The Descent of Man, and Selection in Relation to Sex

Although man may not have been much modified during the latter stages of his existence through the increased or decreased use of parts, the facts now given shew that his liability in this respect has not been lost; and we positively know that the same law holds good with the lower animals. Consequently we may infer that when at a remote epoch the progenitors of man were in a transitional state, and were changing from quadrupeds into bipeds, natural selection would probably have been greatly aided by the inherited effects of the increased or diminished use of the different parts of the body.

ARRESTS OF DEVELOPMENT

There is a difference between arrested development and arrested growth, for parts in the former state continue to grow whilst still retaining their early condition. Various monstrosities come under this head; and some, as a cleft palate, are known to be occasionally inherited. It will suffice for our purpose to refer to the arrested brain-development of microcephalous idiots, as described in Vogt's memoir. (36. 'Mémoire sur les Microcephales,' 1867, pp. 50, 125, 169, 171, 184-198.) Their skulls are smaller, and the convolutions of the brain are less complex than in normal men. The frontal sinus, or the projection over the eye-brows, is largely developed, and the jaws are prognathous to an "effrayant" degree; so that these idiots somewhat resemble the lower types of mankind. Their intelligence, and most of their mental faculties, are extremely feeble. They cannot acquire the power of speech, and are wholly incapable of prolonged attention, but are much given to imitation. They are strong and remarkably active, continually gambolling and jumping about, and making grimaces. They often ascend stairs on all-fours; and are curiously fond of climbing up furniture or trees. We are thus reminded of the delight shewn by almost all boys in climbing trees; and this again reminds us how lambs and kids, originally alpine animals, delight to frisk on any hillock, however small. Idiots also resemble the lower animals in some other respects; thus several cases are recorded of their carefully smelling every mouthful of food before eating it. One idiot is described as often using his mouth in aid of his hands, whilst hunting for lice. They are often filthy in their habits, and have no sense of decency; and several cases have been published of their bodies being remarkably hairy. (37. Prof. Laycock sums up the character of brute-like idiots by calling them "theroid;" 'Journal of Mental Science,' July 1863. Dr. Scott ('The Deaf and Dumb,' 2nd ed. 1870, p. 10) has often observed the imbecile smelling their food. See, on this same subject, and on the hairiness of idiots, Dr. Maudsley, 'Body and Mind,' 1870, pp. 46-51. Pinel has also given a striking case of hairiness in an idiot.)

REVERSION

Many of the cases to be here given, might have been introduced under the last heading. When a structure is arrested in its development, but still continues growing, until it closely resembles a corresponding structure in some lower and adult member of the same group, it may in one sense be considered as a case of reversion. The lower members in a group give us some idea how the common progenitor was probably constructed; and it is hardly credible that a complex part, arrested at an early phase of embryonic development, should go on growing so as ultimately to perform its proper function, unless it had acquired such power during some earlier state of existence, when the present exceptional or arrested structure was normal. The simple brain of a microcephalous idiot, in as far as it resembles that of an ape, may in this sense be said to offer a case of reversion. (38. In my 'Variation of Animals under Domestication' (vol. ii. p. 57), I attributed the not very rare cases of supernumerary mammae in women to reversion. I was led to this as a probable conclusion, by the additional mammae being generally placed symmetrically on the breast; and more especially from one case, in which a single efficient mamma occurred in the inguinal region of a woman, the daughter of another woman with supernumerary mammae. But I now find (see, for instance, Prof. Preyer, 'Der Kampf um das Dasein,' 1869, s. 45) that mammae erraticae, occur in other situations, as on the back, in the armpit, and on the thigh; the mammae in this latter instance having given so much milk that the child was thus nourished. The probability that the additional mammae are due to reversion is thus much weakened; nevertheless, it still seems to me probable, because two pairs are often found symmetrically on the breast; and of this I myself have received information in several cases. It is well known that some Lemurs normally have two pairs of mammae on the breast. Five cases have been recorded of the presence of more than a pair of mammae (of course rudimentary) in the male sex of mankind; see 'Journal of Anat. and Physiology,' 1872, p. 56, for a case given by Dr. Handyside, in which two brothers exhibited this peculiarity; see also a paper by Dr. Bartels, in 'Reichert's and du Bois-Reymond's Archiv.,' 1872, p. 304. In one of the cases alluded to by Dr. Bartels, a man bore five mammae, one being medial and placed above the navel; Meckel von Hemsbach thinks that this latter case is illustrated by a medial mamma occurring in certain Cheiroptera. On the whole, we may well doubt if additional mammae would ever have been developed in both sexes of mankind, had not his early progenitors been provided with more than a single pair.

