The Descent of Man, and Selection in Relation to Sex

This subject, though not intrinsically more important than the two last, will for several reasons be treated here more fully. (22. I had written a rough copy of this chapter before reading a valuable paper, "Caratteri rudimentali in ordine all' origine dell' uomo" ('Annuario della Soc. d. Naturalisti,' Modena, 1867, p. 81), by G. Canestrini, to which paper I am considerably indebted. Haeckel has given admirable discussions on this whole subject, under the title of Dysteleology, in his 'Generelle Morphologie' and 'Schöpfungsgeschichte.') Not one of the higher animals can be named which does not bear some part in a rudimentary condition; and man forms no exception to the rule. Rudimentary organs must be distinguished from those that are nascent; though in some cases the distinction is not easy. The former are either absolutely useless, such as the mammae of male quadrupeds, or the incisor teeth of ruminants which never cut through the gums; or they are of such slight service to their present possessors, that we can hardly suppose that they were developed under the conditions which now exist. Organs in this latter state are not strictly rudimentary, but they are tending in this direction. Nascent organs, on the other hand, though not fully developed, are of high service to their possessors, and are capable of further development. Rudimentary organs are eminently variable; and this is partly intelligible, as they are useless, or nearly useless, and consequently are no longer subjected to natural selection. They often become wholly suppressed. When this occurs, they are nevertheless liable to occasional reappearance through reversion – a circumstance well worthy of attention.

The chief agents in causing organs to become rudimentary seem to have been disuse at that period of life when the organ is chiefly used (and this is generally during maturity), and also inheritance at a corresponding period of life. The term "disuse" does not relate merely to the lessened action of muscles, but includes a diminished flow of blood to a part or organ, from being subjected to fewer alternations of pressure, or from becoming in any way less habitually active. Rudiments, however, may occur in one sex of those parts which are normally present in the other sex; and such rudiments, as we shall hereafter see, have often originated in a way distinct from those here referred to. In some cases, organs have been reduced by means of natural selection, from having become injurious to the species under changed habits of life. The process of reduction is probably often aided through the two principles of compensation and economy of growth; but the later stages of reduction, after disuse has done all that can fairly be attributed to it, and when the saving to be effected by the economy of growth would be very small (23. Some good criticisms on this subject have been given by Messrs. Murie and Mivart, in 'Transact. Zoological Society,' 1869, vol. vii. p. 92.), are difficult to understand. The final and complete suppression of a part, already useless and much reduced in size, in which case neither compensation nor economy can come into play, is perhaps intelligible by the aid of the hypothesis of pangenesis. But as the whole subject of rudimentary organs has been discussed and illustrated in my former works (24. 'Variation of Animals and Plants under Domestication,' vol. ii pp. 317 and 397. See also 'Origin of Species,' 5th Edition p. 535.), I need here say no more on this head.

Rudiments of various muscles have been observed in many parts of the human body (25. For instance, M. Richard ('Annales des Sciences Nat.,' 3rd series, Zoolog. 1852, tom. xviii. p. 13) describes and figures rudiments of what he calls the "muscle pedieux de la main," which he says is sometimes "infiniment petit." Another muscle, called "le tibial posterieur," is generally quite absent in the hand, but appears from time to time in a more or less rudimentary condition.); and not a few muscles, which are regularly present in some of the lower animals can occasionally be detected in man in a greatly reduced condition. Every one must have noticed the power which many animals, especially horses, possess of moving or twitching their skin; and this is effected by the panniculus carnosus. Remnants of this muscle in an efficient state are found in various parts of our bodies; for instance, the muscle on the forehead, by which the eyebrows are raised. The platysma myoides, which is well developed on the neck, belongs to this system. Prof. Turner, of Edinburgh, has occasionally detected, as he informs me, muscular fasciculi in five different situations, namely in the axillae, near the scapulae, etc., all of which must be referred to the system of the panniculus. He has also shewn (26. Prof. W. Turner, 'Proceedings of the Royal Society of Edinburgh,' 1866-67, p. 65.) that the musculus sternalis or sternalis brutorum, which is not an extension of the rectus abdominalis, but is closely allied to the panniculus, occurred in the proportion of about three per cent. in upwards of 600 bodies: he adds, that this muscle affords "an excellent illustration of the statement that occasional and rudimentary structures are especially liable to variation in arrangement."

