^B

mmMmsM

■

miffs '■•

■HI

mm

■':; •'••■■ j i *■:■••■:'■;■■ '^ ■:■•">■■■• ";

;•'-'•••■■■•.-•'.■•■' :
■'•■•'•■■':::■■ :- ■■■■.;:■

-:= ••>■:>;•■''• -: , :•■

:;.-.'.':'■•■

:'!■}» WW.:r.
TB Ml

A ■:■
^HHBI

JbHHB

UB

"»

^&^

^

o
□

a

CD

—

OB
P

=

""*"

a

-D

g

JZ

N

ft

Js

&

. 7/

/<//

THE DESCENT OF MAN,

AND

SELECTION IN KELATION TO SEX.

Phot. Druck v. A. Naumann & Schroeder, Leipzig.

C. Lehr d. J. sculp

o

THE

0

. /TV 4

DESCENT OF MAN,

AND

SELECTION W KELATION TO SEX.

By CHAKLES DARWIN, M.A., F.R.S., &c.

IN TWO VOLUMES.— Vol. I,

WITH ILLUSTRATIONS.

LONDON:
JOHN MURRAY, ALBEMARLE . STREET.

1871.

[The right of Translation is reserved.'}

BY THE SAME AUTHOR.

K>»

ON THE ORIGIN OF SPECIES BY MEANS OF NATURAL

SELECTION ; or, The Preservation of Favoured Races in the Struggle
for Life; Fifth Edition {Tenth Thousand), with Additions and Corrections.
1869. Murray.

THE VARIATION OF ANIMALS AND PLANTS UNDER

DOMESTICATION. In two vols. With Illustrations. 1868. Murray.

ON THE VARIOUS CONTRIVANCES by which BRITISH

AND FOREIGN ORCHIDS ARE FERTILISED BY INSECTS ; and on the
Good Effects of Crossing. With numerous Woodcuts. Murray.

A NATURALIST'S VOYAGE ROUND THE WORLD ; or,

A Journal of Researches into the Natural History and Geology of the
Countries visited during the voyage of H.M.S. ' Beagle,' under the command of
Captain FitzRoy, R.N. Eleventh Thousand. Murray.

ON THE STRUCTURE AND DISTRIBUTION OF CORAL

REEFS. Smith, Elder, & Co.

GEOLOGICAL OBSERVATIONS ON VOLCANIC ISLANDS.

Smith, Elder, & Co.

GEOLOGICAL OBSERVATIONS ON SOUTH AMERICA.

Smith, Elder, & Co.

A MONOGRAPH OF THE CIRRIPEDIA. With numerous

Illustrations. 2 vols. 8vo. Hardwickb.

ON THE MOVEMENTS AND HABITS OF CLIMBING

PLANTS. With Woodcuts. Williams & Norgate.

LONDON : PRINTED BY WILLIAM CLOWES AND SONS, STAMFORD STREET,

AND CHARING CROSS.

CONTENTS.

Introduction Pa«;e 1-5

PART I. ^>\CA/

ON THE DESCENT OF MAN. /v> rtoS a,

— ° — / rw ^ "^ < *"'

CHAPTER I. , J L16R AR

The Evidence of the Descent op man from Wfe\ ^k***'**'

Lower Form. \<xV^7/
Nature of the evidence bearing on the origin of man — Homb^g^^s ^ |
structures in man and the lower animals — Miscellaneous pomt
of correspondence — Development — Eudimentary structures,
muscles, sense-organs, hair, hones, reproductive organs, &c. —
The bearing of these three great classes of facts on the origin of
man 9-33

CHAPTER II.

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 — C uriosity — Imitation — Attention — Mem ory — Ima-
gination — Keason — Progressive improvement — Tools and
weapons used by animals — Language — Self-consciousness —
Sense of beauty — Belief in God, spiritual agencies, superstitions

34-69

CHAPTER III.

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 in-
stincts— Man a social animal — The more enduring social instincts

2 4

VI CONTENTS OF VOL. 1.

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 — Trans-
mission of moral tendencies — Summary .. .. Page 70-106

CHAPTEE IV.

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 — Ee-
version — Correlated variation — Eate 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 107-157

CHAPTEE V.