In the above work (vol. ii. p. 12), I also attributed, though with much hesitation, the frequent cases of polydactylism in men and various animals to reversion. I was partly led to this through Prof. Owen's statement, that some of the Ichthyopterygia possess more than five digits, and therefore, as I supposed, had retained a primordial condition; but Prof. Gegenbaur ('Jenaischen Zeitschrift,' B. v. Heft 3, s. 341), disputes Owen's conclusion. On the other hand, according to the opinion lately advanced by Dr. Gunther, on the paddle of Ceratodus, which is provided with articulated bony rays on both sides of a central chain of bones, there seems no great difficulty in admitting that six or more digits on one side, or on both sides, might reappear through reversion. I am informed by Dr. Zouteveen that there is a case on record of a man having twenty-four fingers and twenty-four toes! I was chiefly led to the conclusion that the presence of supernumerary digits might be due to reversion from the fact that such digits, not only are strongly inherited, but, as I then believed, had the power of regrowth after amputation, like the normal digits of the lower vertebrata. But I have explained in the second edition of my Variation under Domestication why I now place little reliance on the recorded cases of such regrowth. Nevertheless it deserves notice, inasmuch as arrested development and reversion are intimately related processes; that various structures in an embryonic or arrested condition, such as a cleft palate, bifid uterus, etc., are frequently accompanied by polydactylism. This has been strongly insisted on by Meckel and Isidore Geoffroy St. – Hilaire. But at present it is the safest course to give up altogether the idea that there is any relation between the development of supernumerary digits and reversion to some lowly organised progenitor of man.) There are other cases which come more strictly under our present head of reversion. Certain structures, regularly occurring in the lower members of the group to which man belongs, occasionally make their appearance in him, though not found in the normal human embryo; or, if normally present in the human embryo, they become abnormally developed, although in a manner which is normal in the lower members of the group. These remarks will be rendered clearer by the following illustrations.

In various mammals the uterus graduates from a double organ with two distinct orifices and two passages, as in the marsupials, into a single organ, which is in no way double except from having a slight internal fold, as in the higher apes and man. The rodents exhibit a perfect series of gradations between these two extreme states. In all mammals the uterus is developed from two simple primitive tubes, the inferior portions of which form the cornua; and it is in the words of Dr. Farre, "by the coalescence of the two cornua at their lower extremities that the body of the uterus is formed in man; while in those animals in which no middle portion or body exists, the cornua remain ununited. As the development of the uterus proceeds, the two cornua become gradually shorter, until at length they are lost, or, as it were, absorbed into the body of the uterus." The angles of the uterus are still produced into cornua, even in animals as high up in the scale as the lower apes and lemurs.

Now in women, anomalous cases are not very infrequent, in which the mature uterus is furnished with cornua, or is partially divided into two organs; and such cases, according to Owen, repeat "the grade of concentrative development," attained by certain rodents. Here perhaps we have an instance of a simple arrest of embryonic development, with subsequent growth and perfect functional development; for either side of the partially double uterus is capable of performing the proper office of gestation. In other and rarer cases, two distinct uterine cavities are formed, each having its proper orifice and passage. (39. See Dr. A. Farre's well-known article in the 'Cyclopaedia of Anatomy and Physiology,' vol. v. 1859, p. 642. Owen, 'Anatomy of Vertebrates,' vol. iii. 1868, p. 687. Professor Turner, in 'Edinburgh Medical Journal,' February, 1865.) No such stage is passed through during the ordinary development of the embryo; and it is difficult to believe, though perhaps not impossible, that the two simple, minute, primitive tubes should know how (if such an expression may be used) to grow into two distinct uteri, each with a well-constructed orifice and passage, and each furnished with numerous muscles, nerves, glands and vessels, if they had not formerly passed through a similar course of development, as in the case of existing marsupials. No one will pretend that so perfect a structure as the abnormal double uterus in woman could be the result of mere chance. But the principle of reversion, by which a long-lost structure is called back into existence, might serve as the guide for its full development, even after the lapse of an enormous interval of time.

