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THE DESCENT OF MAN
SELECTION IN RELATION TO SEX
Works by Charles Darwin, F.R.S.
Life and Letters of Charles Darwin. With an Autobiographical Chapter.
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THE DESCENT OF MAN
SELECTION IN RELATION TO SEX
CHARLES DARWIN, M.A., F.R.S.
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of Favoured Races in the Struggle for Life. Popular Edition, with a
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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have been introduced, and four of the old drawings have been replaced by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
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,
First Edition February 24, 1871.
Second Edition September, 1874.
PART I. THE DESCENT OR ORIGIN OF MAN.
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.
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.
Comparison of the Mental Powers of Man and the Lower Animals.
The difference in mental power between the highest ape and the lowest
savage, immense--Certain instincts in common--The emotions--Curiosity--
--Tools and weapons used by animals--Abstraction, Self-consciousness--
Language--Sense of beauty--Belief in God, spiritual agencies,
Comparison of the Mental Powers of Man and the Lower Animals--continued.
The moral sense--Fundamental proposition--The qualities of social animals--
Origin of sociability--Struggle between opposed instincts--Man a social
animal--The more enduring social instincts conquer other less persistent
instincts--The social virtues alone regarded by savages--The self-regarding
virtues acquired at a later stage of development--The importance of the
judgment of the members of the same community on conduct--Transmission of
On the Development of the Intellectual and Moral Faculties during Primeval
and Civilised times.
Advancement of the intellectual powers through natural selection--
Importance of imitation--Social and moral faculties--Their development
within the limits of the same tribe--Natural selection as affecting
civilised nations--Evidence that civilised nations were once barbarous.
On the Affinities and Genealogy of Man.
Position of man in the animal series--The natural system genealogical--
Adaptive characters of slight value--Various small points of resemblance
between man and the Quadrumana--Rank of man in the natural system--
Birthplace and antiquity of man--Absence of fossil connecting-links--Lower
stages in the genealogy of man, as inferred firstly from his affinities and
secondly from his structure--Early androgynous condition of the Vertebrata
On the Races of Man.
The nature and value of specific characters--Application to the races of
man--Arguments in favour of, and opposed to, ranking the so-called races of
man as distinct species--Sub-species--Monogenists and polygenists--
Convergence of character--Numerous points of resemblance in body and mind
between the most distinct races of man--The state of man when he first
spread over the earth--Each race not descended from a single pair--The
extinction of races--The formation of races--The effects of crossing--
Slight influence of the direct action of the conditions of life--Slight or
no influence of natural selection--Sexual selection.
PART II. SEXUAL SELECTION.
Principles of Sexual Selection.
Secondary sexual characters--Sexual selection--Manner of action--Excess of
males--Polygamy--The male alone generally modified through sexual
selection--Eagerness of the male--Variability of the male--Choice exerted
by the female--Sexual compared with natural selection--Inheritance at
corresponding periods of life, at corresponding seasons of the year, and as
limited by sex--Relations between the several forms of inheritance--Causes
why one sex and the young are not modified through sexual selection--
Supplement on the proportional numbers of the two sexes throughout the
animal kingdom-- The proportion of the sexes in relation to natural
Secondary Sexual Characters in the Lower Classes of the Animal Kingdom.
These characters are absent in the lowest classes--Brilliant colours--
Mollusca--Annelids--Crustacea, secondary sexual characters strongly
developed; dimorphism; colour; characters not acquired before maturity--
Spiders, sexual colours of; stridulation by the males--Myriapoda.
Secondary Sexual Characters of Insects.
Diversified structures possessed by the males for seizing the females--
Differences between the sexes, of which the meaning is not understood--
Difference in size between the sexes--Thysanura--Diptera--Hemiptera--
Homoptera, musical powers possessed by the males alone--Orthoptera, musical
instruments of the males, much diversified in structure; pugnacity;
colours--Neuroptera, sexual differences in colour--Hymenoptera, pugnacity
and odours--Coleoptera, colours; furnished with great horns, apparently as
an ornament; battles; stridulating organs generally common to both sexes.