On the Development of the Intellectual and Moral
Faculties during Primeval and Civilised Times.

The advancement of the intellectual powers through natural selec-
tion— 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"' 158-184

CHAPTEE VI.

On the Affinities and Genealogy of Man.

Position of man in the animal series — The natural system genea-
logical — Adaptive characters of slight value — Similar small
points of resemblance between man and the Quadrumana —
Eank of man in the natural system — Birthplace and antiquity

CONTENTS OF VOL. I.

Vll

of man — Absence of fossil connecting-links — Lower stages in
the genealogy of man, as inferred, firstly from his affinities and
secondly from bis structure — Early androgynous condition of
the Vertebrata — Conclusion .. ..... .. Page 185-213

CHAPTER VII.

Ox THE 1'vACES 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 — Mono-
genists 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 ex-
tinction of races — The formation of races — The effects of cross-
ing—Slight influence of the direct action of the conditions of life
— Slight or no influence of natural selection — Sexual selection

214-250

PART II.

SEXUAL SELECTION.

*&

CHAPTER VIII.

Principles of Sexual Selection.

m

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 — On the limita-
tation of the numbers of the two sexes through natural selection

253-320

Vlll CONTENTS OF VOL. I.

CHAPTER IX.

Secondary Sexual Characters in the Lower Classes of

the Animal Kingdom.

These characters 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 Page 321-340

CHAPTER X.

Secondary Sexual Characters of Insecst.

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, pug-
nacity and colours — Coleoptera, colours ; furnished with great
horns, apparently as an ornament ; battles ; stridulating organs
generally common to both sexes 341-385

CHAPTER XI.

Insects, continued. — Order Lefidoptera.

Courtship of butterflies — Battles — Ticking noise — Colours com-
mon 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 — Per-
ceptive powers of the Lepidoptera ■ — Variability — Causes of the
difference in colour between the males and females — Mimickry,
female butterflies more brilliantly coloured than the males —
Bright colours of caterpillars — Summary and concluding re-
marks on the secondary sexual characters of insects — Birds
and insects compared 386-423

THE DESCENT OE MAN;

AND ON

SELECTION IN BELATION TO SEX.

INTRODUCTION.

The nature of the following work will be best under-
stood 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 suffi-
cient 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
anv general conclusion respecting his manner of appear-
ance on this earth. Now the case wears a wholly dif-
ferent aspect. When a naturalist like Carl Vogt ven-
tures 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 ;

VOL. I. B

2 INTRODUCTION.

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 other men, 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 appli-
cable 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 argu-
ments derived from the nature of the affinities which
connect together whole groups of organisms — their geo-
graphical distribution in past and present times, and
their geological succession. The homological structure,
embryological development, and rudimentary organs of
a species, whether it be man or any other animal, to
which our attention may be directed, remain to be con-
sidered ; 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 sup-
port 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

INTRODUCTION. 6

his development ; and thirdly, the value of the differ-
ences 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 dis-
cussed in many valuable works. The high antiquity of
man has recently been demonstrated by the labours
of a host of eminent meD, 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 conclu-
sively shewn that in every single visible character man
differs less from the higher apes tban 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. La^

b 2

3 INTRODUCTION.

marck 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, Biichner, Kolle, &c.,' and especially by
Hackel. This last naturalist, besides his great work,
' Generelle Morphologie ' (I860), has recently (1868,
with a second edit, in 1870), published his ' Natiirliche
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 1 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.
Hackel's writings, I give his authority in the text, other
statements I leave as they originally stood in my manu-
script, 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 pro-
bable that sexual selection has played an important
part in differentiating the races of man; but in my

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 iiber die Darwin'-
sche Theorie:' zweite Auflage, 1868, von Dr. L. Biichner ; translated
into French under the title ' Conferences sur la Theorie Darwinienne,'
1869. ' Der Mensch,, im Lichte der Darwin'sche Lehre,' 1865, von
Dr. F. Kolle, 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, p. 81) a
very carious paper on rudimentary characters, as bearing on the origin
of man. Another work has (1869) been published by Dr. Barrage.
Francesco, bearing in Italian the title of " Man, made in the imaee of
God, was also made in the image of the ape."

