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The Descent of Man and Selection in Relation to Sex
The Descent of Man and Selection in Relation to SexThe Descent of Man and Selection in Relation to Sex ....................................................... 2 Preface To The Second Edition .......................................................................................... 3 Introduction......................................................................................................................... 4 The Evidence Of The Descent Of Man From Some Lower Form...................................... 6 On The Manner Of Development Of Man From Some Lower Form............................... 20 Comparison Of The Mental Powers Of Man And The Lower Animals........................... 48 Comparison Of The Mental Powers Of Man And The Lower Animals—continued ....... 70On The Development Of The Intellectual And Moral Faculties During Primeval And Civilised Times ................................................................................................................. 91On The Affinities And Genealogy Of Man .................................................................... 104On The Races Of Man .................................................................................................... 118Principles Of Sexual Selection........................................................................................ 148Secondary Sexual Characters In The Lower Classes Of The Animal Kingdom ............ 187Secondary Sexual Characters Of Insects ........................................................................ 197Insects, continued............................................................................................................ 218Secondary Sexual Characters Of Fishes, Amphibians, And Reptiles............................. 235Secondary Sexual Characters Of Birds........................................................................... 252Birds—continued ............................................................................................................ 277Birds--continued. ............................................................................................................ 303Birds—concluded ........................................................................................................... 317Secondary Sexual Characters Of Mammals ................................................................... 344Secondary Sexual Characters Of Mammals--continued. ................................................ 360Secondary Sexual Characters Of Man ............................................................................ 377Secondary Sexual Characters Of Man—continued ........................................................ 397 General Summary And Conclusion ................................................................................ 412
Preface To The Second Edition
During the successive reprints of the first edition of this work, published in 1871, I was able to introduce several important corrections; and now that more time has elapsed, I have endeavoured to profit by the fiery ordeal through which the book has passed, and have taken advantage of all the criticisms which seem to me sound. I am also greatly indebted to a large number of correspondents for the communication of a surprising number of new facts and remarks. These have been so numerous, that I have been able to use only the more important ones; and of these, as well as of the more important corrections, I will append a list. Some new illustrations have been introduced, and four of the old drawings have been replaced by better ones, done from life by Mr. T.W. Wood. I must especially call attention to some observations which I owe to the kindness of Prof. Huxley (given as a supplement at the end of Part I.), on the nature of the differences between the brains of man and the higher apes. I have been particularly glad to give these observations, because during the last few years several memoirs on the subject have appeared on the Continent, and their importance has been, in some cases, greatly exaggerated by popular writers.
I may take this opportunity of remarking that my critics frequently assume that I attribute all changes of corporeal structure and mental power exclusively to the natural selection of such variations as are often called spontaneous; whereas, even in the first edition of the 'Origin of Species,' I distinctly stated that great weight must be attributed to the inherited effects of use and disuse, with respect both to the body and mind. I also attributed some amount of modification to the direct and prolonged action of changed conditions of life. Some allowance, too, must be made for occasional reversions of structure; nor must we forget what I have called "correlated" growth, meaning, thereby, that various parts of the organisation are in some unknown manner so connected, that when one part varies, so do others; and if variations in the one are accumulated by selection, other parts will be modified. Again, it has been said by several critics, that when I found that many details of structure in man could not be explained through natural selection, I invented sexual selection; I gave, however, a tolerably clear sketch of this principle in the first edition of the 'Origin of Species,' and I there stated that it was applicable to man. This subject of sexual selection has been treated at full length in the present work, simply because an opportunity was here first afforded me. I have been struck with the likeness of many of the half-favourable criticisms on sexual selection, with those which appeared at first on natural selection; such as, that it would explain some few details, but certainly was not applicable to the extent to which I have employed it. My conviction of the power of sexual selection remains unshaken; but it is probable, or almost certain, that several of my conclusions will hereafter be found erroneous; this can hardly fail to be the case in the first treatment of a subject. When naturalists have become familiar with the idea of sexual selection, it will, as I believe, be much more largely accepted; and it has already been fully and favourably received by several capable judges.DOWN, BECKENHAM, KENT, September, 1874. First Edition February 24, 1871. Second Edition September, 1874.
The nature of the following work will be best understood by a brief account of how it came to be written. During many years I collected notes on the origin or descent of man, without any intention of publishing on the subject, but rather with the determination not to publish, as I thought that I should thus only add to the prejudices against my views. It seemed to me sufficient to indicate, in the first edition of my 'Origin of Species,' that by this work "light would be thrown on the origin of man and his history;" and this implies that man must be included with other organic beings in any general conclusion respecting his manner of appearance on this earth. Now the case wears a wholly different aspect. When a naturalist like Carl Vogt ventures to say in his address as President of the National Institution of Geneva (1869), "personne, en Europe au moins, n'ose plus soutenir la creation independante et de toutes pieces, des especes," it is manifest that at least a large number of naturalists must admit that species are the modified descendants of other species; and this especially holds good with the younger and rising naturalists. The greater number accept the agency of natural selection; though some urge, whether with justice the future must decide, that I have greatly overrated its importance. Of the older and honoured chiefs in natural science, many unfortunately are still opposed to evolution in every form. In consequence of the views now adopted by most naturalists, and which will ultimately, as in every other case, be followed by others who are not scientific, I have been led to put together my notes, so as to see how far the general conclusions arrived at in my former works were applicable to man. This seemed all the more desirable, as I had never deliberately applied these views to a species taken singly. When we confine our attention to any one form, we are deprived of the weighty arguments derived from the nature of the affinities which connect together whole groups of organisms--their geographical distribution in past and present times, and their geological succession. The homological structure, embryological development, and rudimentary organs of a species remain to be considered, whether it be man or any other animal, to which our attention may be directed; but these great classes of facts afford, as it appears to me, ample and conclusive evidence in favour of the principle of gradual evolution. The strong support derived from the other arguments should, however, always be kept before the mind.
The sole object of this work is to consider, firstly, whether man, like every other species, is descended from some pre-existing form; secondly, the manner of his development; and thirdly, the value of the differences between the so-called races of man. As I shall confine myself to these points, it will not be necessary to describe in detail the differences between the several races--an enormous subject which has been fully described in many valuable works. The high antiquity of man has recently been demonstrated by the labours of a host of eminent men, beginning with M. Boucher de Perthes; and this is the indispensable basis for understanding his origin. I shall, therefore, take this conclusion for granted, and may refer my readers to the admirable treatises of Sir Charles Lyell, Sir John Lubbock, and others. Nor shall I have occasion to do more than to allude to the amount of difference between man and the anthropomorphous apes; for Prof. Huxley, in the opinion of most competent judges, has conclusively shewn that in every visible character man differs less from the higher apes, than these do from the lower members of the same order of Primates.
This work contains hardly any original facts in regard to man; but as the conclusions at which I arrived, after drawing up a rough draft, appeared to me interesting, I thought that they might interest others. It has often and confidently been asserted, that man's origin can never be known: but ignorance more frequently begets confidence than does knowledge: it is those who know little, and not those who know much, who so positively assert that this or that problem will never be solved by science. The conclusion that man is the co-descendant with other species of some ancient, lower, and extinct form, is not in any degree new. Lamarck long ago came to this conclusion, which has lately been maintained by several eminent naturalists and philosophers; for instance, by Wallace, Huxley, Lyell, Vogt, Lubbock, Buchner, Rolle, etc. (1. As the works of the first- named authors are so well known, I need not give the titles; but as those of the latter are less well known in England, I will give them:--'Sechs Vorlesungen uber die Darwin'sche Theorie:' zweite Auflage, 1868, von Dr L. Buchner; translated into French under the title 'Conferences sur la Theorie Darwinienne,' 1869. 'Der Mensch im Lichte der Darwin'sche Lehre,' 1865, von Dr. F. Rolle. I will not attempt to give references to all the authors who have taken the same side of the question. Thus G. Canestrini has published ('Annuario della Soc. d. Nat.,' Modena, 1867, page 81) a very curious paper on rudimentary characters, as bearing on the origin of man. Another work has (1869) been published by Dr. Francesco Barrago, bearing in Italian the title of "Man, made in the image of God, was also made in the image of the ape."), and especially by Haeckel. This last naturalist, besides his great work, 'Generelle Morphologie' (1866), has recently (1868, with a second edition in 1870), published his 'Naturliche Schopfungsgeschichte,' in which he fully discusses the genealogy of man. If this work had appeared before my essay had been written, I should probably never have completed it. Almost all the conclusions at which I have arrived I find confirmed by this naturalist, whose knowledge on many points is much fuller than mine. Wherever I have added any fact or view from Prof. Haeckel's writings, I give his authority in the text; other statements I leave as they originally stood in my manuscript, occasionally giving in the foot-notes references to his works, as a confirmation of the more doubtful or interesting points.
During many years it has seemed to me highly probable that sexual selection has played an important part in differentiating the races of man; but in my 'Origin of Species' (first edition, page 199) I contented myself by merely alluding to this belief. When I came to apply this view to man, I found it indispensable to treat the whole subject in full detail. (2. Prof. Haeckel was the only author who, at the time when this work first appeared, had discussed the subject of sexual selection, and had seen its full importance, since the publication of the 'Origin'; and this he did in a very able manner in his various works.) Consequently the second part of the present work, treating of sexual selection, has extended to an inordinate length, compared with the first part; but this could not be avoided. I had intended adding to the present volumes an essay on the expression of the various emotions by man and the lower animals. My attention was called to this subject many years ago by Sir Charles Bell's admirable work. This illustrious anatomist maintains that man is endowed with certain muscles solely for the sake of expressing his emotions. As this view is obviously opposed to the belief that man is descended from some other and lower form, it was necessary for me to consider it. I likewise wished to ascertain how far the emotions are expressed in the same manner by the different races of man. But owing to the length of the present work, I have thought it better to reserve my essay for separate publication.
The Evidence Of The Descent Of Man From Some Lower Form
Nature of the evidence bearing on the origin of man--Homologous structures in man and the lower animals--Miscellaneous points of correspondence-- Development-Rudimentary structures, muscles, sense-organs, hair, bones, reproductive organs, etc.-The bearing of these three great classes of facts on the origin of man.