Professor Canestrini, after discussing the foregoing and various analogous cases, arrives at the same conclusion as that just given. He adduces another instance, in the case of the malar bone (40. 'Annuario della Soc. dei Naturalisti,' Modena, 1867, p. 83. Prof. Canestrini gives extracts on this subject from various authorities. Laurillard remarks, that as he has found a complete similarity in the form, proportions, and connection of the two malar bones in several human subjects and in certain apes, he cannot consider this disposition of the parts as simply accidental. Another paper on this same anomaly has been published by Dr. Saviotti in the 'Gazzetta delle Cliniche,' Turin, 1871, where he says that traces of the division may be detected in about two per cent. of adult skulls; he also remarks that it more frequently occurs in prognathous skulls, not of the Aryan race, than in others. See also G. Delorenzi on the same subject; 'Tre nuovi casi d'anomalia dell' osso malare,' Torino, 1872. Also, E. Morselli, 'Sopra una rara anomalia dell' osso malare,' Modena, 1872. Still more recently Gruber has written a pamphlet on the division of this bone. I give these references because a reviewer, without any grounds or scruples, has thrown doubts on my statements.), which, in some of the Quadrumana and other mammals, normally consists of two portions. This is its condition in the human foetus when two months old; and through arrested development, it sometimes remains thus in man when adult, more especially in the lower prognathous races. Hence Canestrini concludes that some ancient progenitor of man must have had this bone normally divided into two portions, which afterwards became fused together. In man the frontal bone consists of a single piece, but in the embryo, and in children, and in almost all the lower mammals, it consists of two pieces separated by a distinct suture. This suture occasionally persists more or less distinctly in man after maturity; and more frequently in ancient than in recent crania, especially, as Canestrini has observed, in those exhumed from the Drift, and belonging to the brachycephalic type. Here again he comes to the same conclusion as in the analogous case of the malar bones. In this, and other instances presently to be given, the cause of ancient races approaching the lower animals in certain characters more frequently than do the modern races, appears to be, that the latter stand at a somewhat greater distance in the long line of descent from their early semi-human progenitors.

Various other anomalies in man, more or less analogous to the foregoing, have been advanced by different authors, as cases of reversion; but these seem not a little doubtful, for we have to descend extremely low in the mammalian series, before we find such structures normally present. (41. A whole series of cases is given by Isidore Geoffroy St. – Hilaire, 'Hist. des Anomalies,' tom, iii, p. 437. A reviewer ('Journal of Anatomy and Physiology,' 1871, p. 366) blames me much for not having discussed the numerous cases, which have been recorded, of various parts arrested in their development. He says that, according to my theory, "every transient condition of an organ, during its development, is not only a means to an end, but once was an end in itself." This does not seem to me necessarily to hold good. Why should not variations occur during an early period of development, having no relation to reversion; yet such variations might be preserved and accumulated, if in any way serviceable, for instance, in shortening and simplifying the course of development? And again, why should not injurious abnormalities, such as atrophied or hypertrophied parts, which have no relation to a former state of existence, occur at an early period, as well as during maturity?)

In man, the canine teeth are perfectly efficient instruments for mastication. But their true canine character, as Owen (42. 'Anatomy of Vertebrates,' vol. iii. 1868, p. 323.) remarks, "is indicated by the conical form of the crown, which terminates in an obtuse point, is convex outward and flat or sub-concave within, at the base of which surface there is a feeble prominence. The conical form is best expressed in the Melanian races, especially the Australian. The canine is more deeply implanted, and by a stronger fang than the incisors." Nevertheless, this tooth no longer serves man as a special weapon for tearing his enemies or prey; it may, therefore, as far as its proper function is concerned, be considered as rudimentary. In every large collection of human skulls some may be found, as Haeckel (43. 'Generelle Morphologie,' 1866, B. ii. s. clv.) observes, with the canine teeth projecting considerably beyond the others in the same manner as in the anthropomorphous apes, but in a less degree. In these cases, open spaces between the teeth in the one jaw are left for the reception of the canines of the opposite jaw. An inter-space of this kind in a Kaffir skull, figured by Wagner, is surprisingly wide. (44. Carl Vogt's 'Lectures on Man,' Eng. translat., 1864, p. 151.) Considering how few are the ancient skulls which have been examined, compared to recent skulls, it is an interesting fact that in at least three cases the canines project largely; and in the Naulette jaw they are spoken of as enormous. (45. C. Carter Blake, on a jaw from La Naulette, 'Anthropological Review,' 1867, p. 295. Schaaffhausen, ibid. 1868, p. 426.)