INTRODUCTION. 0

'Origin of Species' (first edition, p. 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 Consequently
the second part of the present work, treating of sexual
selection, has extended to an inordinate length, com-
pared with the first part ; but this could not be
avoided.

I had intended adding to the present volumes an
essay on the expression of the various emotions by man
and the lower animals. My attention wTas 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, which is
partially completed, for separate publication.

2 Prof. Hiickel is the sole author who, since the publication of the
' Origin,' has discussed, in his various works, in a very able manner
the subject of sexual selection, and has seen its full importance.

-7

II! RV

Part I.

THE DESCENT OR ORIGIN OF MAN.

■CNV00°S "

&\ -

pART I.—THE DESCENT OF MAN.

CHAPTEK I.

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, &c. —
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; such as that of the transmission of characters
to the same age or sex. Again, are the variations the re-
sult, 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 in-
stance by correlation, the inherited effects of use and
disuse, &c. ? Is man subject to similar malconformations,
the result of arrested development, "of reduplication of
parts, &o, 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

10 THE DESCENT OF MAN. Part I.

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 thev react on
each other, both in the first and succeeding genera-
tions ? 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 each other, 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 conve-
niently deferred for a time ; and we will first see how
far the bodily structure of man shows traces, more or
less plain, of his descent from some lower form. In the
two succeeding chapters the mental powers of man, in
comparison with those of the lower animals, will be con-
sidered.

The Bodily Structure of Man. — It is notorious that
man is constructed on the same general type or model
with 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 im-
portant of all the organs, follows the same law, as shewn
by Huxley and other anatomists. Bischoff,1 who is a
hostile witness, admits that every chief fissure and fold

1 ' Grosslrirawmdungen des Menschen/ 1S(3S, s. 96.

Chap. I. HOMOLOGICAL STRUCTURE. 11

in the brain of man has its analogy in that of the orang ;
but he acids that at no period of development do their
brains perfectly agree ; nor could this be expected, for
otherwise their mental powers would have been the same.
Vulpian 2 remarks : " Les differences reelles qui existent
" entre l'encephale de l'homme et celui des singes supe-
" rieurs, sont bien minimes. II ne faut pas se faire
" d'illusions a cet egard. L'homme est bien plus ~pves
" des singes anthropomorphes par les caracteres anato-
(S miques de son cerveau que ceux-ci ne le sont non-
" seulement des autres mammiferes, mais memes 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 struc-
ture, 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 hydro-
phobia, variola, the glanders, &c. ; and this fact proves
the close similarity 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,3 who carefully observed for a long
time the Cebus Azarss in its native land, found it liable
to catarrh, with the usual symptoms, and which when

2 'Lee. sur la Phys.' 1866, p. 890, as quoted by M. Dally, 'L'Ordre
des Primates et le Transformisme,' 1868, p. 29.

3 ' Naturgeschickte der S'augethiere von Paraguay,' 1S30, s. 50.

12 THE DESCENT OF MAN. Part I.

often recurrent led to consumption. These monkeys
suffered also from apoplexy, inflammation of the bowels,
and cataract in the eye. The younger ones when shed-
ding 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
spirituous liquors : they will also, as I have myself seen,
smoke tobacco with pleasure. Brehm asserts that the
natives of north-eastern xAirica 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.4 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 simi-
larly their whole nervous system is affected.

Man is infested with internal parasites, sometimes
causing fatal effects, and is plagued by external para-
sites, all of which belong to the same genera or families
with those infesting other mammals. Man is subject like
other mammals, birds, and even insects, to that mys-
terious law, which causes certain normal processes, such
as gestation, as well as the maturation and duration of
various diseases, to follow lunar periods.5 His wounds

4 Brehm, ' Thierleben,' B. i. 1864, s. 75, 86. On the Ateles, s. 105.
For other analogous statements, see s. 25, 107.

5 With respect to insects see Dr. Laycock ' On a General Law of
Vital Periodicity,' British Association, 1842. Dr. Macculloch, ' Silli-

Chap. I. HOMOLOGICAL STRUCTURE. 13

are repaired by the same process of healing ; and the
stumps left after the amputation of his limbs occa-
sionally possess, especially during an early embryonic
period, some power of regeneration, as in the lowest
animals.6

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 7 to the birth and nurturing of the young. Mon-
keys 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.8 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.9 Man

man's North American Journal of Science,' vol. xvii. p. 305, lias seen
a dog suffering from tertian ague.