He who wishes to decide whether man is the modified descendant of some pre- existing form, would probably first enquire whether man varies, however slightly, in bodily structure and in mental faculties; and if so, whether the variations are transmitted to his offspring in accordance with the laws which prevail with the lower animals. Again, are the variations the result, as far as our ignorance permits us to judge, of the same general causes, and are they governed by the same general laws, as in the case of other organisms; for instance, by correlation, the inherited effects of use and disuse, etc.? Is man subject to similar malconformations, the result of arrested development, of reduplication of parts, etc., and does he display in any of his anomalies reversion to some former and ancient type of structure? It might also naturally be enquired whether man, like so many other animals, has given rise to varieties and sub-races, differing but slightly from each other, or to races differing so much that they must be classed as doubtful species? How are such races distributed over the world; and how, when crossed, do they react on each other in the first and succeeding generations? And so with many other points.
The enquirer would next come to the important point, whether man tends to increase at so rapid a rate, as to lead to occasional severe struggles for existence; and consequently to beneficial variations, whether in body or mind, being preserved, and injurious ones eliminated. Do the races or species of men, whichever term may be applied, encroach on and replace one another, so that some finally become extinct? We shall see that all these questions, as indeed is obvious in respect to most of them, must be answered in the affirmative, in the same manner as with the lower animals. But the several considerations just referred to may be conveniently deferred for a time: and we will first see how far the bodily structure of man shews traces, more or less plain, of his descent from some lower form. In succeeding chapters the mental powers of man, in comparison with those of the lower animals, will be considered.THE BODILY STRUCTURE OF MAN.
It is notorious that man is constructed on the same general type or model as other mammals. All the bones in his skeleton can be compared with corresponding bones in a monkey, bat, or seal. So it is with his muscles, nerves, blood-vessels and internal viscera. The brain, the most important of all the organs, follows the same law, as shewn by Huxley and other anatomists. Bischoff (1. 'Grosshirnwindungen des Menschen,' 1868, s. 96. The conclusions of this author, as well as those of Gratiolet and Aeby, concerning the brain, will be discussed by Prof. Huxley in the Appendix alluded to in the Preface to this edition.), who is a hostile witness, admits that every chief fissure and fold in the brain of man has its analogy in that of the orang; but he adds that at no period of development do their brains perfectly agree; nor could perfect agreement be expected, for otherwise their mental powers would have been the same. Vulpian (2. 'Lec. sur la Phys.' 1866, page 890, as quoted by M. Dally, 'L'Ordre des Primates et le Transformisme,' 1868, page 29.), remarks: "Les differences reelles qui existent entre l'encephale de l'homme et celui des singes superieurs, sont bien minimes. Il ne faut pas se faire d'illusions a cet egard. L'homme est bien plus pres des singes anthropomorphes par les caracteres anatomiques de son cerveau que ceux-ci ne le sont non seulement des autres mammiferes, mais meme de certains quadrumanes, des guenons et des macaques." But it would be superfluous here to give further details on the correspondence between man and the higher mammals in the structure of the brain and all other parts of the body.It may, however, be worth while to specify a few points, not directly or obviously connected with structure, by which this correspondence or relationship is well shewn.
Man is liable to receive from the lower animals, and to communicate to them, certain diseases, as hydrophobia, variola, the glanders, syphilis, cholera, herpes, etc. (3. Dr. W. Lauder Lindsay has treated this subject at some length in the 'Journal of Mental Science,' July 1871; and in the 'Edinburgh Veterinary Review,' July 1858.); and this fact proves the close similarity (4. A Reviewer has criticised ('British Quarterly Review,' Oct. 1st, 1871, page 472) what I have here said with much severity and contempt; but as I do not use the term identity, I cannot see that I am greatly in error. There appears to me a strong analogy between the same infection or contagion producing the same result, or one closely similar, in two distinct animals, and the testing of two distinct fluids by the same chemical reagent.) of their tissues and blood, both in minute structure and composition, far more plainly than does their comparison under the best microscope, or by the aid of the best chemical analysis. Monkeys are liable to many of the same non-contagious diseases as we are; thus Rengger (5. 'Naturgeschichte der Saugethiere von Paraguay,' 1830, s. 50.), who carefully observed for a long time the Cebus Azarae in its native land, found it liable to catarrh, with the usual symptoms, and which, when often recurrent, led to consumption. These monkeys suffered also from apoplexy, inflammation of the bowels, and cataract in the eye. The younger ones when shedding their milk-teeth often died from fever. Medicines produced the same effect on them as on us. Many kinds of monkeys have a strong taste for tea, coffee, and spiritous liquors: they will also, as I have myself seen, smoke tobacco with pleasure. (6. The same tastes are common to some animals much lower in the scale. Mr. A. Nichols informs me that he kept in Queensland, in Australia, three individuals of the Phaseolarctus cinereus; and that, without having been taught in any way, they acquired a strong taste for rum, and for smoking tobacco.) Brehm asserts that the natives of north-eastern Africa catch the wild baboons by exposing vessels with strong beer, by which they are made drunk. He has seen some of these animals, which he kept in confinement, in this state; and he gives a laughable account of their behaviour and strange grimaces. On the following morning they were very cross and dismal; they held their aching heads with both hands, and wore a most pitiable expression: when beer or wine was offered them, they turned away with disgust, but relished the juice of lemons. (7. Brehm, 'Thierleben,' B. i. 1864, s. 75, 86. On the Ateles, s. 105. For other analogous statements, see s. 25, 107.) An American monkey, an Ateles, after getting drunk on brandy, would never touch it again, and thus was wiser than many men. These trifling facts prove how similar the nerves of taste must be in monkeys and man, and how similarly their whole nervous system is affected.
Man is infested with internal parasites, sometimes causing fatal effects; and is plagued by external parasites, all of which belong to the same genera or families as those infesting other mammals, and in the case of scabies to the same species. (8. Dr. W. Lauder Lindsay, 'Edinburgh Vet. Review,' July 1858, page 13.) Man is subject, like other mammals, birds, and even insects (9. With respect to insects see Dr. Laycock, "On a General Law of Vital Periodicity," 'British Association,' 1842. Dr. Macculloch, 'Silliman's North American Journal of Science,' vol. XVII. page 305, has seen a dog suffering from tertian ague. Hereafter I shall return to this subject.), to that mysterious law, which causes certain normal processes, such as gestation, as well as the maturation and duration of various diseases, to follow lunar periods. His wounds are repaired by the same process of healing; and the stumps left after the amputation of his limbs, especially during an early embryonic period, occasionally possess some power of regeneration, as in the lowest animals. (10. I have given the evidence on this head in my 'Variation of Animals and Plants under Domestication,' vol. ii. page 15, and more could be added.)
The whole process of that most important function, the reproduction of the species, is strikingly the same in all mammals, from the first act of courtship by the male (11. Mares e diversis generibus Quadrumanorum sine dubio dignoscunt feminas humanas a maribus. Primum, credo, odoratu, postea aspectu. Mr. Youatt, qui diu in Hortis Zoologicis (Bestiariis) medicus animalium erat, vir in rebus observandis cautus et sagax, hoc mihi certissime probavit, et curatores ejusdem loci et alii e ministris confirmaverunt. Sir Andrew Smith et Brehm notabant idem in Cynocephalo. Illustrissimus Cuvier etiam narrat multa de hac re, qua ut opinor, nihil turpius potest indicari inter omnia hominibus et Quadrumanis communia. Narrat enim Cynocephalum quendam in furorem incidere aspectu feminarum aliquarem, sed nequaquam accendi tanto furore ab omnibus. Semper eligebat juniores, et dignoscebat in turba, et advocabat voce gestuque.), to the birth and nurturing of the young. Monkeys are born in almost as helpless a condition as our own infants; and in certain genera the young differ fully as much in appearance from the adults, as do our children from their full-grown parents. (12. This remark is made with respect to Cynocephalus and the anthropomorphous apes by Geoffroy Saint-Hilaire and F. Cuvier, 'Histoire Nat. des Mammiferes,' tom. i. 1824.) It has been urged by some writers, as an important distinction, that with man the young arrive at maturity at a much later age than with any other animal: but if we look to the races of mankind which inhabit tropical countries the difference is not great, for the orang is believed not to be adult till the age of from ten to fifteen years. (13. Huxley, 'Man's Place in Nature,' 1863, p. 34.) Man differs from woman in size, bodily strength, hairiness, etc., as well as in mind, in the same manner as do the two sexes of many mammals. So that the correspondence in general structure, in the minute structure of the tissues, in chemical composition and in constitution, between man and the higher animals, especially the anthropomorphous apes, is extremely close.EMBRYONIC DEVELOPMENT. [Fig. 1. Shows a human embryo, from Ecker, and a dog embryo, from Bischoff. Labelled in each are:
a. Fore-brain, cerebral hemispheres, etc.
b. Mid-brain, corpora quadrigemina.
c. Hind-brain, cerebellum, medulla oblongata.
f. First visceral arch.
g. Second visceral arch.
H. Vertebral columns and muscles in process of development. i. Anterior extremities.
K. Posterior extremities.
Man is developed from an ovule, about the 125th of an inch in diameter, which differs in no respect from the ovules of other animals. The embryo itself at a very early period can hardly be distinguished from that of other members of the vertebrate kingdom. At this period the arteries run in arch-like branches, as if to carry the blood to branchiae which are not present in the higher Vertebrata, though the slits on the sides of the neck still remain (see f, g, fig. 1), marking their former position. At a somewhat later period, when the extremities are developed, "the feet of lizards and mammals," as the illustrious Von Baer remarks, "the wings and feet of birds, no less than the hands and feet of man, all arise from the same fundamental form." It is, says Prof. Huxley (14. 'Man's Place in Nature,' 1863, p. 67.), "quite in the later stages of development that the young human being presents marked differences from the young ape, while the latter departs as much from the dog in its developments, as the man does. Startling as this last assertion may appear to be, it is demonstrably true."
As some of my readers may never have seen a drawing of an embryo, I have given one of man and another of a dog, at about the same early stage of development, carefully copied from two works of undoubted accuracy. (15. The human embryo (upper fig.) is from Ecker, 'Icones Phys.,' 1851-1859, tab. xxx. fig. 2. This embryo was ten lines in length, so that the drawing is much magnified. The embryo of the dog is from Bischoff, 'Entwicklungsgeschichte des Hunde-Eies,' 1845, tab. xi. fig. 42B. This drawing is five times magnified, the embryo being twenty-five days old. The internal viscera have been omitted, and the uterine appendages in both drawings removed. I was directed to these figures by Prof. Huxley, from whose work, 'Man's Place in Nature,' the idea of giving them was taken. Haeckel has also given analogous drawings in his 'Schopfungsgeschichte.')