Insects, continued.--Order Lepidoptera.
(Butterflies and Moths.)
Courtship of Butterflies--Battles--Ticking noise--Colours common to both
sexes, or more brilliant in the males--Examples--Not due to the direct
action of the conditions of life--Colours adapted for protection--Colours
of moths--Display--Perceptive powers of the Lepidoptera--Variability--
Causes of the difference in colour between the males and females--Mimicry,
female butterflies more brilliantly coloured than the males--Bright colours
of caterpillars--Summary and concluding remarks on the secondary sexual
character of insects--Birds and insects compared.
Secondary Sexual Characters of Fishes, Amphibians, and Reptiles.
Fishes: Courtship and battles of the males--Larger size of the females--
Males, bright colours and ornamental appendages; other strange characters--
Colours and appendages acquired by the males during the breeding-season
alone--Fishes with both sexes brilliantly coloured--Protective colours--The
less conspicuous colours of the female cannot be accounted for on the
principle of protection--Male fishes building nests, and taking charge of
the ova and young. AMPHIBIANS: Differences in structure and colour
between the sexes--Vocal organs. REPTILES: Chelonians--Crocodiles--
Snakes, colours in some cases protective--Lizards, battles of--Ornamental
appendages--Strange differences in structure between the sexes--Colours--
Sexual differences almost as great as with birds.
Secondary Sexual Characters of Birds.
Sexual differences--Law of battle--Special weapons--Vocal organs--
Instrumental music--Love-antics and dances--Decorations, permanent and
seasonal--Double and single annual moults--Display of ornaments by the
Choice exerted by the female--Length of courtship--Unpaired birds--Mental
qualities and taste for the beautiful--Preference or antipathy shewn by the
female for particular males--Variability of birds--Variations sometimes
abrupt--Laws of variation--Formation of ocelli--Gradations of character--
Case of Peacock, Argus pheasant, and Urosticte.
Discussion as to why the males alone of some species, and both sexes of
others are brightly coloured--On sexually-limited inheritance, as applied
to various structures and to brightly-coloured plumage--Nidification in
relation to colour--Loss of nuptial plumage during the winter.
The immature plumage in relation to the character of the plumage in both
sexes when adult--Six classes of cases--Sexual differences between the
males of closely-allied or representative species--The female assuming the
characters of the male--Plumage of the young in relation to the summer and
winter plumage of the adults--On the increase of beauty in the birds of the
world--Protective colouring--Conspicuously coloured birds--Novelty
appreciated--Summary of the four chapters on birds.
Secondary Sexual Characters of Mammals.
The law of battle--Special weapons, confined to the males--Cause of absence
of weapons in the female--Weapons common to both sexes, yet primarily
acquired by the male--Other uses of such weapons--Their high importance--
Greater size of the male--Means of defence--On the preference shewn by
either sex in the pairing of quadrupeds.
Secondary Sexual Characters of Mammals--continued.
Voice--Remarkable sexual peculiarities in seals--Odour--Development of the
hair--Colour of the hair and skin--Anomalous case of the female being more
ornamented than the male--Colour and ornaments due to sexual selection--
Colour acquired for the sake of protection--Colour, though common to both
sexes, often due to sexual selection--On the disappearance of spots and
stripes in adult quadrupeds--On the colours and ornaments of the
PART III. SEXUAL SELECTION IN RELATION TO MAN, AND CONCLUSION.
Secondary Sexual Characters of Man.
Differences between man and woman--Causes of such differences, and of
certain characters common to both sexes--Law of battle--Differences in
mental powers, and voice--On the influence of beauty in determining the
marriages of mankind--Attention paid by savages to ornaments--Their ideas
of beauty in women--The tendency to exaggerate each natural peculiarity.
Secondary Sexual Characters of Man--continued.
On the effects of the continued selection of women according to a different
standard of beauty in each race--On the causes which interfere with sexual
selection in civilised and savage nations--Conditions favourable to sexual
selection during primeval times--On the manner of action of sexual
selection with mankind--On the women in savage tribes having some power to
choose their husbands--Absence of hair on the body, and development of the
beard--Colour of the skin--Summary.