6 I have given the evidence on this head in my ' Variation of
Animals and Plants under Domestication,' vol. ii. p. 15.

" " Mares e diversis generibus Quadrumanorum sine dubio dignoseunt
" feminas hum anas a maribus. Primum, credo, odoratu, postea aspectu.
" Mr. Youatt, qui diu in Hortis Zoologicis (Bestiariis) medicus animal-
" ium erat, vir in rebus observandis cautus et sagax, hoc mibi certissime
" probavit, et curatores ejusdem loci et alii e ministris confirmaverunt.
' ' Sir Andrew Smith et Brelim notabant idem in Cynocephalo. Illus-
" trissimus Cuvier etiam narrat multa de hac re qua ut opinor nihil
" turpius potest indicari inter omnia liominibus et Quadrumanis com-
*' munia. Narrat enim Cynocephalum quendam in furorem incidere
" aspectu feminarum aliquarum, sed nequaquam accendi tanto furore
" ab omnibus. Semper eligebat juniores, et dignoscebat in turba, et
" advocabat voce gestuque."

s This remark is made with respect to Cynocephalus and the an-
thropomorphous apes by Geoffroy Saint-Hilaire and F. Cuvier, ' Hist.
Nat. des Mammiteres,' torn. i. 1824.

9 Huxley, ■ Man's Place in Nature,' 1863, p. 34.

14 THE DESCENT OF MAN. Part I.

differs from woman in size, bodily strength, hairyness,
&c, as well as in mind, in the same manner as do the
two sexes of many mammals. It is, in short, scarcely
possible to exaggerate the close correspondence in gene-
ral structure, in the minute structure of the tissues, in
chemical composition and in constitution, between man
and the higher animals, especially the anthropomor-
phous apes.

Embryonic Development. — 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 verte-
brate 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 (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 funda-
" mental form." It is, says Prof. Huxley,10 " quite in
" the later stages of development that the young human
" being presents marked differences from the young
" ape, while the latter departs as much from the dog
" in its developments, as the man does. Startling as
•* this last assertion may appear to be, it is demonstrably
" true."

As some of my readers may never nave seen a draw-
ing of an embryo, I have given one of man and another
of a dog, at about the same early stage of development,

10 ' Man's Place in Nature,' 1SC3, p. 67.

Chap. I.

EMBRYONIC DEVELOPMENT.

15

Fig. 1 . Upper figure human embryo, from Ecker. Lower figure that of a dog,

f'om Bischoff.

a. Fore-brain, cerebral hemispheres, &c.

b. Mid-brain, corpora quadrigemina.

c. Hind-brain, cerebellum, medulla ob
longata.

d. Eye.

e. Ear.

/. First visceral arch.

g. Second visceral arch.
H. Vertebral columns and muscles in
process of development.
i. Anterior )
K. Posterior } extremities.

L. Tail or os coccyx.

16 THE DESCENT OF MAN. Part I.

carefully copied from two works of undoubted accu-

racy.11

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 like-
wise resembles in various points of structure cer-
tain low forms when adult. 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." 12 In the embryos of
all air-breathing vertebrates, certain glands called the cor-
pora Wolffiana, correspond with and act like the kidneys
of mature fishes.13 Even at a later embryonic period,
some striking resemblances between man and the lower
animals may be observed. Bischoff says that the con-
volutions of the brain in a human foetus at the end of
the seventh month reach about the same stage of deve-
lopment as in a baboon when adult.14 The great toe, as
Prof. Owen remarks,15 " which forms the fulcrum when
" standing or walking, is perhaps the most characteristic

11 The human emhryo (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, ' Entwicklungsgesckichte des Hunde-Eies,' 1845, tab. xi. fig.
42 b. This drawing is five times magnified, the embryo being 25 days
old. The internal viscera have been omitted, and the uterine appen-
dages 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. Hackel has also given analogous drawings in
his ' Schopfungsgeschichte.'

12 Prof. Wyman in ' Proc. of American Acad, of Sciences,' vol. iv.
I860, p. 17. '

13 Owen, ' Anatomy of Vertebrates,' vol. i. p. 533.

14 'Die Grosshimwindungen des Menschen,' 1868, s. 95.