After the foregoing statements made by such high authorities, it would be superfluous on my part to give a number of borrowed details, shewing that the embryo of man closely resembles that of other mammals. It may, however, be added, that the human embryo likewise resembles certain low forms when adult in various points of structure. For instance, the heart at first exists as a simple pulsating vessel; the excreta are voided through a cloacal passage; and the os coccyx projects like a true tail, "extending considerably beyond the rudimentary legs." (16. Prof. Wyman in 'Proceedings of the American Academy of Sciences,' vol. iv. 1860, p. 17.) In the embryos of all air-breathing vertebrates, certain glands, called the corpora Wolffiana, correspond with, and act like the kidneys of mature fishes. (17. Owen, 'Anatomy of Vertebrates,' vol. i. p. 533.) Even at a later embryonic period, some striking resemblances between man and the lower animals may be observed. Bischoff says that "the convolutions of the brain in a human foetus at the end of the seventh month reach about the same stage of development as in a baboon when adult." (18. 'Die Grosshirnwindungen des Menschen,' 1868, s. 95.) The great toe, as Professor Owen remarks (19. 'Anatomy of Vertebrates,' vol. ii. p. 553.), "which forms the fulcrum when standing or walking, is perhaps the most characteristic peculiarity in the human structure;" but in an embryo, about an inch in length, Prof. Wyman (20. 'Proc. Soc. Nat. Hist.' Boston, 1863, vol. ix. p. 185.) found "that the great toe was shorter than the others; and, instead of being parallel to them, projected at an angle from the side of the foot, thus corresponding with the permanent condition of this part in the quadrumana." I will conclude with a quotation from Huxley (21. 'Man's Place in Nature,' p. 65.) who after asking, does man originate in a different way from a dog, bird, frog or fish? says, "the reply is not doubtful for a moment; without question, the mode of origin, and the early stages of the development of man, are identical with those of the animals immediately below him in the scale: without a doubt in these respects, he is far nearer to apes than the apes are to the dog."RUDIMENTS.
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 'Schopfungsgeschichte.') 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 semihuman 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 ihabiting 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. 'Uber 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 ('Etudes sur les Facultes 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, Uber die Richtung der Haare am menschlichen Korper, 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 thirtyfive, 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'Unite 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.)
In some of the lower Quadrumana, in the Lemuridae and Carnivora, as well as in many marsupials, there is a passage near the lower end of the humerus, called the supracondyloid foramen, through which the great nerve of the fore limb and often the great artery pass. Now in the humerus of man, there is generally a trace of this passage, which is sometimes fairly well developed, being formed by a depending hook-like process of bone, completed by a band of ligament. Dr. Struthers (49. With respect to inheritance, see Dr. Struthers in the 'Lancet,' Feb. 15, 1873, and another important paper, ibid. Jan. 24, 1863, p. 83. Dr. Knox, as I am informed, was the first anatomist who drew attention to this peculiar structure in man; see his 'Great Artists and Anatomists,' p. 63. See also an important memoir on this process by Dr. Gruber, in the 'Bulletin de l'Acad. Imp. de St. Petersbourg,' tom. xii. 1867, p. 448.), who has closely attended to the subject, has now shewn that this peculiarity is sometimes inherited, as it has occurred in a father, and in no less than four out of his seven children. When present, the great nerve invariably passes through it; and this clearly indicates that it is the homologue and rudiment of the supracondyloid foramen of the lower animals. Prof. Turner estimates, as he informs me, that it occurs in about one per cent. of recent skeletons. But if the occasional development of this structure in man is, as seems probable, due to reversion, it is a return to a very ancient state of things, because in the higher Quadrumana it is absent.
There is another foramen or perforation in the humerus, occasionally present in man, which may be called the inter-condyloid. This occurs, but not constantly, in various anthropoid and other apes (50. Mr. St. George Mivart, 'Transactions Phil. Soc.' 1867, p. 310.), and likewise in many of the lower animals. It is remarkable that this perforation seems to have been present in man much more frequently during ancient times than recently. Mr. Busk (51. "On the Caves of Gibraltar," 'Transactions of the International Congress of Prehistoric Archaeology,' Third Session, 1869, p. 159. Prof. Wyman has lately shewn (Fourth Annual Report, Peabody Museum, 1871, p. 20), that this perforation is present in thirty-one per cent. of some human remains from ancient mounds in the Western United States, and in Florida. It frequently occurs in the negro.) has collected the following evidence on this head: Prof. Broca "noticed the perforation in four and a half per cent. of the arm-bones collected in the 'Cimetiere du Sud,' at Paris; and in the Grotto of Orrony, the contents of which are referred to the Bronze period, as many as eight humeri out of thirty-two were perforated; but this extraordinary proportion, he thinks, might be due to the cavern having been a sort of 'family vault.' Again, M. Dupont found thirty per cent. of perforated bones in the caves of the Valley of the Lesse, belonging to the Reindeer period; whilst M. Leguay, in a sort of dolmen at Argenteuil, observed twenty-five per cent. to be perforated; and M. Pruner-Bey found twenty-six per cent. in the same condition in bones from Vaureal. Nor should it be left unnoticed that M. PrunerBey states that this condition is common in Guanche skeletons." It is an interesting fact that ancient races, in this and several other cases, more frequently present structures which resemble those of the lower animals than do the modern. One chief cause seems to be that the ancient races stand somewhat nearer in the long line of descent to their remote animal-like progenitors.
In man, the os coccyx, together with certain other vertebrae hereafter to be described, though functionless as a tail, plainly represent this part in other vertebrate animals. At an early embryonic period it is free, and projects beyond the lower extremities; as may be seen in the drawing (Fig. 1.) of a human embryo. Even after birth it has been known, in certain rare and anomalous cases (52. Quatrefages has lately collected the evidence on this subject. 'Revue des Cours Scientifiques,' 1867-1868, p. 625. In 1840 Fleischmann exhibited a human foetus bearing a free tail, which, as is not always the case, included vertebral bodies; and this tail was critically examined by the many anatomists present at the meeting of naturalists at Erlangen (see Marshall in Niederlandischen Archiv fur Zoologie, December 1871).), to form a small external rudiment of a tail. The os coccyx is short, usually including only four vertebrae, all anchylosed together: and these are in a rudimentary condition, for they consist, with the exception of the basal one, of the centrum alone. (53. Owen, 'On the Nature of Limbs,' 1849, p. 114.) They are furnished with some small muscles; one of which, as I am informed by Prof. Turner, has been expressly described by Theile as a rudimentary repetition of the extensor of the tail, a muscle which is so largely developed in many mammals.
The spinal cord in man extends only as far downwards as the last dorsal or first lumbar vertebra; but a thread-like structure (the filum terminale) runs down the axis of the sacral part of the spinal canal, and even along the back of the coccygeal bones. The upper part of this filament, as Prof. Turner informs me, is undoubtedly homologous with the spinal cord; but the lower part apparently consists merely of the pia mater, or vascular investing membrane. Even in this case the os coccyx may be said to possess a vestige of so important a structure as the spinal cord, though no longer enclosed within a bony canal. The following fact, for which I am also indebted to Prof. Turner, shews how closely the os coccyx corresponds with the true tail in the lower animals: Luschka has recently discovered at the extremity of the coccygeal bones a very peculiar convoluted body, which is continuous with the middle sacral artery; and this discovery led Krause and Meyer to examine the tail of a monkey (Macacus), and of a cat, in both of which they found a similarly convoluted body, though not at the extremity.
The reproductive system offers various rudimentary structures; but these differ in one important respect from the foregoing cases. Here we are not concerned with the vestige of a part which does not belong to the species in an efficient state, but with a part efficient in the one sex, and represented in the other by a mere rudiment. Nevertheless, the occurrence of such rudiments is as difficult to explain, on the belief of the separate creation of each species, as in the foregoing cases. Hereafter I shall have to recur to these rudiments, and shall shew that their presence generally depends merely on inheritance, that is, on parts acquired by one sex having been partially transmitted to the other. I will in this place only give some instances of such rudiments. It is well known that in the males of all mammals, including man, rudimentary mammae exist. These in several instances have become well developed, and have yielded a copious supply of milk. Their essential identity in the two sexes is likewise shewn by their occasional sympathetic enlargement in both during an attack of the measles. The vesicula prostatica, which has been observed in many male mammals, is now universally acknowledged to be the homologue of the female uterus, together with the connected passage. It is impossible to read Leuckart's able description of this organ, and his reasoning, without admitting the justness of his conclusion. This is especially clear in the case of those mammals in which the true female uterus bifurcates, for in the males of these the vesicula likewise bifurcates. (54. Leuckart, in Todd's 'Cyclopaedia of Anatomy' 1849-52, vol. iv. p. 1415. In man this organ is only from three to six lines in length, but, like so many other rudimentary parts, it is variable in development as well as in other characters.) Some other rudimentary structures belonging to the reproductive system might have been here adduced. (55. See, on this subject, Owen, 'Anatomy of Vertebrates,' vol. iii. pp. 675, 676, 706.)
