderfully diversified instincts, mental powers, and affections of ants are generally known, yet their cerebral ganglia are not so large as the quarter of a small pin’s head. Under this latter point of view, the brain of an ant is one of the most marvellous atoms of matter in the world, perhaps more marvellous than the brain of man.
The belief that there exists in man some close relation between the size of the brain and the development of the intellectual faculties is supported by the comparison of the skulls of savage and civilised races, of ancient and modern people, and by the analogy of the whole vertebrate series. Dr. J. Barnard Davis has proved[209] by many careful measurements, that the mean internal capacity of the skull in Europeans is 92·3 cubic inches; in Americans 87·5; in Asiatics 87·1; and in Australians only 81·9 inches. Professor Broca[210] found that skulls from graves in Paris of the nineteenth century, were larger than those from vaults of the twelfth century, in the proportion of 1484 to 1426; and Prichard is persuaded that the present inhabitants of Britain have “much more capacious brain-cases” than the ancient inhabitants. Nevertheless it must be admitted that some skulls of very high antiquity, such as the famous one of Neanderthal, are well developed and capacious. With respect to the lower animals, M. E. Lartet,[211] by comparing the crania of tertiary and recent mammals, belonging to the same groups, has come to the remarkable conclusion that the brain is generally larger and the convolutions more complex in the more recent form. On the other hand I have shewn[212] that the brains of domestic rabbits are considerably reduced in bulk, in comparison with those of the wild rabbit or hare; and this may be attributed to their having been closely confined during many generations, so that they have exerted but little their intellect, instincts, senses, and voluntary movements.
The gradually increasing weight of the brain and skull in man must have influenced the development of the supporting spinal column, more especially whilst he was becoming erect. As this change of position was being brought about, the internal pressure of the brain, will, also, have influenced the form of the skull; for many facts shew how easily the skull is thus affected. Ethnologists believe that it is modified by the kind of cradle in which infants sleep. Habitual spasms of the muscles and a cicatrix from a severe burn have permanently modified the facial bones. In young persons whose heads from disease have become fixed either sideways or backwards, one of the eyes has changed its position, and the bones of the skull have been modified; and this apparently results from the brain pressing in a new direction.[213] I have shewn that with long-eared rabbits, even so trifling a cause as the lopping forward of one ear drags forward on that side almost every bone of the skull; so that the bones on the opposite sides no longer strictly correspond. Lastly, if any animal were to increase or diminish much in general size, without any change in its mental powers; or if the mental powers were to be much increased or diminished without any great change in the size of the body; the shape of the skull would almost certainly be altered. I infer this from my observations on domestic rabbits, some kinds of which have become very much larger than the wild animal, whilst others have retained nearly the same size, but in both cases the brain has been much reduced relatively to the size of the body. Now I was at first much surprised by finding that in all these rabbits the skull had become elongated or dolichocephalic; for instance, of two skulls of nearly equal breadth, the one from a wild rabbit and the other from a large domestic kind, the former was only 3·15 and the latter 4·3 inches in length.[214] One of the most marked distinctions in different races of man is that the skull in some is elongated, and in others rounded; and here the explanation suggested by the case of the rabbits may partially hold good; for Welcker finds that short “men incline more to brachycephaly, and tall men to dolichocephaly;”[215] and tall men may be compared with the larger and longer-bodied rabbits, all of which have elongated skulls, or are dolichocephalic.
From these several facts we can to a certain extent understand the means through which the great size and more or less rounded form of the skull has been acquired by man; and these are characters eminently distinctive of him in comparison with the lower animals.
Another most conspicuous difference between man and the lower animals is the nakedness of his skin. Whales and dolphins (Cetacea), dugongs (Sirenia) and the hippopotamus are naked; and this may be advantageous to them for gliding through the water; nor would it be injurious to them from the loss of warmth, as the species which inhabit the colder regions are protected by a thick layer of blubber, serving the same purpose as the fur of seals and otters. Elephants and rhinoceroses are almost hairless; and as certain extinct species which formerly lived under an arctic climate were covered with long wool or hair, it would almost appear as if the existing species of both genera had lost their hairy covering from exposure to heat. This appears the more probable, as the elephants in India which live on elevated and cool districts are more hairy[216] than those on the lowlands. May we then infer that man became divested of hair from having aboriginally inhabited some tropical land? The fact of the hair being chiefly retained in the male sex on the chest and face, and in both sexes at the junction of all four limbs with the trunk, favours this inference, assuming that the hair was lost before man became erect; for the parts which now retain most hair would then have been most protected from the heat of the sun. The crown of the head, however, offers a curious exception, for at all times it must have been one of the most exposed parts, yet it is thickly clothed with hair. In this respect man agrees with the great majority of quadrupeds, which generally have their upper and exposed surfaces more thickly clothed than the lower surface. Nevertheless, the fact that the other members of the order of Primates, to which man belongs, although inhabiting various hot regions, are well clothed with hair, generally thickest on the upper surface,[217] is strongly opposed to the supposition that man became naked through the action of the sun. I am inclined to believe, as we shall see under sexual selection, that man, or rather primarily woman, became divested of hair for ornamental purposes; and according to this belief it is not surprising that man should differ so greatly in hairiness from all his lower brethren, for characters gained through sexual selection often differ in closely-related forms to an extraordinary degree.