Of the anthropomorphous apes the males alone have their canines fully developed; but in the female gorilla, and in a less degree in the female orang, these teeth project considerably beyond the others; therefore the fact, of which I have been assured, that women sometimes have considerably projecting canines, is no serious objection to the belief that their occasional great development in man is a case of reversion to an ape-like progenitor. He who rejects with scorn the belief that the shape of his own canines, and their occasional great development in other men, are due to our early forefathers having been provided with these formidable weapons, will probably reveal, by sneering, the line of his descent. For though he no longer intends, nor has the power, to use these teeth as weapons, he will unconsciously retract his "snarling muscles" (thus named by Sir C. Bell) (46. The Anatomy of Expression, 1844, pp. 110, 131.), so as to expose them ready for action, like a dog prepared to fight.

Many muscles are occasionally developed in man, which are proper to the Quadrumana or other mammals. Professor Vlacovich (47. Quoted by Prof. Canestrini in the 'Annuario della Soc. dei Naturalisti,' 1867, p. 90.) examined forty male subjects, and found a muscle, called by him the ischio- pubic, in nineteen of them; in three others there was a ligament which represented this muscle; and in the remaining eighteen no trace of it. In only two out of thirty female subjects was this muscle developed on both sides, but in three others the rudimentary ligament was present. This muscle, therefore, appears to be much more common in the male than in the female sex; and on the belief in the descent of man from some lower form, the fact is intelligible; for it has been detected in several of the lower animals, and in all of these it serves exclusively to aid the male in the act of reproduction.

Mr. J. Wood, in his valuable series of papers (48. These papers deserve careful study by any one who desires to learn how frequently our muscles vary, and in varying come to resemble those of the Quadrumana. The following references relate to the few points touched on in my text: 'Proc. Royal Soc.' vol. xiv. 1865, pp. 379-384; vol. xv. 1866, pp. 241, 242; vol. xv. 1867, p. 544; vol. xvi. 1868, p. 524. I may here add that Dr. Murie and Mr. St. George Mivart have shewn in their Memoir on the Lemuroidea ('Transactions, Zoological Society,' vol. vii. 1869, p. 96), how extraordinarily variable some of the muscles are in these animals, the lowest members of the Primates. Gradations, also, in the muscles leading to structures found in animals still lower in the scale, are numerous in the Lemuroidea.), has minutely described a vast number of muscular variations in man, which resemble normal structures in the lower animals. The muscles which closely resemble those regularly present in our nearest allies, the Quadrumana, are too numerous to be here even specified. In a single male subject, having a strong bodily frame, and well-formed skull, no less than seven muscular variations were observed, all of which plainly represented muscles proper to various kinds of apes. This man, for instance, had on both sides of his neck a true and powerful "levator claviculae," such as is found in all kinds of apes, and which is said to occur in about one out of sixty human subjects. (49. See also Prof. Macalister in 'Proceedings, Royal Irish Academy,' vol. x. 1868, p. 124.) Again, this man had "a special abductor of the metatarsal bone of the fifth digit, such as Professor Huxley and Mr. Flower have shewn to exist uniformly in the higher and lower apes." I will give only two additional cases; the acromio-basilar muscle is found in all mammals below man, and seems to be correlated with a quadrupedal gait, (50. Mr. Champneys in 'Journal of Anatomy and Physiology,' Nov. 1871, p. 178.) and it occurs in about one out of sixty human subjects. In the lower extremities Mr. Bradley (51. Ibid. May 1872, p. 421.) found an abductor ossis metatarsi quinti in both feet of man; this muscle had not up to that time been recorded in mankind, but is always present in the anthropomorphous apes. The muscles of the hands and arms – parts which are so eminently characteristic of man – are extremely liable to vary, so as to resemble the corresponding muscles in the lower animals. (52. Prof. Macalister (ibid. p. 121) has tabulated his observations, and finds that muscular abnormalities are most frequent in the fore-arms, secondly, in the face, thirdly, in the foot, etc.) Such resemblances are either perfect or imperfect; yet in the latter case they are manifestly of a transitional nature. Certain variations are more common in man, and others in woman, without our being able to assign any reason. Mr. Wood, after describing numerous variations, makes the following pregnant remark. "Notable departures from the ordinary type of the muscular structures run in grooves or directions, which must be taken to indicate some unknown factor, of much importance to a comprehensive knowledge of general and scientific anatomy." (53. The Rev. Dr. Haughton, after giving ('Proc. R. Irish Academy,' June 27, 1864, p. 715) a remarkable case of variation in the human flexor pollicis longus, adds, "This remarkable example shews that man may sometimes possess the arrangement of tendons of thumb and fingers characteristic of the macaque; but whether such a case should be regarded as a macaque passing upwards into a man, or a man passing downwards into a macaque, or as a congenital freak of nature, I cannot undertake to say." It is satisfactory to hear so capable an anatomist, and so embittered an opponent of evolutionism, admitting even the possibility of either of his first propositions. Prof. Macalister has also described ('Proceedings Royal Irish Academy,' vol. x. 1864, p. 138) variations in the flexor pollicis longus, remarkable from their relations to the same muscle in the Quadrumana.)