The bearing of the three great classes of facts now given is unmistakeable. But it would be superfluous fully to recapitulate the line of argument given in detail in my 'Origin of Species.' The homological construction of the whole frame in the members of the same class is intelligible, if we admit their descent from a common progenitor, together with their subsequent adaptation to diversified conditions. On any other view, the similarity of pattern between the hand of a man or monkey, the foot of a horse, the flipper of a seal, the wing of a bat, etc., is utterly inexplicable. (56. Prof. Bianconi, in a recently published work, illustrated by admirable engravings ('La Theorie Darwinienne et la creation dite independante,' 1874), endeavours to shew that homological structures, in the above and other cases, can be fully explained on mechanical principles, in accordance with their uses. No one has shewn so well, how admirably such structures are adapted for their final purpose; and this adaptation can, as I believe, be explained through natural selection. In considering the wing of a bat, he brings forward (p. 218) what appears to me (to use Auguste Comte's words) a mere metaphysical principle, namely, the preservation "in its integrity of the mammalian nature of the animal." In only a few cases does he discuss rudiments, and then only those parts which are partially rudimentary, such as the little hoofs of the pig and ox, which do not touch the ground; these he shews clearly to be of service to the animal. It is unfortunate that he did not consider such cases as the minute teeth, which never cut through the jaw in the ox, or the mammae of male quadrupeds, or the wings of certain beetles, existing under the soldered wing-covers, or the vestiges of the pistil and stamens in various flowers, and many other such cases. Although I greatly admire Prof. Bianconi's work, yet the belief now held by most naturalists seems to me left unshaken, that homological structures are inexplicable on the principle of mere adaptation.) It is no scientific explanation to assert that they have all been formed on the same ideal plan. With respect to development, we can clearly understand, on the principle of variations supervening at a rather late embryonic period, and being inherited at a corresponding period, how it is that the embryos of wonderfully different forms should still retain, more or less perfectly, the structure of their common progenitor. No other explanation has ever been given of the marvellous fact that the embryos of a man, dog, seal, bat, reptile, etc., can at first hardly be distinguished from each other. In order to understand the existence of rudimentary organs, we have only to suppose that a former progenitor possessed the parts in question in a perfect state, and that under changed habits of life they became greatly reduced, either from simple disuse, or through the natural selection of those individuals which were least encumbered with a superfluous part, aided by the other means previously indicated.
Thus we can understand how it has come to pass that man and all other vertebrate animals have been constructed on the same general model, why they pass through the same early stages of development, and why they retain certain rudiments in common. Consequently we ought frankly to admit their community of descent: to take any other view, is to admit that our own structure, and that of all the animals around us, is a mere snare laid to entrap our judgment. This conclusion is greatly strengthened, if we look to the members of the whole animal series, and consider the evidence derived from their affinities or classification, their geographical distribution and geological succession. It is only our natural prejudice, and that arrogance which made our forefathers declare that they were descended from demi-gods, which leads us to demur to this conclusion. But the time will before long come, when it will be thought wonderful that naturalists, who were well acquainted with the comparative structure and development of man, and other mammals, should have believed that each was the work of a separate act of creation.
On The Manner Of Development Of Man From Some Lower Form
Variability of body and mind in man--Inheritance--Causes of variability-- Laws of variation the same in man as in the lower animals--Direct action of the conditions of life-Effects of the increased use and disuse of parts-- Arrested development--Reversion-Correlated variation--Rate of increase-- Checks to increase--Natural selection--Man the most dominant animal in the world--Importance of his corporeal structure--The causes which have led to his becoming erect--Consequent changes of structure--Decrease in size of the canine teeth--Increased size and altered shape of the skull--Nakedness --Absence of a tail--Defenceless condition of man.
It is manifest that man is now subject to much variability. No two individuals of the same race are quite alike. We may compare millions of faces, and each will be distinct. There is an equally great amount of diversity in the proportions and dimensions of the various parts of the body; the length of the legs being one of the most variable points. (1. 'Investigations in Military and Anthropological Statistics of American Soldiers,' by B.A. Gould, 1869, p. 256.) Although in some quarters of the world an elongated skull, and in other quarters a short skull prevails, yet there is great diversity of shape even within the limits of the same race, as with the aborigines of America and South Australia--the latter a race "probably as pure and homogeneous in blood, customs, and language as any in existence"--and even with the inhabitants of so confined an area as the Sandwich Islands. (2. With respect to the "Cranial forms of the American aborigines," see Dr. Aitken Meigs in 'Proc. Acad. Nat. Sci.' Philadelphia, May 1868. On the Australians, see Huxley, in Lyell's 'Antiquity of Man,' 1863, p. 87. On the Sandwich Islanders, Prof. J. Wyman, 'Observations on Crania,' Boston, 1868, p. 18.) An eminent dentist assures me that there is nearly as much diversity in the teeth as in the features. The chief arteries so frequently run in abnormal courses, that it has been found useful for surgical purposes to calculate from 1040 corpses how often each course prevails. (3. 'Anatomy of the Arteries,' by R. Quain. Preface, vol. i. 1844.) The muscles are eminently variable: thus those of the foot were found by Prof. Turner (4. 'Transactions of the Royal Society of Edinburgh,' vol. xxiv. pp. 175, 189.) not to be strictly alike in any two out of fifty bodies; and in some the deviations were considerable. He adds, that the power of performing the appropriate movements must have been modified in accordance with the several deviations. Mr. J. Wood has recorded (5. 'Proceedings Royal Society,' 1867, p. 544; also 1868, pp. 483, 524. There is a previous paper, 1866, p. 229.) the occurrence of 295 muscular variations in thirty-six subjects, and in another set of the same number no less than 558 variations, those occurring on both sides of the body being only reckoned as one. In the last set, not one body out of the thirty-six was "found totally wanting in departures from the standard descriptions of the muscular system given in anatomical text books." A single body presented the extraordinary number of twenty-five distinct abnormalities. The same muscle sometimes varies in many ways: thus Prof. Macalister describes (6. 'Proc. R. Irish Academy,' vol. x. 1868, p. 141.) no less than twenty distinct variations in the palmaris accessorius.
The famous old anatomist, Wolff (7. 'Act. Acad. St. Petersburg,' 1778, part ii. p. 217.), insists that the internal viscera are more variable than the external parts: Nulla particula est quae non aliter et aliter in aliis se habeat hominibus. He has even written a treatise on the choice of typical examples of the viscera for representation. A discussion on the beauideal of the liver, lungs, kidneys, etc., as of the human face divine, sounds strange in our ears.
The variability or diversity of the mental faculties in men of the same race, not to mention the greater differences between the men of distinct races, is so notorious that not a word need here be said. So it is with the lower animals. All who have had charge of menageries admit this fact, and we see it plainly in our dogs and other domestic animals. Brehm especially insists that each individual monkey of those which he kept tame in Africa had its own peculiar disposition and temper: he mentions one baboon remarkable for its high intelligence; and the keepers in the Zoological Gardens pointed out to me a monkey, belonging to the New World division, equally remarkable for intelligence. Rengger, also, insists on the diversity in the various mental characters of the monkeys of the same species which he kept in Paraguay; and this diversity, as he adds, is partly innate, and partly the result of the manner in which they have been treated or educated. (8. Brehm, 'Thierleben,' B. i. ss. 58, 87. Rengger, 'Saugethiere von Paraguay,' s. 57.)
I have elsewhere (9. 'Variation of Animals and Plants under Domestication,' vol. ii. chap. xii.) so fully discussed the subject of Inheritance, that I need here add hardly anything. A greater number of facts have been collected with respect to the transmission of the most trifling, as well as of the most important characters in man, than in any of the lower animals; though the facts are copious enough with respect to the latter. So in regard to mental qualities, their transmission is manifest in our dogs, horses, and other domestic animals. Besides special tastes and habits, general intelligence, courage, bad and good temper, etc., are certainly transmitted. With man we see similar facts in almost every family; and we now know, through the admirable labours of Mr. Galton (10. 'Hereditary Genius: an Inquiry into its Laws and Consequences,' 1869.), that genius which implies a wonderfully complex combination of high faculties, tends to be inherited; and, on the other hand, it is too certain that insanity and deteriorated mental powers likewise run in families.
With respect to the causes of variability, we are in all cases very ignorant; but we can see that in man as in the lower animals, they stand in some relation to the conditions to which each species has been exposed, during several generations. Domesticated animals vary more than those in a state of nature; and this is apparently due to the diversified and changing nature of the conditions to which they have been subjected. In this respect the different races of man resemble domesticated animals, and so do the individuals of the same race, when inhabiting a very wide area, like that of America. We see the influence of diversified conditions in the more civilised nations; for the members belonging to different grades of rank, and following different occupations, present a greater range of character than do the members of barbarous nations. But the uniformity of savages has often been exaggerated, and in some cases can hardly be said to exist. (11. Mr. Bates remarks ('The Naturalist on the Amazons,' 1863, vol. ii p. 159), with respect to the Indians of the same South American tribe, "no two of them were at all similar in the shape of the head; one man had an oval visage with fine features, and another was quite Mongolian in breadth and prominence of cheek, spread of nostrils, and obliquity of eyes.") It is, nevertheless, an error to speak of man, even if we look only to the conditions to which he has been exposed, as "far more domesticated" (12. Blumenbach, 'Treatises on Anthropology.' Eng. translat., 1865, p. 205.) than any other animal. Some savage races, such as the Australians, are not exposed to more diversified conditions than are many species which have a wide range. In another and much more important respect, man differs widely from any strictly domesticated animal; for his breeding has never long been controlled, either by methodical or unconscious selection. No race or body of men has been so completely subjugated by other men, as that certain individuals should be preserved, and thus unconsciously selected, from somehow excelling in utility to their masters. Nor have certain male and female individuals been intentionally picked out and matched, except in the well-known case of the Prussian grenadiers; and in this case man obeyed, as might have been expected, the law of methodical selection; for it is asserted that many tall men were reared in the villages inhabited by the grenadiers and their tall wives. In Sparta, also, a form of selection was followed, for it was enacted that all children should be examined shortly after birth; the well-formed and vigorous being preserved, the others left to perish. (13. Mitford's 'History of Greece,' vol. i. p. 282. It appears also from a passage in Xenophon's 'Memorabilia,' B. ii. 4 (to which my attention has been called by the Rev. J.N. Hoare), that it was a well recognised principle with the Greeks, that men ought to select their wives with a view to the health and vigour of their children. The Grecian poet, Theognis, who lived 550 B.C., clearly saw how important selection, if carefully applied, would be for the improvement of mankind. He saw, likewise, that wealth often checks the proper action of sexual selection. He thus writes:
"With kine and horses, Kurnus! we proceed By reasonable rules, and choose a breed For profit and increase, at any price:
Of a sound stock, without defect or vice. But, in the daily matches that we make, The price is everything: for money's sake, Men marry: women are in marriage given The churl or ruffian, that in wealth has thriven, May match his offspring with the proudest race: Thus everything is mix'd, noble and base! If then in outward manner, form, and mind, You find us a degraded, motley kind,
Wonder no more, my friend! the cause is plain, And to lament the consequence is vain."
If we consider all the races of man as forming a single species, his range is enormous; but some separate races, as the Americans and Polynesians, have very wide ranges. It is a well-known law that widely-ranging species are much more variable than species with restricted ranges; and the variability of man may with more truth be compared with that of widely- ranging species, than with that of domesticated animals.