15 'Anatomy of Vertebrates,' vol. ii. p. 553.

Chap. I. RUDIMENTS. 17

" peculiarity in the human structure ; " but in an em-
bryo, about an inch in length, Prof. Wyman 16 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,17 who after asking, does man originate in a
different way from a dog, bird, frog or fish ? says, " the
" reply is uot doubtful for a moment; without question,
" the mode of origin and the early stages of the develop-
" ment 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."

Budiments. — This subject, though not intrinsically
more important than the two last, will for several rea-
sons be here treated with more fullness.18 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 cannot
suppose that they were developed under the conditions

16 'Proc. Soc. Nat. Hist.' Boston, 1863, vol. ix. p. 185.

17 ' Man's Place in Nature,' p. G5.

18 I had written a rough copy of this chapter before reading a valu-
able paper, " Caratteri rudimc ntali in ordine all' origine del uoino "
(' Annuario della Soc. d. Nat.,' Modena, 1867, p. 81), by G. Canestrini.
to which paper I am considerably indebted. H'ackel has given admi-
rable discussions on this whole subject, under the title of Dysteleology,
in his ' Generelle Morphologie' and ' Schopfungsgeschichte.'

VOL. 1. C

18 THE DESCENT OF MAN. Part I.

which now exist. Organs in this latter state are not
strictly rudimentary, but they are tending in this direc-
tion. Nascent organs, on the other hand, though not
fully developed, are of high service to their possessors,
and are capable of further development. Kudimentary
organs are eminently variable; and this is partly in-
telligible, as they are useless or nearly useless, and
consequently are no longer subjected to natural selec-
tion. They often become wholly suppressed. When
this occurs, they are nevertheless liable to occasional
reappearance through reversion ; and this is a circum-
stance well worthy of attention.

Disuse at that period of life, when an organ is chiefly
used, and this is generally during maturity, together
with inheritance at a corresponding period of life, seem
to have been the chief agents in causing organs to be-
come rudimentary. 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. Rudi-
ments, however, may occur in one sex of parts normally
present in the other sex ; and such rudiments, as we
shall hereafter see, have often originated in a distinct
manner. In some cases organs have been reduced by
means of natural selection, from having become inju-
rious 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 savins:
to be effected by the economy of growth would be very
small,19 are difficult to understand. The final and com-

19 Some good criticisms on this subject have been given by Messrs.
Murie and Mivart, in < Transact. Zoolug. Soc' 1869, vol. vii. p. 92.

Chap. I. RUDIMENTS. 19

plete 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, and apparently in
no other way. But as the whole subject of rudimentary
organs has been fully discussed and illustrated in my
former works,20 1 need here say no more on this head.

Eudiments of various muscles have been observed in
many parts of the human body ; 21 and not a few muscles,
which are regularly present in some of the lower ani-
mals 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, on the forehead, by which the
eyebrows are raised. The platysma myoides, which is
well developed on the neck, belongs to this system, but
cannot be voluntarily brought into action. Prof.
Turner, of Edinburgh, has occasionally detected, as he
informs me, muscular fasciculi in five different situa-
tions, namely in the axillae, near the scapulae, &c, all of
which must be referred to the system of the panniculus.
He has also shewn 22 that the musculus sternalis or ster-
nalis brutorum, which is not an extension of the rectus
abdominal is, but is closely allied to the panniculus, oc-

20 ' Variation of Animals and Plants under Domestication,' vol. ii.
pp. 317 and 397. See also 'Origin of Species,' 5th edit. p. 535.

21 For instance M. Richard (' Annales des Sciences Nat.' 3rd series,
Zoolog. 1852, torn, 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.

22 Prof. W. Turner, ' Proc. Ptoyal Soc. Edinburgh,' 18GG-G7, p. Q5.

c 2

20 THE DESCENT OF MAN. Part I.

curred in the proportion of about 3 per cent, in upwards
of 600 bodies : he adds, that this muscle affords " an
" excellent illustration of the statement that occasional
" and rudimentary structures are especially liable to
" variation in arrangement."