That this unknown factor is reversion to a former state of existence may be admitted as in the highest degree probable. (54. Since the first edition of this book appeared, Mr. Wood has published another memoir in the Philosophical Transactions, 1870, p. 83, on the varieties of the muscles of the human neck, shoulder, and chest. He here shews how extremely variable these muscles are, and how often and how closely the variations resemble the normal muscles of the lower animals. He sums up by remarking, "It will be enough for my purpose if I have succeeded in shewing the more important forms which, when occurring as varieties in the human subject, tend to exhibit in a sufficiently marked manner what may be considered as proofs and examples of the Darwinian principle of reversion, or law of inheritance, in this department of anatomical science.") It is quite incredible that a man should through mere accident abnormally resemble certain apes in no less than seven of his muscles, if there had been no genetic connection between them. On the other hand, if man is descended from some ape-like creature, no valid reason can be assigned why certain muscles should not suddenly reappear after an interval of many thousand generations, in the same manner as with horses, asses, and mules, dark- coloured stripes suddenly reappear on the legs, and shoulders, after an interval of hundreds, or more probably of thousands of generations.

These various cases of reversion are so closely related to those of rudimentary organs given in the first chapter, that many of them might have been indifferently introduced either there or here. Thus a human uterus furnished with cornua may be said to represent, in a rudimentary condition, the same organ in its normal state in certain mammals. Some parts which are rudimentary in man, as the os coccyx in both sexes, and the mammae in the male sex, are always present; whilst others, such as the supracondyloid foramen, only occasionally appear, and therefore might have been introduced under the head of reversion. These several reversionary structures, as well as the strictly rudimentary ones, reveal the descent of man from some lower form in an unmistakable manner.

CORRELATED VARIATION

In man, as in the lower animals, many structures are so intimately related, that when one part varies so does another, without our being able, in most cases, to assign any reason. We cannot say whether the one part governs the other, or whether both are governed by some earlier developed part. Various monstrosities, as I. Geoffroy repeatedly insists, are thus intimately connected. Homologous structures are particularly liable to change together, as we see on the opposite sides of the body, and in the upper and lower extremities. Meckel long ago remarked, that when the muscles of the arm depart from their proper type, they almost always imitate those of the leg; and so, conversely, with the muscles of the legs. The organs of sight and hearing, the teeth and hair, the colour of the skin and of the hair, colour and constitution, are more or less correlated. (55. The authorities for these several statements are given in my 'Variation of Animals under Domestication,' vol. ii. pp. 320-335.) Professor Schaaffhausen first drew attention to the relation apparently existing between a muscular frame and the strongly-pronounced supra-orbital ridges, which are so characteristic of the lower races of man.

Besides the variations which can be grouped with more or less probability under the foregoing heads, there is a large class of variations which may be provisionally called spontaneous, for to our ignorance they appear to arise without any exciting cause. It can, however, be shewn that such variations, whether consisting of slight individual differences, or of strongly-marked and abrupt deviations of structure, depend much more on the constitution of the organism than on the nature of the conditions to which it has been subjected. (56. This whole subject has been discussed in chap. xxiii. vol. ii. of my 'Variation of Animals and Plants under Domestication.')