Not only does variability appear to be induced in man and the lower animals by the same general causes, but in both the same parts of the body are affected in a closely analogous manner. This has been proved in such full detail by Godron and Quatrefages, that I need here only refer to their works. (14. Godron, 'De l'Espece,' 1859, tom. ii. livre 3. Quatrefages, 'Unite de l'Espece Humaine,' 1861. Also Lectures on Anthropology, given in the 'Revue des Cours Scientifiques,' 1866-1868.) Monstrosities, which graduate into slight variations, are likewise so similar in man and the lower animals, that the same classification and the same terms can be used for both, as has been shewn by Isidore Geoffroy St.-Hilaire. (15. 'Hist. Gen. et Part. des Anomalies de l'Organisation,' in three volumes, tom. i. 1832.) In my work on the variation of domestic animals, I have attempted to arrange in a rude fashion the laws of variation under the following heads:-The direct and definite action of changed conditions, as exhibited by all or nearly all the individuals of the same species, varying in the same manner under the same circumstances. The effects of the long- continued use or disuse of parts. The cohesion of homologous parts. The variability of multiple parts. Compensation of growth; but of this law I have found no good instance in the case of man. The effects of the mechanical pressure of one part on another; as of the pelvis on the cranium of the infant in the womb. Arrests of development, leading to the diminution or suppression of parts. The reappearance of long-lost characters through reversion. And lastly, correlated variation. All these so-called laws apply equally to man and the lower animals; and most of them even to plants. It would be superfluous here to discuss all of them (16. I have fully discussed these laws in my 'Variation of Animals and Plants under Domestication,' vol. ii. chap. xxii. and xxiii. M. J.P. Durand has lately (1868) published a valuable essay, 'De l'Influence des Milieux,' etc. He lays much stress, in the case of plants, on the nature of the soil.); but several are so important, that they must be treated at considerable length.THE DIRECT AND DEFINITE ACTION OF CHANGED CONDITIONS.
This is a most perplexing subject. It cannot be denied that changed conditions produce some, and occasionally a considerable effect, on organisms of all kinds; and it seems at first probable that if sufficient time were allowed this would be the invariable result. But I have failed to obtain clear evidence in favour of this conclusion; and valid reasons may be urged on the other side, at least as far as the innumerable structures are concerned, which are adapted for special ends. There can, however, be no doubt that changed conditions induce an almost indefinite amount of fluctuating variability, by which the whole organisation is rendered in some degree plastic.
In the United States, above 1,000,000 soldiers, who served in the late war, were measured, and the States in which they were born and reared were recorded. (17. 'Investigations in Military and Anthrop. Statistics,' etc., 1869, by B.A. Gould, pp. 93, 107, 126, 131, 134.) From this astonishing number of observations it is proved that local influences of some kind act directly on stature; and we further learn that "the State where the physical growth has in great measure taken place, and the State of birth, which indicates the ancestry, seem to exert a marked influence on the stature." For instance, it is established, "that residence in the Western States, during the years of growth, tends to produce increase of stature." On the other hand, it is certain that with sailors, their life delays growth, as shewn "by the great difference between the statures of soldiers and sailors at the ages of seventeen and eighteen years." Mr. B.A. Gould endeavoured to ascertain the nature of the influences which thus act on stature; but he arrived only at negative results, namely that they did not relate to climate, the elevation of the land, soil, nor even "in any controlling degree" to the abundance or the need of the comforts of life. This latter conclusion is directly opposed to that arrived at by Villerme, from the statistics of the height of the conscripts in different parts of France. When we compare the differences in stature between the Polynesian chiefs and the lower orders within the same islands, or between the inhabitants of the fertile volcanic and low barren coral islands of the same ocean (18. For the Polynesians, see Prichard's 'Physical History of Mankind,' vol. v. 1847, pp. 145, 283. Also Godron, 'De l'Espece,' tom. ii. p. 289. There is also a remarkable difference in appearance between the closely-allied Hindoos inhabiting the Upper Ganges and Bengal; see Elphinstone's 'History of India,' vol. i. p. 324.) or again between the Fuegians on the eastern and western shores of their country, where the means of subsistence are very different, it is scarcely possible to avoid the conclusion that better food and greater comfort do influence stature. But the preceding statements shew how difficult it is to arrive at any precise result. Dr. Beddoe has lately proved that, with the inhabitants of Britain, residence in towns and certain occupations have a deteriorating influence on height; and he infers that the result is to a certain extent inherited, as is likewise the case in the United States. Dr. Beddoe further believes that wherever a "race attains its maximum of physical development, it rises highest in energy and moral vigour." (19. 'Memoirs, Anthropological Society,' vol. iii. 1867-69, pp. 561, 565, 567.)
Whether external conditions produce any other direct effect on man is not known. It might have been expected that differences of climate would have had a marked influence, inasmuch as the lungs and kidneys are brought into activity under a low temperature, and the liver and skin under a high one. (20. Dr. Brakenridge, 'Theory of Diathesis,' 'Medical Times,' June 19 and July 17, 1869.) It was formerly thought that the colour of the skin and the character of the hair were determined by light or heat; and although it can hardly be denied that some effect is thus produced, almost all observers now agree that the effect has been very small, even after exposure during many ages. But this subject will be more properly discussed when we treat of the different races of mankind. With our domestic animals there are grounds for believing that cold and damp directly affect the growth of the hair; but I have not met with any evidence on this head in the case of man.EFFECTS OF THE INCREASED USE AND DISUSE OF PARTS.
It is well known that use strengthens the muscles in the individual, and complete disuse, or the destruction of the proper nerve, weakens them. When the eye is destroyed, the optic nerve often becomes atrophied. When an artery is tied, the lateral channels increase not only in diameter, but in the thickness and strength of their coats. When one kidney ceases to act from disease, the other increases in size, and does double work. Bones increase not only in thickness, but in length, from carrying a greater weight. (21. I have given authorities for these several statements in my 'Variation of Animals and Plants under Domestication,' vol. ii. pp. 297- 300. Dr. Jaeger, "Uber das Langenwachsthum der Knochen," 'Jenaischen Zeitschrift,' B. v. Heft. i.) Different occupations, habitually followed, lead to changed proportions in various parts of the body. Thus it was ascertained by the United States Commission (22. 'Investigations,' etc., by B.A. Gould, 1869, p. 288.) that the legs of the sailors employed in the late war were longer by 0.217 of an inch than those of the soldiers, though the sailors were on an average shorter men; whilst their arms were shorter by 1.09 of an inch, and therefore, out of proportion, shorter in relation to their lesser height. This shortness of the arms is apparently due to their greater use, and is an unexpected result: but sailors chiefly use their arms in pulling, and not in supporting weights. With sailors, the girth of the neck and the depth of the instep are greater, whilst the circumference of the chest, waist, and hips is less, than in soldiers.
Whether the several foregoing modifications would become hereditary, if the same habits of life were followed during many generations, is not known, but it is probable. Rengger (23. 'Saugethiere von Paraguay,' 1830, s. 4.) attributes the thin legs and thick arms of the Payaguas Indians to successive generations having passed nearly their whole lives in canoes, with their lower extremities motionless. Other writers have come to a similar conclusion in analogous cases. According to Cranz (24. 'History of Greenland,' Eng. translat., 1767, vol. i. p. 230.), who lived for a long time with the Esquimaux, "the natives believe that ingenuity and dexterity in seal-catching (their highest art and virtue) is hereditary; there is really something in it, for the son of a celebrated seal-catcher will distinguish himself, though he lost his father in childhood." But in this case it is mental aptitude, quite as much as bodily structure, which appears to be inherited. It is asserted that the hands of English labourers are at birth larger than those of the gentry. (25. 'Intermarriage,' by Alex. Walker, 1838, p. 377.) From the correlation which exists, at least in some cases (26. 'The Variation of Animals under Domestication,' vol. i. p. 173.), between the development of the extremities and of the jaws, it is possible that in those classes which do not labour much with their hands and feet, the jaws would be reduced in size from this cause. That they are generally smaller in refined and civilised men than in hard-working men or savages, is certain. But with savages, as Mr. Herbert Spencer (27. 'Principles of Biology,' vol. i. p. 455.) has remarked, the greater use of the jaws in chewing coarse, uncooked food, would act in a direct manner on the masticatory muscles, and on the bones to which they are attached. In infants, long before birth, the skin on the soles of the feet is thicker than on any other part of the body; (28. Paget, 'Lectures on Surgical Pathology,' vol. ii, 1853, p. 209.) and it can hardly be doubted that this is due to the inherited effects of pressure during a long series of generations.
It is familiar to every one that watchmakers and engravers are liable to be short-sighted, whilst men living much out of doors, and especially savages, are generally long-sighted. (29. It is a singular and unexpected fact that sailors are inferior to landsmen in their mean distance of distinct vision. Dr. B.A. Gould ('Sanitary Memoirs of the War of the Rebellion,' 1869, p. 530), has proved this to be the case; and he accounts for it by the ordinary range of vision in sailors being "restricted to the length of the vessel and the height of the masts.") Short-sight and long-sight certainly tend to be inherited. (30. 'The Variation of Animals under Domestication,' vol. i. p. 8.) The inferiority of Europeans, in comparison with savages, in eyesight and in the other senses, is no doubt the accumulated and transmitted effect of lessened use during many generations; for Rengger (31. 'Saugethiere von Paraguay,' s. 8, 10. I have had good opportunities for observing the extraordinary power of eyesight in the Fuegians. See also Lawrence ('Lectures on Physiology,' etc., 1822, p. 404) on this same subject. M. Giraud-Teulon has recently collected ('Revue des Cours Scientifiques,' 1870, p. 625) a large and valuable body of evidence proving that the cause of short-sight, "C'est le travail assidu, de pres.") states that he has repeatedly observed Europeans, who had been brought up and spent their whole lives with the wild Indians, who nevertheless did not equal them in the sharpness of their senses. The same naturalist observes that the cavities in the skull for the reception of the several sense- organs are larger in the American aborigines than in Europeans; and this probably indicates a corresponding difference in the dimensions of the organs themselves. Blumenbach has also remarked on the large size of the nasal cavities in the skulls of the American aborigines, and connects this fact with their remarkably acute power of smell. The Mongolians of the plains of northern Asia, according to Pallas, have wonderfully perfect senses; and Prichard believes that the great breadth of their skulls across the zygomas follows from their highly-developed sense organs. (32. Prichard, 'Physical History of Mankind,' on the authority of Blumenbach, vol. i. 1851, p. 311; for the statement by Pallas, vol. iv. 1844, p. 407.)