Some few persons have the power of contracting the superficial muscles on their scalps; and these muscles are in a variable and partially rudimentary condition. M. A. de Candolle has communicated to me a curious instance of the long-continued persistence or inheritance of this power, as well as of its unusual development. He knows a family, in which one member, the present head of the family, could, when a youth, pitch several heavy books from his head by the movement of the scalp alone; and he won wagers by performing this feat. His father, uncle, grandfather, and his three children possess the same power to the same unusual degree. This family became divided eight generations ago into two branches; so that the head of the above-mentioned branch is cousin in the seventh degree to the head of the other branch. This distant cousin resides in another part of France; and on being asked whether he possessed the same faculty, immediately exhibited his power. This case offers a good illustration how persistent may be the transmission of an absolutely useless faculty, probably derived from our remote semi-human progenitors; since many monkeys have, and frequently use the power, of largely moving their scalps up and down. (27. See my 'Expression of the Emotions in Man and Animals,' 1872, p. 144.)

The extrinsic muscles which serve to move the external ear, and the intrinsic muscles which move the different parts, are in a rudimentary condition in man, and they all belong to the system of the panniculus; they are also variable in development, or at least in function. I have seen one man who could draw the whole ear forwards; other men can draw it upwards; another who could draw it backwards (28. Canestrini quotes Hyrtl. ('Annuario della Soc. dei Naturalisti,' Modena, 1867, p. 97) to the same effect.); and from what one of these persons told me, it is probable that most of us, by often touching our ears, and thus directing our attention towards them, could recover some power of movement by repeated trials. The power of erecting and directing the shell of the ears to the various points of the compass, is no doubt of the highest service to many animals, as they thus perceive the direction of danger; but I have never heard, on sufficient evidence, of a man who possessed this power, the one which might be of use to him. The whole external shell may be considered a rudiment, together with the various folds and prominences (helix and anti-helix, tragus and anti-tragus, etc.) which in the lower animals strengthen and support the ear when erect, without adding much to its weight. Some authors, however, suppose that the cartilage of the shell serves to transmit vibrations to the acoustic nerve; but Mr. Toynbee (29. 'The Diseases of the Ear,' by J. Toynbee, F.R.S., 1860, p. 12. A distinguished physiologist, Prof. Preyer, informs me that he had lately been experimenting on the function of the shell of the ear, and has come to nearly the same conclusion as that given here.), after collecting all the known evidence on this head, concludes that the external shell is of no distinct use. The ears of the chimpanzee and orang are curiously like those of man, and the proper muscles are likewise but very slightly developed. (30. Prof. A. Macalister, 'Annals and Magazine of Natural History,' vol. vii. 1871, p. 342.) I am also assured by the keepers in the Zoological Gardens that these animals never move or erect their ears; so that they are in an equally rudimentary condition with those of man, as far as function is concerned. Why these animals, as well as the progenitors of man, should have lost the power of erecting their ears, we cannot say. It may be, though I am not satisfied with this view, that owing to their arboreal habits and great strength they were but little exposed to danger, and so during a lengthened period moved their ears but little, and thus gradually lost the power of moving them. This would be a parallel case with that of those large and heavy birds, which, from inhabiting oceanic islands, have not been exposed to the attacks of beasts of prey, and have consequently lost the power of using their wings for flight. The inability to move the ears in man and several apes is, however, partly compensated by the freedom with which they can move the head in a horizontal plane, so as to catch sounds from all directions. It has been asserted that the ear of man alone possesses a lobule; but "a rudiment of it is found in the gorilla" (31. Mr. St. George Mivart, 'Elementary Anatomy,' 1873, p. 396.); and, as I hear from Prof. Preyer, it is not rarely absent in the negro.

[Fig. 2. Human Ear, modelled and drawn by Mr. Woolner. The projecting point is labelled a.]