Some few persons have the power of contracting the
superficial muscles on their scalps ; and these muscles
are in a variable and partially rudimentary condition.
M. A. de Candolle has communicated to me a curious
instance of the long-continued persistence or inheritance
of this power, as well as of its unusual development.
He knows a family, in which one member, the present
head of a 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 all his three chil-
dren 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 persistently an
absolutely useless faculty may be transmitted.

The extrinsic muscles which serve to move the whole
external ear, and the intrinsic muscles which move the
different parts, all of which belong to the system of the
panniculus, are in a rudimentary condition in man ; they
are also variable in development, or at least in function.
I have seen one man who could draw his ears for-
wards, and another who could draw them backwards;23

23 Cabestrini quotes Hyrt. (' Annuario del]a Soc. dei Naturalisti,'
Modena, 1807, p. 97) to the same effect.

Chap. I. KUDIMENTS. 21

and from what one of these persons told ine, it is pro-
bable that most of us by often touching our ears and
thus directing our attention towards them, could by
repeated trials recover some power of movement. The
faculty of erecting the ears and of directing them to
different points of the compass, is no doubt of the
highest service to many animals, as they thus perceive
the point of danger ; but I have never heard of a man
who possessed the least power of erecting his ears, —
the one movement which might be of use to him. The
whole external shell of the ear may be considered a
rudiment, together with the various folds and promi-
nences (helix and anti-helix, tragus and anti-tragus, &c.)
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,24 after collecting all the
known evidence on this head, concludes that the exter-
nal shell is of no distinct use. The ears of the chim-
panzee and orang are curiously like those of man, and I
am assured by the keepers in the Zoological Gardens
that these animals never move or erect them ; so that
they are in an equally rudimentary condition, as far as
function is concerned, as in man. 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 quite 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^bffls,y^7^s

The Diseases of the Ear,' by J. Toynbee, F.K.S., lwgph*2.^ *"»- <*

H LIBRARY

24 i

22

THE DESCENT OF MAN.

PaPwT I.

which from inhabiting oceanic islands have not been
exposed to the attacks of beasts of prey, and have con-
sequently lost the power of using their wings for flight.
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 signification. 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 pecu-
liarity consists in a little blunt point, projecting from
the inwardly folded margin, or helix. Mr. Woolner
made an exact model of one such case, and has sent

me the accompanying drawing.
(Fig. 2.) These points not only
project inwards, but often a little
outwards, so that they are visible
when the head is viewed from di-
rectly in front or behind. They
are variable in size and some-
what in position, standing either
a little higher or lower ; and they
sometimes occur on one ear and
not on the other. Now the mean-
ing of these projections is not,
I think, doubtful; but it may
be thought that they offer too
trifling a character to be worth notice. This thought,
however, is as false as it is natural. Every character,
however slight, must be the result of some definite
cause ; and if it occurs in many individuals deserves
consideration. The helix obviously consists of the ex-
treme margin of the ear folded inwards ; and this fold-
ing appears to be in some manner connected with the

Fig. 2. Human Ear, modelled
and drawn by Mr. Woolner.

a. The projecting point.

Chap. I. RUDIMENTS. 23

whole external ear being permanently pressed back-
wards. In many monkeys, which do not stand high in
the order, as baboons and some species of macacus,23 the
upper portion of t^e 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 and probably a little outwards. This
could actually be observed in a specimen of the Ateles
beelzebuth in the Zoological Gardens ; and we may safely
conclude that it is a similar structure — a vestige of
formerly pointed ears — which occasionally reappears in
man.

The nictitating membrane, or third eyelid, with its
accessory muscles and other structures, is especially
well developed in birds, and is of much functional im-
portance to them, as it can be rapidly drawn across the
whole eye-ball. It is found in some reptiles and amphi-
bians, and in certain fishes, as in sharks. It is fairly
well developed in the two lower divisions of the mam-
malian series, namely, in the monotremata and marsu-
pials, 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.26

The sense of smell is of the highest importance to
the greater number of mammals — to some, as the rumi-
nants, in warning them of danger ; to others, as the

85 See also some remarks, and the drawings of the ears of the Lemu-
roidea, in Messrs. Murie and Mivart's excellent paper in ' Transact.
Zoolog. Soc.' vol. vii. 1869, pp. 6 and 90.