The Quechua Indians inhabit the lofty plateaux of Peru; and Alcide d'Orbigny states (33. Quoted by Prichard, 'Researches into the Physical History of Mankind,' vol. v. p. 463.) that, from continually breathing a highly rarefied atmosphere, they have acquired chests and lungs of extraordinary dimensions. The cells, also, of the lungs are larger and more numerous than in Europeans. These observations have been doubted, but Mr. D. Forbes carefully measured many Aymaras, an allied race, living at the height of between 10,000 and 15,000 feet; and he informs me (34. Mr. Forbes' valuable paper is now published in the 'Journal of the Ethnological Society of London,' new series, vol. ii. 1870, p.193.) that they differ conspicuously from the men of all other races seen by him in the circumference and length of their bodies. In his table of measurements, the stature of each man is taken at 1000, and the other measurements are reduced to this standard. It is here seen that the extended arms of the Aymaras are shorter than those of Europeans, and much shorter than those of Negroes. The legs are likewise shorter; and they present this remarkable peculiarity, that in every Aymara measured, the femur is actually shorter than the tibia. On an average, the length of the femur to that of the tibia is as 211 to 252; whilst in two Europeans, measured at the same time, the femora to the tibiae were as 244 to 230; and in three Negroes as 258 to 241. The humerus is likewise shorter relatively to the forearm. This shortening of that part of the limb which is nearest to the body, appears to be, as suggested to me by Mr. Forbes, a case of compensation in relation with the greatly increased length of the trunk. The Aymaras present some other singular points of structure, for instance, the very small projection of the heel.
These men are so thoroughly acclimatised to their cold and lofty abode, that when formerly carried down by the Spaniards to the low eastern plains, and when now tempted down by high wages to the gold-washings, they suffer a frightful rate of mortality. Nevertheless Mr. Forbes found a few pure families which had survived during two generations: and he observed that they still inherited their characteristic peculiarities. But it was manifest, even without measurement, that these peculiarities had all decreased; and on measurement, their bodies were found not to be so much elongated as those of the men on the high plateau; whilst their femora had become somewhat lengthened, as had their tibiae, although in a less degree. The actual measurements may be seen by consulting Mr. Forbes's memoir. From these observations, there can, I think, be no doubt that residence during many generations at a great elevation tends, both directly and indirectly, to induce inherited modifications in the proportions of the body. (35. Dr. Wilckens ('Landwirthschaft. Wochenblatt,' No. 10, 1869) has lately published an interesting essay shewing how domestic animals, which live in mountainous regions, have their frames modified.)
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 braindevelopment of microcephalous idiots, as described in Vogt's memoir. (36. 'Memoires 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 interspace 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.
Malthus has discussed these several checks, but he does not lay stress enough on what is probably the most important of all, namely infanticide, especially of female infants, and the habit of procuring abortion. These practices now prevail in many quarters of the world; and infanticide seems formerly to have prevailed, as Mr. M'Lennan (61. 'Primitive Marriage,' 1865.) has shewn, on a still more extensive scale. These practices appear to have originated in savages recognising the difficulty, or rather the impossibility of supporting all the infants that are born. Licentiousness may also be added to the foregoing checks; but this does not follow from failing means of subsistence; though there is reason to believe that in some cases (as in Japan) it has been intentionally encouraged as a means of keeping down the population.
If we look back to an extremely remote epoch, before man had arrived at the dignity of manhood, he would have been guided more by instinct and less by reason than are the lowest savages at the present time. Our early semi- human progenitors would not have practised infanticide or polyandry; for the instincts of the lower animals are never so perverted (62. A writer in the 'Spectator' (March 12, 1871, p. 320) comments as follows on this passage:--"Mr. Darwin finds himself compelled to reintroduce a new doctrine of the fall of man. He shews that the instincts of the higher animals are far nobler than the habits of savage races of men, and he finds himself, therefore, compelled to re-introduce,
-in a form of the substantial orthodoxy of which he appears to be quite unconscious,--and to introduce as a scientific hypothesis the doctrine that man's gain of KNOWLEDGE was the cause of a temporary but long-enduring moral deterioration as indicated by the many foul customs, especially as to marriage, of savage tribes. What does the Jewish tradition of the moral degeneration of man through his snatching at a knowledge forbidden him by his highest instinct assert beyond this?") as to lead them regularly to destroy their own offspring, or to be quite devoid of jealousy. There would have been no prudential restraint from marriage, and the sexes would have freely united at an early age. Hence the progenitors of man would have tended to increase rapidly; but checks of some kind, either periodical or constant, must have kept down their numbers, even more severely than with existing savages. What the precise nature of these checks were, we cannot say, any more than with most other animals. We know that horses and cattle, which are not extremely prolific animals, when first turned loose in South America, increased at an enormous rate. The elephant, the slowest breeder of all known animals, would in a few thousand years stock the whole world. The increase of every species of monkey must be checked by some means; but not, as Brehm remarks, by the attacks of beasts of prey. No one will assume that the actual power of reproduction in the wild horses and cattle of America, was at first in any sensible degree increased; or that, as each district became fully stocked, this same power was diminished. No doubt, in this case, and in all others, many checks concur, and different checks under different circumstances; periodical dearths, depending on unfavourable seasons, being probably the most important of all. So it will have been with the early progenitors of man.
We have now seen that man is variable in body and mind; and that the variations are induced, either directly or indirectly, by the same general causes, and obey the same general laws, as with the lower animals. Man has spread widely over the face of the earth, and must have been exposed, during his incessant migrations (63. See some good remarks to this effect by W. Stanley Jevons, "A Deduction from Darwin's Theory," 'Nature,' 1869, p. 231.), to the most diversified conditions. The inhabitants of Tierra del Fuego, the Cape of Good Hope, and Tasmania in the one hemisphere, and of the arctic regions in the other, must have passed through many climates, and changed their habits many times, before they reached their present homes. (64. Latham, 'Man and his Migrations,' 1851, p. 135.) The early progenitors of man must also have tended, like all other animals, to have increased beyond their means of subsistence; they must, therefore, occasionally have been exposed to a struggle for existence, and consequently to the rigid law of natural selection. Beneficial variations of all kinds will thus, either occasionally or habitually, have been preserved and injurious ones eliminated. I do not refer to stronglymarked deviations of structure, which occur only at long intervals of time, but to mere individual differences. We know, for instance, that the muscles of our hands and feet, which determine our powers of movement, are liable, like those of the lower animals, (65. Messrs. Murie and Mivart in their 'Anatomy of the Lemuroidea' ('Transact. Zoolog. Soc.' vol. vii. 1869, pp. 96-98) say, "some muscles are so irregular in their distribution that they cannot be well classed in any of the above groups." These muscles differ even on the opposite sides of the same individual.) to incessant variability. If then the progenitors of man inhabiting any district, especially one undergoing some change in its conditions, were divided into two equal bodies, the one half which included all the individuals best adapted by their powers of movement for gaining subsistence, or for defending themselves, would on an average survive in greater numbers, and procreate more offspring than the other and less well endowed half.
Man in the rudest state in which he now exists is the most dominant animal that has ever appeared on this earth. He has spread more widely than any other highly organised form: and all others have yielded before him. He manifestly owes this immense superiority to his intellectual faculties, to his social habits, which lead him to aid and defend his fellows, and to his corporeal structure. The supreme importance of these characters has been proved by the final arbitrament of the battle for life. Through his powers of intellect, articulate language has been evolved; and on this his wonderful advancement has mainly depended. As Mr. Chauncey Wright remarks (66. Limits of Natural Selection, 'North American Review,' Oct. 1870, p. 295.): "a psychological analysis of the faculty of language shews, that even the smallest proficiency in it might require more brain power than the greatest proficiency in any other direction." He has invented and is able to use various weapons, tools, traps, etc., with which he defends himself, kills or catches prey, and otherwise obtains food. He has made rafts or canoes for fishing or crossing over to neighbouring fertile islands. He has discovered the art of making fire, by which hard and stringy roots can be rendered digestible, and poisonous roots or herbs innocuous. This discovery of fire, probably the greatest ever made by man, excepting language, dates from before the dawn of history. These several inventions, by which man in the rudest state has become so pre-eminent, are the direct results of the development of his powers of observation, memory, curiosity, imagination, and reason. I cannot, therefore, understand how it is that Mr. Wallace (67. 'Quarterly Review,' April 1869, p. 392. This subject is more fully discussed in Mr. Wallace's 'Contributions to the Theory of Natural Selection,' 1870, in which all the essays referred to in this work are re-published. The 'Essay on Man,' has been ably criticised by Prof. Claparede, one of the most distinguished zoologists in Europe, in an article published in the 'Bibliotheque Universelle,' June 1870. The remark quoted in my text will surprise every one who has read Mr. Wallace's celebrated paper on 'The Origin of Human Races Deduced from the Theory of Natural Selection,' originally published in the 'Anthropological Review,' May 1864, p. clviii. I cannot here resist quoting a most just remark by Sir J. Lubbock ('Prehistoric Times,' 1865, p. 479) in reference to this paper, namely, that Mr. Wallace, "with characteristic unselfishness, ascribes it (i.e. the idea of natural selection) unreservedly to Mr. Darwin, although, as is well known, he struck out the idea independently, and published it, though not with the same elaboration, at the same time.") maintains, that "natural selection could only have endowed the savage with a brain a little superior to that of an ape."
Although the intellectual powers and social habits of man are of paramount importance to him, we must not underrate the importance of his bodily structure, to which subject the remainder of this chapter will be devoted; the development of the intellectual and social or moral faculties being discussed in a later chapter.