The celebrated sculptor, Mr. Woolner, informs me of one little peculiarity in the external ear, which he has often observed both in men and women, and of which he perceived the full significance. His attention was first called to the subject whilst at work on his figure of Puck, to which he had given pointed ears. He was thus led to examine the ears of various monkeys, and subsequently more carefully those of man. The peculiarity consists in a little blunt point, projecting from the inwardly folded margin, or helix. When present, it is developed at birth, and, according to Prof. Ludwig Meyer, more frequently in man than in woman. Mr. Woolner made an exact model of one such case, and sent me the accompanying drawing. (Fig. 2). These points not only project inwards towards the centre of the ear, but often a little outwards from its plane, so as to be visible when the head is viewed from directly in front or behind. They are variable in size, and somewhat in position, standing either a little higher or lower; and they sometimes occur on one ear and not on the other. They are not confined to mankind, for I observed a case in one of the spider-monkeys (Ateles beelzebuth) in our Zoological Gardens; and Mr. E. Ray Lankester informs me of another case in a chimpanzee in the gardens at Hamburg. The helix obviously consists of the extreme margin of the ear folded inwards; and this folding appears to be in some manner connected with the whole external ear being permanently pressed backwards. In many monkeys, which do not stand high in the order, as baboons and some species of macacus (32. See also some remarks, and the drawings of the ears of the Lemuroidea, in Messrs. Murie and Mivart's excellent paper in 'Transactions of the Zoological Society,' vol. vii. 1869, pp. 6 and 90.), the upper portion of the ear is slightly pointed, and the margin is not at all folded inwards; but if the margin were to be thus folded, a slight point would necessarily project inwards towards the centre, and probably a little outwards from the plane of the ear; and this I believe to be their origin in many cases. On the other hand, Prof. L. Meyer, in an able paper recently published (33. 'Über das Darwin'sche Spitzohr,' Archiv fur Path. Anat. und Phys., 1871, p. 485.), maintains that the whole case is one of mere variability; and that the projections are not real ones, but are due to the internal cartilage on each side of the points not having been fully developed. I am quite ready to admit that this is the correct explanation in many instances, as in those figured by Prof. Meyer, in which there are several minute points, or the whole margin is sinuous. I have myself seen, through the kindness of Dr. L. Down, the ear of a microcephalous idiot, on which there is a projection on the outside of the helix, and not on the inward folded edge, so that this point can have no relation to a former apex of the ear. Nevertheless in some cases, my original view, that the points are vestiges of the tips of formerly erect and pointed ears, still seems to me probable. I think so from the frequency of their occurrence, and from the general correspondence in position with that of the tip of a pointed ear. In one case, of which a photograph has been sent me, the projection is so large, that supposing, in accordance with Prof. Meyer's view, the ear to be made perfect by the equal development of the cartilage throughout the whole extent of the margin, it would have covered fully one-third of the whole ear. Two cases have been communicated to me, one in North America, and the other in England, in which the upper margin is not at all folded inwards, but is pointed, so that it closely resembles the pointed ear of an ordinary quadruped in outline. In one of these cases, which was that of a young child, the father compared the ear with the drawing which I have given (34. 'The Expression of the Emotions,' p. 136.) of the ear of a monkey, the Cynopithecus niger, and says that their outlines are closely similar. If, in these two cases, the margin had been folded inwards in the normal manner, an inward projection must have been formed. I may add that in two other cases the outline still remains somewhat pointed, although the margin of the upper part of the ear is normally folded inwards – in one of them, however, very narrowly. [Fig.3. Foetus of an Orang(?). Exact copy of a photograph, shewing the form of the ear at this early age.] The following woodcut (No. 3) is an accurate copy of a photograph of the foetus of an orang (kindly sent me by Dr. Nitsche), in which it may be seen how different the pointed outline of the ear is at this period from its adult condition, when it bears a close general resemblance to that of man. It is evident that the folding over of the tip of such an ear, unless it changed greatly during its further development, would give rise to a point projecting inwards. On the whole, it still seems to me probable that the points in question are in some cases, both in man and apes, vestiges of a former condition.

The nictitating membrane, or third eyelid, with its accessory muscles and other structures, is especially well developed in birds, and is of much functional importance to them, as it can be rapidly drawn across the whole eye-ball. It is found in some reptiles and amphibians, and in certain fishes, as in sharks. It is fairly well developed in the two lower divisions of the mammalian series, namely, in the monotremata and marsupials, and in some few of the higher mammals, as in the walrus. But in man, the quadrumana, and most other mammals, it exists, as is admitted by all anatomists, as a mere rudiment, called the semilunar fold. (35. Muller's 'Elements of Physiology,' Eng. translat. 1842, vol. ii. p. 1117. Owen, 'Anatomy of Vertebrates,' vol. iii. p. 260; ibid. on the Walrus, 'Proceedings of the Zoological Society,' November 8, 1854. See also R. Knox, 'Great Artists and Anatomists,' p. 106. This rudiment apparently is somewhat larger in Negroes and Australians than in Europeans, see Carl Vogt, 'Lectures on Man,' Eng. translat. p. 129.)