General Summary and Conclusion.
Main conclusion that man is descended from some lower form--Manner of
development--Genealogy of man--Intellectual and moral faculties--Sexual
THE DESCENT OF MAN; AND SELECTION IN RELATION TO SEX.
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
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.
PART I. THE DESCENT OR ORIGIN OF MAN.
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
[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.
L. Tail or os coccyx.]
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
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."
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
Some few persons have the power of contracting the superficial muscles on
their scalps; and these muscles are in a variable and partially rudimentary
condition. M. A. de Candolle has communicated to me a curious instance of
the long-continued persistence or inheritance of this power, as well as of
its unusual development. He knows a family, in which one member, the
present head of the family, could, when a youth, pitch several heavy books
from his head by the movement of the scalp alone; and he won wagers by
performing this feat. His father, uncle, grandfather, and his three
children possess the same power to the same unusual degree. This family
became divided eight generations ago into two branches; so that the head of
the above-mentioned branch is cousin in the seventh degree to the head of
the other branch. This distant cousin resides in another part of France;
and on being asked whether he possessed the same faculty, immediately
exhibited his power. This case offers a good illustration how persistent
may be the transmission of an absolutely useless faculty, probably derived
from our remote semi-human progenitors; since many monkeys have, and
frequently use the power, of largely moving their scalps up and down. (27.
See my 'Expression of the Emotions in Man and Animals,' 1872, p. 144.)
The extrinsic muscles which serve to move the external ear, and the
intrinsic muscles which move the different parts, are in a rudimentary
condition in man, and they all belong to the system of the panniculus; they
are also variable in development, or at least in function. I have seen one
man who could draw the whole ear forwards; other men can draw it upwards;
another who could draw it backwards (28. Canestrini quotes Hyrtl.
('Annuario della Soc. dei Naturalisti,' Modena, 1867, p. 97) to the same
effect.); and from what one of these persons told me, it is probable that
most of us, by often touching our ears, and thus directing our attention
towards them, could recover some power of movement by repeated trials. The
power of erecting and directing the shell of the ears to the various points
of the compass, is no doubt of the highest service to many animals, as they
thus perceive the direction of danger; but I have never heard, on
sufficient evidence, of a man who possessed this power, the one which might
be of use to him. The whole external shell may be considered a rudiment,
together with the various folds and prominences (helix and anti-helix,
tragus and anti-tragus, etc.) which in the lower animals strengthen and
support the ear when erect, without adding much to its weight. Some
authors, however, suppose that the cartilage of the shell serves to
transmit vibrations to the acoustic nerve; but Mr. Toynbee (29. 'The
Diseases of the Ear,' by J. Toynbee, F.R.S., 1860, p. 12. A distinguished
physiologist, Prof. Preyer, informs me that he had lately been
experimenting on the function of the shell of the ear, and has come to
nearly the same conclusion as that given here.), after collecting all the
known evidence on this head, concludes that the external shell is of no
distinct use. The ears of the chimpanzee and orang are curiously like
those of man, and the proper muscles are likewise but very slightly
developed. (30. Prof. A. Macalister, 'Annals and Magazine of Natural
History,' vol. vii. 1871, p. 342.) I am also assured by the keepers in the
Zoological Gardens that these animals never move or erect their ears; so
that they are in an equally rudimentary condition with those of man, as far
as function is concerned. Why these animals, as well as the progenitors of
man, should have lost the power of erecting their ears, we cannot say. It
may be, though I am not satisfied with this view, that owing to their
arboreal habits and great strength they were but little exposed to danger,
and so during a lengthened period moved their ears but little, and thus
gradually lost the power of moving them. This would be a parallel case
with that of those large and heavy birds, which, from 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
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 thirty-five, with the lanugo of a foetus, and finds it
quite similar in texture; therefore, as he remarks, the case may be
attributed to an arrest of development in the hair, together with its
continued growth. Many delicate children, as I have been assured by a
surgeon to a hospital for children, have their backs covered by rather long
silky hairs; and such cases probably come under the same head.
