does not relate exclusively to man. The tail often differs remarkably in length within the same genus: thus in some species of Macacus it is longer than the whole body, and is formed of twenty-four vertebrae; in others it consists of a scarcely visible stump, containing only three or four vertebrae. In some kinds of baboons there are twenty-five, whilst in the mandrill there are ten very small stunted caudal vertebrae, or, according to Cuvier (90. Mr. St. George Mivart, 'Proc. Zoolog. Soc.' 1865, pp. 562, 583. Dr. J.E. Gray, 'Cat. Brit. Mus.: 'Skeletons.' Owen, 'Anatomy of Vertebrates,' vol. ii. p. 517. Isidore Geoffroy, 'Hist. Nat. Gen.' tom. ii. p. 244.), sometimes only five. The tail, whether it be long or short, almost always tapers towards the end; and this, I presume, results from the atrophy of the terminal muscles, together with their arteries and nerves, through disuse, leading to the atrophy of the terminal bones. But no explanation can at present be given of the great diversity which often occurs in its length. Here, however, we are more specially concerned with the complete external disappearance of the tail. Professor Broca has recently shewn (91. 'Revue d'Anthropologie,' 1872; 'La Constitution des vertèbres caudales.') that the tail in all quadrupeds consists of two portions, generally separated abruptly from each other; the basal portion consists of vertebrae, more or less perfectly channelled and furnished with apophyses like ordinary vertebrae; whereas those of the terminal portion are not channelled, are almost smooth, and scarcely resemble true vertebrae. A tail, though not externally visible, is really present in man and the anthropomorphous apes, and is constructed on exactly the same pattern in both. In the terminal portion the vertebrae, constituting the os coccyx, are quite rudimentary, being much reduced in size and number. In the basal portion, the vertebrae are likewise few, are united firmly together, and are arrested in development; but they have been rendered much broader and flatter than the corresponding vertebrae in the tails of other animals: they constitute what Broca calls the accessory sacral vertebrae. These are of functional importance by supporting certain internal parts and in other ways; and their modification is directly connected with the erect or semi-erect attitude of man and the anthropomorphous apes. This conclusion is the more trustworthy, as Broca formerly held a different view, which he has now abandoned. The modification, therefore, of the basal caudal vertebrae in man and the higher apes may have been effected, directly or indirectly, through natural selection.
But what are we to say about the rudimentary and variable vertebrae of the terminal portion of the tail, forming the os coccyx? A notion which has often been, and will no doubt again be ridiculed, namely, that friction has had something to do with the disappearance of the external portion of the tail, is not so ridiculous as it at first appears. Dr. Anderson (92. 'Proceedings Zoological Society,' 1872, p. 210.) states that the extremely short tail of Macacus brunneus is formed of eleven vertebrae, including the imbedded basal ones. The extremity is tendinous and contains no vertebrae; this is succeeded by five rudimentary ones, so minute that together they are only one line and a half in length, and these are permanently bent to one side in the shape of a hook. The free part of the tail, only a little above an inch in length, includes only four more small vertebrae. This short tail is carried erect; but about a quarter of its total length is doubled on to itself to the left; and this terminal part, which includes the hook-like portion, serves "to fill up the interspace between the upper divergent portion of the callosities;" so that the animal sits on it, and thus renders it rough and callous. Dr. Anderson thus sums up his observations: "These facts seem to me to have only one explanation; this tail, from its short size, is in the monkey's way when it sits down, and frequently becomes placed under the animal while it is in this attitude; and from the circumstance that it does not extend beyond the extremity of the ischial tuberosities, it seems as if the tail originally had been bent round by the will of the animal, into the interspace between the callosities, to escape being pressed between them and the ground, and that in time the curvature became permanent, fitting in of itself when the organ happens to be sat upon." Under these circumstances it is not surprising that the surface of the tail should have been roughened and rendered callous, and Dr. Murie (93. 'Proceedings Zoological Society,' 1872, p. 786.), who carefully observed this species in the Zoological Gardens, as well as three other closely allied forms with slightly longer tails, says that when the animal sits down, the tail "is necessarily thrust to one side of the buttocks; and whether long or short its root is consequently liable to be rubbed or chafed." As we now have evidence that mutilations occasionally produce an inherited effect (94. I allude to Dr. Brown-Sequard's observations on the transmitted effect of an operation causing epilepsy in guinea-pigs, and likewise more recently on the analogous effects of cutting the sympathetic nerve in the neck. I shall hereafter have occasion to refer to Mr. Salvin's interesting case of the apparently inherited effects of mot-mots biting off the barbs of their own tail-feathers. See also on the general subject 'Variation of Animals and Plants under Domestication,' vol. ii. pp. 22–24.), it is not very improbable that in short-tailed monkeys, the projecting part of the tail, being functionally useless, should after many generations have become rudimentary and distorted, from being continually rubbed and chafed. We see the projecting part in this condition in the Macacus brunneus, and absolutely aborted in the M. ecaudatus and in several of the higher apes. Finally, then, as far as we can judge, the tail has disappeared in man and the anthropomorphous apes, owing to the terminal portion having been injured by friction during a long lapse of time; the basal and embedded portion having been reduced and modified, so as to become suitable to the erect or semi-erect position.
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 do not serve to adapt an organism 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 new kinds of food. Nor must we forget the principle of correlation, by which, as Isidore Geoffroy has shewn in the case of man, many strange 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 (95. The 'Variation of Animals and Plants under Domestication,' vol. ii. pp. 280, 282.); for we can thus see that the fluids of the system, if altered for some special purpose, might induce other 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 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 Nageli 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 perhaps 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; but I am convinced, from the light gained during even the last few years, that very many structures which now appear to us useless, will hereafter be proved to be useful, and will therefore come within the range of natural selection. Nevertheless, I did not formerly consider sufficiently the existence of structures, which, as far as we can at present judge, are 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. I was not, however, able to annul the influence of my former belief, then almost universal, that each species had been purposely created; and this led to my tacit assumption 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 too far the action of natural selection, either during