The sense of smell is of the highest importance to the greater number of mammals – to some, as the ruminants, in warning them of danger; to others, as the Carnivora, in finding their prey; to others, again, as the wild boar, for both purposes combined. But the sense of smell is of extremely slight service, if any, even to the dark coloured races of men, in whom it is much more highly developed than in the white and civilised races. (36. The account given by Humboldt of the power of smell possessed by the natives of South America is well known, and has been confirmed by others. M. Houzeau ('Études sur les Facultés Mentales,' etc., tom. i. 1872, p. 91) asserts that he repeatedly made experiments, and proved that Negroes and Indians could recognise persons in the dark by their odour. Dr. W. Ogle has made some curious observations on the connection between the power of smell and the colouring matter of the mucous membrane of the olfactory region as well as of the skin of the body. I have, therefore, spoken in the text of the dark-coloured races having a finer sense of smell than the white races. See his paper, 'Medico-Chirurgical Transactions,' London, vol. liii. 1870, p. 276.) Nevertheless it does not warn them of danger, nor guide them to their food; nor does it prevent the Esquimaux from sleeping in the most fetid atmosphere, nor many savages from eating half-putrid meat. In Europeans the power differs greatly in different individuals, as I am assured by an eminent naturalist who possesses this sense highly developed, and who has attended to the subject. Those who believe in the principle of gradual evolution, will not readily admit that the sense of smell in its present state was originally acquired by man, as he now exists. He inherits the power in an enfeebled and so far rudimentary condition, from some early progenitor, to whom it was highly serviceable, and by whom it was continually used. In those animals which have this sense highly developed, such as dogs and horses, the recollection of persons and of places is strongly associated with their odour; and we can thus perhaps understand how it is, as Dr. Maudsley has truly remarked (37. 'The Physiology and Pathology of Mind,' 2nd ed. 1868, p. 134.), that the sense of smell in man "is singularly effective in recalling vividly the ideas and images of forgotten scenes and places."

Man differs conspicuously from all the other primates in being almost naked. But a few short straggling hairs are found over the greater part of the body in the man, and fine down on that of the woman. The different races differ much in hairiness; and in the individuals of the same race the hairs are highly variable, not only in abundance, but likewise in position: thus in some Europeans the shoulders are quite naked, whilst in others they bear thick tufts of hair. (38. Eschricht, Über die Richtung der Haare am menschlichen Körper, Muller's 'Archiv fur Anat. und Phys.' 1837, s. 47. I shall often have to refer to this very curious paper.) There can be little doubt that the hairs thus scattered over the body are the rudiments of the uniform hairy coat of the lower animals. This view is rendered all the more probable, as it is known that fine, short, and pale-coloured hairs on the limbs and other parts of the body, occasionally become developed into "thickset, long, and rather coarse dark hairs," when abnormally nourished near old-standing inflamed surfaces. (39. Paget, 'Lectures on Surgical Pathology,' 1853, vol. i. p. 71.)

I am informed by Sir James Paget that often several members of a family have a few hairs in their eyebrows much longer than the others; so that even this slight peculiarity seems to be inherited. These hairs, too, seem to have their representatives; for in the chimpanzee, and in certain species of Macacus, there are scattered hairs of considerable length rising from the naked skin above the eyes, and corresponding to our eyebrows; similar long hairs project from the hairy covering of the superciliary ridges in some baboons.