Even to hammer with precision is no easy matter, as every one who has tried to learn carpentry will admit. To throw a stone with as true an aim as a Fuegian in defending himself, or in killing birds, requires the most consummate perfection in the correlated action of the muscles of the hand, arm, and shoulder, and, further, a fine sense of touch. In throwing a stone or spear, and in many other actions, a man must stand firmly on his feet; and this again demands the perfect co-adaptation of numerous muscles. To chip a flint into the rudest tool, or to form a barbed spear or hook from a bone, demands the use of a perfect hand; for, as a most capable judge, Mr. Schoolcraft (68. Quoted by Mr. Lawson Tait in his 'Law of Natural Selection,' 'Dublin Quarterly Journal of Medical Science,' Feb. 1869. Dr. Keller is likewise quoted to the same effect.), remarks, the shaping fragments of stone into knives, lances, or arrow-heads, shews "extraordinary ability and long practice." This is to a great extent proved by the fact that primeval men practised a division of labour; each man did not manufacture his own flint tools or rude pottery, but certain individuals appear to have devoted themselves to such work, no doubt receiving in exchange the produce of the chase. Archaeologists are convinced that an enormous interval of time elapsed before our ancestors thought of grinding chipped flints into smooth tools. One can hardly doubt, that a man-like animal who possessed a hand and arm sufficiently perfect to throw a stone with precision, or to form a flint into a rude tool, could, with sufficient practice, as far as mechanical skill alone is concerned, make almost anything which a civilised man can make. The structure of the hand in this respect may be compared with that of the vocal organs, which in the apes are used for uttering various signal-cries, or, as in one genus, musical cadences; but in man the closely similar vocal organs have become adapted through the inherited effects of use for the utterance of articulate language.
Turning now to the nearest allies of men, and therefore to the best representatives of our early progenitors, we find that the hands of the Quadrumana are constructed on the same general pattern as our own, but are far less perfectly adapted for diversified uses. Their hands do not serve for locomotion so well as the feet of a dog; as may be seen in such monkeys as the chimpanzee and orang, which walk on the outer margins of the palms, or on the knuckles. (69. Owen, 'Anatomy of Vertebrates,' vol. iii. p. 71.) Their hands, however, are admirably adapted for climbing trees. Monkeys seize thin branches or ropes, with the thumb on one side and the fingers and palm on the other, in the same manner as we do. They can thus also lift rather large objects, such as the neck of a bottle, to their mouths. Baboons turn over stones, and scratch up roots with their hands. They seize nuts, insects, or other small objects with the thumb in opposition to the fingers, and no doubt they thus extract eggs and young from the nests of birds. American monkeys beat the wild oranges on the branches until the rind is cracked, and then tear it off with the fingers of the two hands. In a wild state they break open hard fruits with stones. Other monkeys open mussel-shells with the two thumbs. With their fingers they pull out thorns and burs, and hunt for each other's parasites. They roll down stones, or throw them at their enemies: nevertheless, they are clumsy in these various actions, and, as I have myself seen, are quite unable to throw a stone with precision.
It seems to me far from true that because "objects are grasped clumsily" by monkeys, "a much less specialised organ of prehension" would have served them (70. 'Quarterly Review,' April 1869, p. 392.) equally well with their present hands. On the contrary, I see no reason to doubt that more perfectly constructed hands would have been an advantage to them, provided that they were not thus rendered less fitted for climbing trees. We may suspect that a hand as perfect as that of man would have been disadvantageous for climbing; for the most arboreal monkeys in the world, namely, Ateles in America, Colobus in Africa, and Hylobates in Asia, are either thumbless, or their toes partially cohere, so that their limbs are converted into mere grasping hooks. (71. In Hylobates syndactylus, as the name expresses, two of the toes regularly cohere; and this, as Mr. Blyth informs me, is occasionally the case with the toes of H. agilis, lar, and leuciscus. Colobus is strictly arboreal and extraordinarily active (Brehm, 'Thierleben,' B. i. s. 50), but whether a better climber than the species of the allied genera, I do not know. It deserves notice that the feet of the sloths, the most arboreal animals in the world, are wonderfully hook- like.
As soon as some ancient member in the great series of the Primates came to be less arboreal, owing to a change in its manner of procuring subsistence, or to some change in the surrounding conditions, its habitual manner of progression would have been modified: and thus it would have been rendered more strictly quadrupedal or bipedal. Baboons frequent hilly and rocky districts, and only from necessity climb high trees (72. Brehm, 'Thierleben,' B. i. s. 80.); and they have acquired almost the gait of a dog. Man alone has become a biped; and we can, I think, partly see how he has come to assume his erect attitude, which forms one of his most conspicuous characters. Man could not have attained his present dominant position in the world without the use of his hands, which are so admirably adapted to act in obedience to his will. Sir C. Bell (73. 'The Hand,' etc., 'Bridgewater Treatise,' 1833, p. 38.) insists that "the hand supplies all instruments, and by its correspondence with the intellect gives him universal dominion." But the hands and arms could hardly have become perfect enough to have manufactured weapons, or to have hurled stones and spears with a true aim, as long as they were habitually used for locomotion and for supporting the whole weight of the body, or, as before remarked, so long as they were especially fitted for climbing trees. Such rough treatment would also have blunted the sense of touch, on which their delicate use largely depends. From these causes alone it would have been an advantage to man to become a biped; but for many actions it is indispensable that the arms and whole upper part of the body should be free; and he must for this end stand firmly on his feet. To gain this great advantage, the feet have been rendered flat; and the great toe has been peculiarly modified, though this has entailed the almost complete loss of its power of prehension. It accords with the principle of the division of physiological labour, prevailing throughout the animal kingdom, that as the hands became perfected for prehension, the feet should have become perfected for support and locomotion. With some savages, however, the foot has not altogether lost its prehensile power, as shewn by their manner of climbing trees, and of using them in other ways. (74. Haeckel has an excellent discussion on the steps by which man became a biped: 'Naturliche Schopfungsgeschicte,' 1868, s. 507. Dr. Buchner ('Conferences sur la Theorie Darwinienne,' 1869, p. 135) has given good cases of the use of the foot as a prehensile organ by man; and has also written on the manner of progression of the higher apes, to which I allude in the following paragraph: see also Owen ('Anatomy of Vertebrates,' vol. iii. p. 71) on this latter subject.
If it be an advantage to man to stand firmly on his feet and to have his hands and arms free, of which, from his pre-eminent success in the battle of life there can be no doubt, then I can see no reason why it should not have been advantageous to the progenitors of man to have become more and more erect or bipedal. They would thus have been better able to defend themselves with stones or clubs, to attack their prey, or otherwise to obtain food. The best built individuals would in the long run have succeeded best, and have survived in larger numbers. If the gorilla and a few allied forms had become extinct, it might have been argued, with great force and apparent truth, that an animal could not have been gradually converted from a quadruped into a biped, as all the individuals in an intermediate condition would have been miserably ill-fitted for progression. But we know (and this is well worthy of reflection) that the anthropomorphous apes are now actually in an intermediate condition; and no one doubts that they are on the whole well adapted for their conditions of life. Thus the gorilla runs with a sidelong shambling gait, but more commonly progresses by resting on its bent hands. The long-armed apes occasionally use their arms like crutches, swinging their bodies forward between them, and some kinds of Hylobates, without having been taught, can walk or run upright with tolerable quickness; yet they move awkwardly, and much less securely than man. We see, in short, in existing monkeys a manner of progression intermediate between that of a quadruped and a biped; but, as an unprejudiced judge (75. Prof. Broca, La Constitution des Vertebres caudales; 'La Revue d'Anthropologie,' 1872, p. 26, (separate copy).) insists, the anthropomorphous apes approach in structure more nearly to the bipedal than to the quadrupedal type.
As the progenitors of man became more and more erect, with their hands and arms more and more modified for prehension and other purposes, with their feet and legs at the same time transformed for firm support and progression, endless other changes of structure would have become necessary. The pelvis would have to be broadened, the spine peculiarly curved, and the head fixed in an altered position, all which changes have been attained by man. Prof. Schaaffhausen (76. 'On the Primitive Form of the Skull,' translated in 'Anthropological Review,' Oct. 1868, p. 428. Owen ('Anatomy of Vertebrates,' vol. ii. 1866, p. 551) on the mastoid processes in the higher apes.) maintains that "the powerful mastoid processes of the human skull are the result of his erect position;" and these processes are absent in the orang, chimpanzee, etc., and are smaller in the gorilla than in man. Various other structures, which appear connected with man's erect position, might here have been added. It is very difficult to decide how far these correlated modifications are the result of natural selection, and how far of the inherited effects of the increased use of certain parts, or of the action of one part on another. No doubt these means of change often co-operate: thus when certain muscles, and the crests of bone to which they are attached, become enlarged by habitual use, this shews that certain actions are habitually performed and must be serviceable. Hence the individuals which performed them best, would tend to survive in greater numbers.
The free use of the arms and hands, partly the cause and partly the result of man's erect position, appears to have led in an indirect manner to other modifications of structure. The early male forefathers of man were, as previously stated, probably furnished with great canine teeth; but as they gradually acquired the habit of using stones, clubs, or other weapons, for fighting with their enemies or rivals, they would use their jaws and teeth less and less. In this case, the jaws, together with the teeth, would become reduced in size, as we may feel almost sure from innumerable analogous cases. In a future chapter we shall meet with a closely parallel case, in the reduction or complete disappearance of the canine teeth in male ruminants, apparently in relation with the development of their horns; and in horses, in relation to their habit of fighting with their incisor teeth and hoofs.