It appears as if the posterior molar or wisdom-teeth were tending to become
rudimentary in the more civilised races of man. These teeth are rather
smaller than the other molars, as is likewise the case with the
corresponding teeth in the chimpanzee and orang; and they have only two
separate fangs. They do not cut through the gums till about the
seventeenth year, and I have been assured that they are much more liable to
decay, and are earlier lost than the other teeth; but this is denied by
some eminent dentists. They are also much more liable to vary, both in
structure and in the period of their development, than the other teeth.
(42. Dr. Webb, 'Teeth in Man and the Anthropoid Apes,' as quoted by Dr. C.
Carter Blake in Anthropological Review, July 1867, p. 299.) In the
Melanian races, on the other hand, the wisdom-teeth are usually furnished
with three separate fangs, and are generally sound; they also differ from
the other molars in size, less than in the Caucasian races. (43. Owen,
'Anatomy of Vertebrates,' vol. iii. pp. 320, 321, and 325.) Prof.
Schaaffhausen accounts for this difference between the races by "the
posterior dental portion of the jaw being always shortened" in those that
are civilised (44. 'On the Primitive Form of the Skull,' Eng. translat.,
in 'Anthropological Review,' Oct. 1868, p. 426), and this shortening may, I
presume, be attributed to civilised men habitually feeding on soft, cooked
food, and thus using their jaws less. I am informed by Mr. Brace that it
is becoming quite a common practice in the United States to remove some of
the molar teeth of children, as the jaw does not grow large enough for the
perfect development of the normal number. (45. Prof. Montegazza writes to
me from Florence, that he has lately been studying the last molar teeth in
the different races of man, and has come to the same conclusion as that
given in my text, viz., that in the higher or civilised races they are on
the road towards atrophy or elimination.)
With respect to the alimentary canal, I have met with an account of only a
single rudiment, namely the vermiform appendage of the caecum. The caecum
is a branch or diverticulum of the intestine, ending in a cul-de-sac, and
is extremely long in many of the lower vegetable-feeding mammals. In the
marsupial koala it is actually more than thrice as long as the whole body.
(46. Owen, 'Anatomy of Vertebrates,' vol. iii. pp. 416, 434, 441.) It is
sometimes produced into a long gradually-tapering point, and is sometimes
constricted in parts. It appears as if, in consequence of changed diet or
habits, the caecum had become much shortened in various animals, the
vermiform appendage being left as a rudiment of the shortened part. That
this appendage is a rudiment, we may infer from its small size, and from
the evidence which Prof. Canestrini (47. 'Annuario della Soc. d. Nat.'
Modena, 1867, p. 94.) has collected of its variability in man. It is
occasionally quite absent, or again is largely developed. The passage is
sometimes completely closed for half or two-thirds of its length, with the
terminal part consisting of a flattened solid expansion. In the orang this
appendage is long and convoluted: in man it arises from the end of the
short caecum, and is commonly from four to five inches in length, being
only about the third of an inch in diameter. Not only is it useless, but
it is sometimes the cause of death, of which fact I have lately heard two
instances: this is due to small hard bodies, such as seeds, entering the
passage, and causing inflammation. (48. M. C. Martins ("De l'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 supra-condyloid 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
supra-condyloid 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. Pruner-Bey 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
beau-ideal 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."
(The Works of J. Hookham Frere, vol. ii. 1872, p. 334.))
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
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 brain-development 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.)