26 Muller's ' Elements of Physiology/ Eng. translat., 1842, vol. ii. p.
1117. Owen, 'Anatomy of Vertebrates,' vol. iii. p. 2G0 ; ibid, on the
Walrus, ' Proc. Zoolog. Soc.' November 8th, 1854. See also E. Knox,
• Great Artists and Anatomists,' p. 10G. This rudiment apparently is
somewhat larger in Negroes and Australians than in Europeans, see Carl
Vogt, 4 Lectures on Man,' Eng. translat. p. 129.

24 THE DESCENT OF MAN. Paht I.

carnivora, in finding their prey ; to others, as the wild
boar, for both purposes combined. But the sense of
smell is of extremely slight service, if any, even to
savages, in whom it is generally more highly developed
than in the civilised races. It does not warn them of
danger, nor guide them to their food ; nor does it pre-
vent the Esquimaux from sleeping in the most fetid
atmosphere, nor many savages from eating half-putrid
meat. Those who believe in the principle of gradual
evolution, will not readily admit that this sense in its
present state was originally acquired by man, as he
now exists. No doubt 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. We can thus
perhaps understand how it is, as Dr. Maudsley has truly
remarked,27 that the sense of smell in man " is singu-
" larly effective in recalling vividly the ideas and images
'•' of forgotten scenes and places ;" for we see in those
animals, which have this sense highly developed, such as
dogs and horses, that old recollections of persons and
places are strongly associated with their odour.

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 male sex, and fine down on that of the female sex.
In individuals belonging to the same race these hairs
are highly variable, not only in abundance, but like-
wise in position : thus the shoulders in some Europeans
are quite naked, whilst in others they bear thick tufts
of hair.28 There can be little doubt that the hairs

27 'The Physiology and Pathology of Mind,' 2nd edit. 1S6S, p. 134.

2S Eschricht, Ueber die Richtung der Haare am menschlicben Korper,
' Miiller s Archiv fur Anat. und Phys.' 1837, s. 47. I shall often have
to refer to this very curious paper.

Chap I. KUDIMENTS. 25

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.29

I am informed by Mr. Paget that persons belonging
to the same family often have a few hairs in their eye-
brows much longer than the others ; so that this slight
peculiarity seems to be inherited. These hairs appa-
rently represent the vibrissa?, which are used as organs
of touch by many of the lower animals. In a young
chimpanzee I observed that a few upright, rather long,
hairs, projected above the eyes, where the true eyebrows,
if present, would have stood.

The tine 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 Eschricht30 on a female foetus ; but this
is not so surprising a circumstance as it may at first ap-
pear, 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

29 Paget, ' Lectures on Surgical Pathology,' 1S53, vol. i. p. 71.

30 Eschricht, ibid. s. 40, 47.

26 THE DESCENT OP MAN. Part I.

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 coinci-
dence, we must consider the woolly covering of the
foetus to be the ruclimental representative of the first
permanent coat of hair in those mammals which are
born hairy. This representation is much more com-
plete, in accordance with the usual law of embryological
development, than that afforded by the straggling hairs
on the body of the adult.

It appears as if the posterior molar or wisdom-teeth
were tending to become rudimentary in the more civi-
lised 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 dentists. They are also much more
liable to vary both in structure and in the period of
their development than the other teeth.31 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.32 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,33 and this
shortening may, 1 presume, be safely attributed to civi-

S1 Dr. Webb, ' Teeth in Man and the Anthropoid Apes,' as quoted by
Dr. C. Carter Blake in ' Anthropological Keview,' July, 1867, p. 299.

32 Owen, ' Anatomy of Vertebrates,' vol. iii. pp. 320, 321, and 325.

33 ' On the Primitive Form of the Skull,' Eng. translat. in ' Anthro-
pological Keview,' Oct. 1868, p. 426.

Chap. I. EUDIMENTS. 27

lised 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.

With respect to the alimentary canal I have met
with an account of only a single rudiment, namely the
vermiform appendage of the caBcum. The ca3cum is
a branch or diverticulum of the intestine, ending in a
cul-de-sac, and it 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.34 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 ap-
pendage is a rudiment, we may infer from its small
size, and from the evidence which Prof. Canestrini 35 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 flat-
tened solid expansion. In the orang this appendage is
long and convoluted : in man it arises from the end of
the short caecuni, 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 some-
times the cause of death, of which fact I have lately
heard two instances : this is due to small hard bodies,

34 Owen, ' Anatomy of Vertebrates,' vol. iii. pp. 416, 434, 441.

35 ' Annuario della Soc. d. Nat.' Modena, 1867, p. 94.