In the adult male anthropomorphous apes, as Rutimeyer (77. 'Die Grenzen der Thierwelt, eine Betrachtung zu Darwin's Lehre,' 1868, s. 51.), and others, have insisted, it is the effect on the skull of the great development of the jaw-muscles that causes it to differ so greatly in many respects from that of man, and has given to these animals "a truly frightful physiognomy." Therefore, as the jaws and teeth in man's progenitors gradually become reduced in size, the adult skull would have come to resemble more and more that of existing man. As we shall hereafter see, a great reduction of the canine teeth in the males would almost certainly affect the teeth of the females through inheritance. As the various mental faculties gradually developed themselves the brain would almost certainly become larger. No one, I presume, doubts that the large proportion which the size of man's brain bears to his body, compared to the same proportion in the gorilla or orang, is closely connected with his higher mental powers. We meet with closely analogous facts with insects, for in ants the cerebral ganglia are of extraordinary dimensions, and in all the Hymenoptera these ganglia are many times larger than in the less intelligent orders, such as beetles. (78. Dujardin, 'Annales des Sciences Nat.' 3rd series, Zoolog., tom. xiv. 1850, p. 203. See also Mr. Lowne, 'Anatomy and Phys. of the Musca vomitoria,' 1870, p. 14. My son, Mr. F. Darwin, dissected for me the cerebral ganglia of the Formica rufa.) 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 notorious, yet their cerebral ganglia are not so large as the quarter of a small pin's head. Under this point of view, the brain of an ant is one of the most marvellous atoms of matter in the world, perhaps more so than the brain of a 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 (79. 'Philosophical Transactions,' 1869, p. 513.), 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 cubic inches. Professor Broca (80. 'Les Selections,' M. P. Broca, 'Revue d'Anthropologies,' 1873; see also, as quoted in C. Vogt's 'Lectures on Man,' Engl. translat., 1864, pp. 88, 90. Prichard, 'Physical History of Mankind,' vol. i. 1838, p. 305.) found that the nineteenth century skulls from graves in Paris were larger than those from vaults of the twelfth century, in the proportion of 1484 to 1426; and that the increased size, as ascertained by measurements, was exclusively in the frontal part of the skull--the seat of the intellectual faculties. 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. (81. In the interesting article just referred to, Prof. Broca has well remarked, that in civilised nations, the average capacity of the skull must be lowered by the preservation of a considerable number of individuals, weak in mind and body, who would have been promptly eliminated in the savage state. On the other hand, with savages, the average includes only the more capable individuals, who have been able to survive under extremely hard conditions of life. Broca thus explains the otherwise inexplicable fact, that the mean capacity of the skull of the ancient Troglodytes of Lozere is greater than that of modern Frenchmen.) With respect to the lower animals, M.E. Lartet (82. 'Comptes-rendus des Sciences,' etc., June 1, 1868.), 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 are more complex in the more recent forms. On the other hand, I have shewn (83. The 'Variation of Animals and Plants under Domestication,' vol. i. pp. 124-129.) 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 their intellect, instincts, senses and voluntary movements but little. 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 have become fixed either sideways or backwards, owing to disease, one of the two eyes has changed its position, and the shape of the skull has been altered apparently by the pressure of the brain in a new direction. (84. Schaaffhausen gives from Blumenbach and Busch, the cases of the spasms and cicatrix, in 'Anthropological Review,' Oct. 1868, p. 420. Dr. Jarrold ('Anthropologia,' 1808, pp. 115, 116) adduces from Camper and from his own observations, cases of the modification of the skull from the head being fixed in an unnatural position. He believes that in certain trades, such as that of a shoemaker, where the head is habitually held forward, the forehead becomes more rounded and prominent.) I have shewn that with long-eared rabbits even so trifling a cause as the lopping forward of one ear drags forward almost every bone of the skull on that side; so that the bones on the opposite side 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 on 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 3.15 and the latter 4.3 inches in length. (85. 'Variation of Animals and Plants under Domestication,' vol. i. p. 117, on the elongation of the skull; p. 119, on the effect of the lopping of one ear.) One of the most marked distinctions in different races of men is that the skull in some is elongated, and in others rounded; and here the explanation suggested by the case of the rabbits may hold good; for Welcker finds that short "men incline more to brachycephaly, and tall men to dolichocephaly" (86. Quoted by Schaaffhausen, in 'Anthropological Review,' Oct. 1868, p. 419.); and tall men may be compared with the larger and longerbodied rabbits, all of which have elongated skulls or are dolichocephalic.
From these several facts we can understand, to a certain extent, the means by which the great size and more or less rounded form of the skull have 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 porpoises (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 (87. Owen, 'Anatomy of Vertebrates,' vol. iii. p. 619.) than those on the lowlands. May we then infer that man became divested of hair from having aboriginally inhabited some tropical land? That the hair is 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--on the assumption 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. The fact, however, 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 (88. Isidore Geoffroy St.-Hilaire remarks ('Histoire Nat. Generale,' tom. ii. 1859, pp. 215-217) on the head of man being covered with long hair; also on the upper surfaces of monkeys and of other mammals being more thickly clothed than the lower surfaces. This has likewise been observed by various authors. Prof. P. Gervais ('Histoire Nat. des Mammiferes,' tom. i. 1854, p. 28), however, states that in the Gorilla the hair is thinner on the back, where it is partly rubbed off, than on the lower surface.), is opposed to the supposition that man became naked through the action of the sun. Mr. Belt believes (89. The 'Naturalist in Nicaragua,' 1874, p. 209. As some confirmation of Mr. Belt's view, I may quote the following passage from Sir W. Denison ('Varieties of Vice-Regal Life,' vol. i. 1870, p. 440): "It is said to be a practice with the Australians, when the vermin get troublesome, to singe themselves.") that within the tropics it is an advantage to man to be destitute of hair, as he is thus enabled to free himself of the multitude of ticks (acari) and other parasites, with which he is often infested, and which sometimes cause ulceration. But whether this evil is of sufficient magnitude to have led to the denudation of his body through natural selection, may be doubted, since none of the many quadrupeds inhabiting the tropics have, as far as I know, acquired any specialised means of relief. The view which seems to me the most probable is that man, or rather primarily woman, became divested of hair for ornamental purposes, as we shall see under Sexual Selection; and, according to this belief, it is not surprising that man should differ so greatly in hairiness from all other Primates, for characters, gained through sexual selection, often differ to an extraordinary degree in closely related forms.
According to a popular impression, the absence of a tail is eminently distinctive of man; but as those apes which come nearest to him are destitute of this organ, its disappearance 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 vertebres 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 vertabrae, 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 past or present times. 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 very 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.
It is, as I can now see, probable that all organic beings, including man, possess peculiarities of structure, which neither are now, nor were formerly of any service to them, and which, therefore, are of no physiological importance. 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 efficient cause. If these causes, whatever they may be, were to act more uniformly and energetically during a lengthened period (and against this no reason can be assigned), the result would probably be not a mere slight individual difference, but a well-marked and constant modification, though one of no physiological importance. Changed structures, which are in no way beneficial, cannot be kept uniform through natural selection, though the injurious will be 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. During successive periods, the same organism might in this manner acquire successive modifications, which 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.CONCLUSION.
In this chapter we have seen that as man at the present day is liable, like every other animal, to multiform individual differences or slight variations, so no doubt were the early progenitors of man; the variations being formerly induced by the same general causes, and governed by the same general and complex laws as at present. As all animals tend to multiply beyond their means of subsistence, so it must have been with the progenitors of man; and this would inevitably lead to a struggle for existence and to natural selection. The latter process would be greatly aided by the inherited effects of the increased use of parts, and these two processes would incessantly react on each other. It appears, also, as we shall hereafter see, that various unimportant characters have been acquired by man through sexual selection. An unexplained residuum of change must be left to the assumed uniform action of those unknown agencies, which occasionally induce strongly marked and abrupt deviations of structure in our domestic productions.
Judging from the habits of savages and of the greater number of the Quadrumana, primeval men, and even their ape-like progenitors, probably lived in society. With strictly social animals, natural selection sometimes acts on the individual, through the preservation of variations which are beneficial to the community. A community which includes a large number of well-endowed individuals increases in number, and is victorious over other less favoured ones; even although each separate member gains no advantage over the others of the same community. Associated insects have thus acquired many remarkable structures, which are of little or no service to the individual, such as the pollen-collecting apparatus, or the sting of the worker-bee, or the great jaws of soldierants. With the higher social animals, I am not aware that any structure has been modified solely for the good of the community, though some are of secondary service to it. For instance, the horns of ruminants and the great canine teeth of baboons appear to have been acquired by the males as weapons for sexual strife, but they are used in defence of the herd or troop. In regard to certain mental powers the case, as we shall see in the fifth chapter, is wholly different; for these faculties have been chiefly, or even exclusively, gained for the benefit of the community, and the individuals thereof have at the same time gained an advantage indirectly.
It has often been objected to such views as the foregoing, that man is one of the most helpless and defenceless creatures in the world; and that during his early and less welldeveloped condition, he would have been still more helpless. The Duke of Argyll, for instance, insists (96. 'Primeval Man,' 1869, p. 66.) that "the human frame has diverged from the structure of brutes, in the direction of greater physical helplessness and weakness. That is to say, it is a divergence which of all others it is most impossible to ascribe to mere natural selection." He adduces the naked and unprotected state of the body, the absence of great teeth or claws for defence, the small strength and speed of man, and his slight power of discovering food or of avoiding danger by smell. To these deficiencies there might be added one still more serious, namely, that he cannot climb quickly, and so escape from enemies. The loss of hair would not have been a great injury to the inhabitants of a warm country. For we know that the unclothed Fuegians can exist under a wretched climate. When we compare the defenceless state of man with that of apes, we must remember that the great canine teeth with which the latter are provided, are possessed in their full development by the males alone, and are chiefly used by them for fighting with their rivals; yet the females, which are not thus provided, manage to survive.
In regard to bodily size or strength, we do not know whether man is descended from some small species, like the chimpanzee, or from one as powerful as the gorilla; and, therefore, we cannot say whether man has become larger and stronger, or smaller and weaker, than his ancestors. We should, however, bear in mind that an animal possessing great size, strength, and ferocity, and which, like the gorilla, could defend itself from all enemies, would not perhaps have become social: and this would most effectually have checked the acquirement of the higher mental qualities, such as sympathy and the love of his fellows. Hence it might have been an immense advantage to man to have sprung from some comparatively weak creature.
The small strength and speed of man, his want of natural weapons, etc., are more than counterbalanced, firstly, by his intellectual powers, through which he has formed for himself weapons, tools, etc., though still remaining in a barbarous state, and, secondly, by his social qualities which lead him to give and receive aid from his fellow-men. No country in the world abounds in a greater degree with dangerous beasts than Southern Africa; no country presents more fearful physical hardships than the Arctic regions; yet one of the puniest of races, that of the Bushmen, maintains itself in Southern Africa, as do the dwarfed Esquimaux in the Arctic regions. The ancestors of man were, no doubt, inferior in intellect, and probably in social disposition, to the lowest existing savages; but it is quite conceivable that they might have existed, or even flourished, if they had advanced in intellect, whilst gradually losing their brute-like powers, such as that of climbing trees, etc. But these ancestors would not have been exposed to any special danger, even if far more helpless and defenceless than any existing savages, had they inhabited some warm continent or large island, such as Australia, New Guinea, or Borneo, which is now the home of the orang. And natural selection arising from the competition of tribe with tribe, in some such large area as one of these, together with the inherited effects of habit, would, under favourable conditions, have sufficed to raise man to his present high position in the organic scale.