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
Professor Canestrini, after discussing the foregoing and various analogous
cases, arrives at the same conclusion as that just given. He adduces
another instance, in the case of the malar bone (40. 'Annuario della Soc.
dei Naturalisti,' Modena, 1867, p. 83. Prof. Canestrini gives extracts on
this subject from various authorities. Laurillard remarks, that as he has
found a complete similarity in the form, proportions, and connection of the
two malar bones in several human subjects and in certain apes, he cannot
consider this disposition of the parts as simply accidental. Another paper
on this same anomaly has been published by Dr. Saviotti in the 'Gazzetta
delle Cliniche,' Turin, 1871, where he says that traces of the division may
be detected in about two per cent. of adult skulls; he also remarks that it
more frequently occurs in prognathous skulls, not of the Aryan race, than
in others. See also G. Delorenzi on the same subject; 'Tre nuovi casi
d'anomalia dell' osso malare,' Torino, 1872. Also, E. Morselli, 'Sopra una
rara anomalia dell' osso malare,' Modena, 1872. Still more recently Gruber
has written a pamphlet on the division of this bone. I give these
references because a reviewer, without any grounds or scruples, has thrown
doubts on my statements.), which, in some of the Quadrumana and other
mammals, normally consists of two portions. This is its condition in the
human foetus when two months old; and through arrested development, it
sometimes remains thus in man when adult, more especially in the lower
prognathous races. Hence Canestrini concludes that some ancient progenitor
of man must have had this bone normally divided into two portions, which
afterwards became fused together. In man the frontal bone consists of a
single piece, but in the embryo, and in children, and in almost all the
lower mammals, it consists of two pieces separated by a distinct suture.
This suture occasionally persists more or less distinctly in man after
maturity; and more frequently in ancient than in recent crania, especially,
as Canestrini has observed, in those exhumed from the Drift, and belonging
to the brachycephalic type. Here again he comes to the same conclusion as
in the analogous case of the malar bones. In this, and other instances
presently to be given, the cause of ancient races approaching the lower
animals in certain characters more frequently than do the modern races,
appears to be, that the latter stand at a somewhat greater distance in the
long line of descent from their early semi-human progenitors.
Various other anomalies in man, more or less analogous to the foregoing,
have been advanced by different authors, as cases of reversion; but these
seem not a little doubtful, for we have to descend extremely low in the
mammalian series, before we find such structures normally present. (41. A
whole series of cases is given by Isidore Geoffroy St.-Hilaire, 'Hist. des
Anomalies,' tom, iii, p. 437. A reviewer ('Journal of Anatomy and
Physiology,' 1871, p. 366) blames me much for not having discussed the
numerous cases, which have been recorded, of various parts arrested in
their development. He says that, according to my theory, "every transient
condition of an organ, during its development, is not only a means to an
end, but once was an end in itself." This does not seem to me necessarily
to hold good. Why should not variations occur during an early period of
development, having no relation to reversion; yet such variations might be
preserved and accumulated, if in any way serviceable, for instance, in
shortening and simplifying the course of development? And again, why
should not injurious abnormalities, such as atrophied or hypertrophied
parts, which have no relation to a former state of existence, occur at an
early period, as well as during maturity?)
In man, the canine teeth are perfectly efficient instruments for
mastication. But their true canine character, as Owen (42. 'Anatomy of
Vertebrates,' vol. iii. 1868, p. 323.) remarks, "is indicated by the
conical form of the crown, which terminates in an obtuse point, is convex
outward and flat or sub-concave within, at the base of which surface there
is a feeble prominence. The conical form is best expressed in the Melanian
races, especially the Australian. The canine is more deeply implanted, and
by a stronger fang than the incisors." Nevertheless, this tooth no longer
serves man as a special weapon for tearing his enemies or prey; it may,
therefore, as far as its proper function is concerned, be considered as
rudimentary. In every large collection of human skulls some may be found,
as Haeckel (43. 'Generelle Morphologie,' 1866, B. ii. s. clv.) observes,
with the canine teeth projecting considerably beyond the others in the same
manner as in the anthropomorphous apes, but in a less degree. In these
cases, open spaces between the teeth in the one jaw are left for the
reception of the canines of the opposite jaw. An inter-space of this kind
in a Kaffir skull, figured by Wagner, is surprisingly wide. (44. Carl
Vogt's 'Lectures on Man,' Eng. translat., 1864, p. 151.) Considering how
few are the ancient skulls which have been examined, compared to recent
skulls, it is an interesting fact that in at least three cases the canines