According to a popular impression, the absence of a tail is eminently distinctive of man; but as those apes which come nearest to man are destitute of this organ, its disappearance does not especially concern us. Nevertheless it may be well to own that no explanation, as far as I am aware, has ever been given of the loss of the tail by certain apes and man. Its loss, however, is not surprising, for it sometimes differs remarkably in length in species of the same genera: thus in some species of Macacus the tail is longer than the whole body, consisting of twenty-four vertebræ; in others it consists of a scarcely visible stump, containing only three or four vertebræ. In some kinds of baboons there are twenty-five, whilst in the mandrill there are ten very small stunted caudal vertebræ, or, according to Cuvier,[218] sometimes only five. This great diversity in the structure and length of the tail in animals belonging to the same genera, and following nearly the same habits of life, renders it probable that the tail is not of much importance to them; and if so, we might have expected that it would sometimes have become more or less rudimentary, in accordance with what we incessantly see with other structures. The tail almost always tapers towards the end whether it be long or short; and this, I presume, results from the atrophy, through disuse, of the terminal muscles together with their arteries and nerves, leading to the atrophy of the terminal bones. With respect to the os coccyx, which in man and the higher apes manifestly consists of the few basal and tapering segments of an ordinary tail, I have heard it asked how could these have become completely embedded within the body; but there is no difficulty in this respect, for in many monkeys the basal segments of the true tail are thus embedded. For instance, Mr. Murie informs me that in the skeleton of a not full-grown Macacus inornatus, he counted nine or ten caudal vertebræ, which altogether were only 1·8 inch in length. Of these the three basal ones appeared to have been embedded; the remainder forming the free part of the tail, which was only one inch in length, and half an inch in diameter. Here, then, the three embedded caudal vertebræ plainly correspond with the four coalesced vertebræ of the human os coccyx.
I have now endeavoured to shew that some of the most distinctive characters of man have in all probability been acquired, either directly, or more commonly indirectly, through natural selection. We should bear in mind that modifications in structure or constitution, which are of no service to an organism in adapting it to its habits of life, to the food which it consumes, or passively to the surrounding conditions, cannot have been thus acquired. We must not, however, be too confident in deciding what modifications are of service to each being: we should remember how little we know about the use of many parts, or what changes in the blood or tissues may serve to fit an organism for a new climate or some new kind of food. Nor must we forget the principle of correlation, by which, as Isidore Geoffroy has shewn in the case of man, many fit-range deviations of structure are tied together. Independently of correlation, a change in one part often leads through the increased or decreased use of other parts, to other changes of a quite unexpected nature. It is also well to reflect on such facts, as the wonderful growth of galls on plants caused by the poison of an insect, and on the remarkable changes of colour in the plumage of parrots when fed on certain fishes, or inoculated with the poison of toads;[219] for we can thus see that the fluids of the system, if altered for some special purpose, might induce other strange changes. We should especially bear in mind that modifications acquired and continually used during past ages for some useful purpose would probably become firmly fixed and might be long inherited.
Thus a very large yet undefined extension may safely be given to the direct and indirect results of natural selection; but I now admit, after reading the essay by Nägeli on plants, and the remarks by various authors with respect to animals, more especially those recently made by Professor Broca, that in the earlier editions of my ‘Origin of Species’ I probably attributed too much to the action of natural selection or the survival of the fittest. I have altered the fifth edition of the Origin so as to confine my remarks to adaptive changes of structure. I had not formerly sufficiently considered the existence of many structures which appear to be, as far as we can judge, neither beneficial nor injurious; and this I believe to be one of the greatest oversights as yet detected in my work. I may be permitted to say as some excuse, that I had two distinct objects in view, firstly, to shew that species had not been separately created, and secondly, that natural selection had been the chief agent of change, though largely aided by the inherited effects of habit, and slightly by the direct action of the surrounding conditions. Nevertheless I was not able to annul the influence of my former belief, then widely prevalent, that each species had been purposely created; and this led to my tacitly assuming that every detail of structure, excepting rudiments, was of some special, though unrecognised, service. Any one with this assumption in his mind would naturally extend the action of natural selection, either during past or present times, too far. Some of those who admit the principle of evolution, but reject natural selection, seem to forget, when criticising my book, that I had the above two objects in view; hence if I have erred in giving to natural selection great power, which I am far from admitting, or in having exaggerated its power, which is in itself probable, I have at least, as I hope, done good service in aiding to overthrow the dogma of separate creations.
That all organic beings, including man, present many modifications of structure which are of no service to them at present, nor have been formerly, is, as I can now see, probable. We know not what produces the numberless slight differences between the individuals of each species, for reversion only carries the problem a few steps backwards; but each peculiarity must have had its own efficient cause. If these causes, whatever they may be, were to act more uniformly and energetically during a lengthened period (and no reason can be assigned why this should not sometimes occur), the result would probably be not mere slight individual differences, but well-marked, constant modifications. Modifications which are in no way beneficial cannot have been kept uniform through natural selection, though any which were injurious would have been thus eliminated. Uniformity of character would, however, naturally follow from, the assumed uniformity of the exciting causes, and likewise from the free intercrossing of many individuals. The same organism might acquire in this manner during successive periods successive modifications, and these would be transmitted in a nearly uniform state as long as the exciting causes remained the same and there was free intercrossing. With respect to the exciting causes we can only say, as when speaking of so-called spontaneous variations, that they relate much more closely to the constitution of the varying organism, than to the nature of the conditions to which it has been subjected.