The fine wool-like hair, or so-called lanugo, with which the human foetus during the sixth month is thickly covered, offers a more curious case. It is first developed, during the fifth month, on the eyebrows and face, and especially round the mouth, where it is much longer than that on the head. A moustache of this kind was observed by Eschricht (40. Eschricht, ibid. s. 40, 47.) on a female foetus; but this is not so surprising a circumstance as it may at first appear, for the two sexes generally resemble each other in all external characters during an early period of growth. The direction and arrangement of the hairs on all parts of the foetal body are the same as in the adult, but are subject to much variability. The whole surface, including even the forehead and ears, is thus thickly clothed; but it is a significant fact that the palms of the hands and the soles of the feet are quite naked, like the inferior surfaces of all four extremities in most of the lower animals. As this can hardly be an accidental coincidence, the woolly covering of the foetus probably represents the first permanent coat of hair in those mammals which are born hairy. Three or four cases have been recorded of persons born with their whole bodies and faces thickly covered with fine long hairs; and this strange condition is strongly inherited, and is correlated with an abnormal condition of the teeth. (41. See my 'Variation of Animals and Plants under Domestication,' vol. ii. p. 327. Prof. Alex. Brandt has recently sent me an additional case of a father and son, born in Russia, with these peculiarities. I have received drawings of both from Paris.) Prof. Alex. Brandt informs me that he has compared the hair from the face of a man thus characterised, aged thirty-five, with the lanugo of a foetus, and finds it quite similar in texture; therefore, as he remarks, the case may be attributed to an arrest of development in the hair, together with its continued growth. Many delicate children, as I have been assured by a surgeon to a hospital for children, have their backs covered by rather long silky hairs; and such cases probably come under the same head.

It appears as if the posterior molar or wisdom-teeth were tending to become rudimentary in the more civilised races of man. These teeth are rather smaller than the other molars, as is likewise the case with the corresponding teeth in the chimpanzee and orang; and they have only two separate fangs. They do not cut through the gums till about the seventeenth year, and I have been assured that they are much more liable to decay, and are earlier lost than the other teeth; but this is denied by some eminent dentists. They are also much more liable to vary, both in structure and in the period of their development, than the other teeth. (42. Dr. Webb, 'Teeth in Man and the Anthropoid Apes,' as quoted by Dr. C. Carter Blake in Anthropological Review, July 1867, p. 299.) In the Melanian races, on the other hand, the wisdom-teeth are usually furnished with three separate fangs, and are generally sound; they also differ from the other molars in size, less than in the Caucasian races. (43. Owen, 'Anatomy of Vertebrates,' vol. iii. pp. 320, 321, and 325.) Prof. Schaaffhausen accounts for this difference between the races by "the posterior dental portion of the jaw being always shortened" in those that are civilised (44. 'On the Primitive Form of the Skull,' Eng. translat., in 'Anthropological Review,' Oct. 1868, p. 426), and this shortening may, I presume, be attributed to civilised men habitually feeding on soft, cooked food, and thus using their jaws less. I am informed by Mr. Brace that it is becoming quite a common practice in the United States to remove some of the molar teeth of children, as the jaw does not grow large enough for the perfect development of the normal number. (45. Prof. Montegazza writes to me from Florence, that he has lately been studying the last molar teeth in the different races of man, and has come to the same conclusion as that given in my text, viz., that in the higher or civilised races they are on the road towards atrophy or elimination.)

With respect to the alimentary canal, I have met with an account of only a single rudiment, namely the vermiform appendage of the caecum. The caecum is a branch or diverticulum of the intestine, ending in a cul-de-sac, and is extremely long in many of the lower vegetable-feeding mammals. In the marsupial koala it is actually more than thrice as long as the whole body. (46. Owen, 'Anatomy of Vertebrates,' vol. iii. pp. 416, 434, 441.) It is sometimes produced into a long gradually-tapering point, and is sometimes constricted in parts. It appears as if, in consequence of changed diet or habits, the caecum had become much shortened in various animals, the vermiform appendage being left as a rudiment of the shortened part. That this appendage is a rudiment, we may infer from its small size, and from the evidence which Prof. Canestrini (47. 'Annuario della Soc. d. Nat.' Modena, 1867, p. 94.) has collected of its variability in man. It is occasionally quite absent, or again is largely developed. The passage is sometimes completely closed for half or two-thirds of its length, with the terminal part consisting of a flattened solid expansion. In the orang this appendage is long and convoluted: in man it arises from the end of the short caecum, and is commonly from four to five inches in length, being only about the third of an inch in diameter. Not only is it useless, but it is sometimes the cause of death, of which fact I have lately heard two instances: this is due to small hard bodies, such as seeds, entering the passage, and causing inflammation. (48. M. C. Martins ("De l'Unité Organique," in 'Revue des Deux Mondes,' June 15, 1862, p. 16) and Haeckel ('Generelle Morphologie,' B. ii. s. 278), have both remarked on the singular fact of this rudiment sometimes causing death.)