RATE OF INCREASE

Civilised populations have been known under favourable conditions, as in the United States, to double their numbers in twenty-five years; and, according to a calculation, by Euler, this might occur in a little over twelve years. (57. See the ever memorable 'Essay on the Principle of Population,' by the Rev. T. Malthus, vol. i. 1826. pp. 6, 517.) At the former rate, the present population of the United States (thirty millions), would in 657 years cover the whole terraqueous globe so thickly, that four men would have to stand on each square yard of surface. The primary or fundamental check to the continued increase of man is the difficulty of gaining subsistence, and of living in comfort. We may infer that this is the case from what we see, for instance, in the United States, where subsistence is easy, and there is plenty of room. If such means were suddenly doubled in Great Britain, our number would be quickly doubled. With civilised nations this primary check acts chiefly by restraining marriages. The greater death-rate of infants in the poorest classes is also very important; as well as the greater mortality, from various diseases, of the inhabitants of crowded and miserable houses, at all ages. The effects of severe epidemics and wars are soon counterbalanced, and more than counterbalanced, in nations placed under favourable conditions. Emigration also comes in aid as a temporary check, but, with the extremely poor classes, not to any great extent.

There is reason to suspect, as Malthus has remarked, that the reproductive power is actually less in barbarous, than in civilised races. We know nothing positively on this head, for with savages no census has been taken; but from the concurrent testimony of missionaries, and of others who have long resided with such people, it appears that their families are usually small, and large ones rare. This may be partly accounted for, as it is believed, by the women suckling their infants during a long time; but it is highly probable that savages, who often suffer much hardship, and who do not obtain so much nutritious food as civilised men, would be actually less prolific. I have shewn in a former work (58. 'Variation of Animals and Plants under Domestication,' vol ii. pp. 111-113, 163.), that all our domesticated quadrupeds and birds, and all our cultivated plants, are more fertile than the corresponding species in a state of nature. It is no valid objection to this conclusion that animals suddenly supplied with an excess of food, or when grown very fat; and that most plants on sudden removal from very poor to very rich soil, are rendered more or less sterile. We might, therefore, expect that civilised men, who in one sense are highly domesticated, would be more prolific than wild men. It is also probable that the increased fertility of civilised nations would become, as with our domestic animals, an inherited character: it is at least known that with mankind a tendency to produce twins runs in families. (59. Mr. Sedgwick, 'British and Foreign Medico-Chirurgical Review,' July 1863, p. 170.)

Notwithstanding that savages appear to be less prolific than civilised people, they would no doubt rapidly increase if their numbers were not by some means rigidly kept down. The Santali, or hill-tribes of India, have recently afforded a good illustration of this fact; for, as shewn by Mr. Hunter (60. 'The Annals of Rural Bengal,' by W.W. Hunter, 1868, p. 259.), they have increased at an extraordinary rate since vaccination has been introduced, other pestilences mitigated, and war sternly repressed. This increase, however, would not have been possible had not these rude people spread into the adjoining districts, and worked for hire. Savages almost always marry; yet there is some prudential restraint, for they do not commonly marry at the earliest possible age. The young men are often required to shew that they can support a wife; and they generally have first to earn the price with which to purchase her from her parents. With savages the difficulty of obtaining subsistence occasionally limits their number in a much more direct manner than with civilised people, for all tribes periodically suffer from severe famines. At such times savages are forced to devour much bad food, and their health can hardly fail to be injured. Many accounts have been published of their protruding stomachs and emaciated limbs after and during famines. They are then, also, compelled to wander much, and, as I was assured in Australia, their infants perish in large numbers. As famines are periodical, depending chiefly on extreme seasons, all tribes must fluctuate in number. They cannot steadily and regularly increase, as there is no artificial increase in the supply of food. Savages, when hard pressed, encroach on each other's territories, and war is the result; but they are indeed almost always at war with their neighbours. They are liable to many accidents on land and water in their search for food; and in some countries they suffer much from the larger beasts of prey. Even in India, districts have been depopulated by the ravages of tigers.