28 THE DESCENT OF MAN. Part I.

such as seeds, entering the passage and causing inflam-
mation.36

In the Quadrumana and some other orders of mam-
mals, especially in the Carnivora, 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
passes, and often the great artery. Now in the humerus
of man, as Dr. Struthers37 and others have shewn, there
is generally a trace of this passage, and it is sometimes
fairly well developed, being formed by a depending
hook-like process of bone, completed by a band of
ligament. When present the great nerve invariably
passes through it, and this clearly indicates that it is the
liomologue and rudiment of the supra-condyloid fora-
men of the lower animals. Prof. Turner estimates, as
he informs me, that it occurs in about one per cent, of
recent skeletons.

There is another foramen in the humerus, which
may be called the inter-condyloid ; and this occurs in
various genera of anthropoid and other apes,38 and occa-
sionallv in man. It is remarkable that this foramen
seems to have been much more frequently present
during ancient than during recent times. Mr. Busk 39
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

36 M. C. Martins ("De l'Unite Organique," in 'Kevuo des Deux
Mondes,' June 15, 1862, p. 16), and H'ackel ('Generelle Morphologie,'
B. ii. s. 278), have both remarked on the singular fact of this rudiment
sometimes causing death.

37 ' The Lancet,' Jan. 24, 18G3, p. 83. Dr. Knox, ' Great Artists and
Anatomists,' p. 63. See also an important memoir on this process by
Dr. Grube, in the ' Bulletin de l'Acad. Imp. de St. Pe'tersbourg,' torn.
xii. 1867, p. 448.

38 Mr. St. George Mivart, ' Transact. Phil. Soc' 18 .7, p. 310.

39 " On the Caves of Gibraltar," ' Transact. Intermit. Congress of
Prehist. Arch.' Third Session, 1869, p. 159.

Chap. I. KUDIMENTS. 29

" Sud ' at Paris ; and in the Grotto of Orrony, the con-
11 tents of which are referred to the Bronze period, as
" many as eight humeri out of thirty-two were perfo-
" rated ; but this extraordinary proportion, he thinks,
" might be due to the cavern having been a sort of
" ' family vault.' Again, M. Dupont found 30 per cent.
" of perforated bones in the caves of the Valley of the
" Lesse, belonging to the Keindeer period ; whilst M.
"• Leguay, in a sort of dolmen at Argentenil, 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 unno-
" ticed that M. Prnner-Bey states that this condition is
" common in Guanche skeletons." The 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 races, is interesting. One
chief cause seems to be that ancient races stand some-
what nearer than modern races in the long line of
descent to their remote animal-like progenitors.

The os coccyx in man, though functionless as a tail,
plainly represents this part in other vertebrate animals.
At an early embryonic period it is free, and, as we have
seen, projects beyond the lower extremities. In certain
rare and anomalous cases it has been known, according
to Isidore Geoffroy St.-Hilaire and others,40 to form a
small external rudiment of a tail. The os coccyx is
short, usually including only four vertebrae : and these
are in a rudimental condition, for they consist, with the
exception of the basal one, of the centrum alone.41 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

40 Quatrefages lias lately collected the evidence on this subject.
'Kevue des Cours h-'cientifiques/ 1867-1868, p. 625.

41

Owen, < On the Nature of Limbs,' 1849, p. 114.

30 THE DESCENT OF MAN. Part I.

extensor of the tail, which is so largely developed in
many mammals.

The spinal cord in man extends only as far down-
wards as the last dorsal or first lumbar vertebra ; but a
thread-like structure (the filum terminate) 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 im-
portant 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 bodv, 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, though not at the extre-
mity, a similarly convoluted body.

The reproductive system offers various rudimentary
structures ; but these differ in one important respect
from the foregoing cases. We are not here concerned
with a vestige of a part which does not belong to the
species in an efficient state; but with a part which
is always present and efficient in the one sex, being
represented in the other by a mere rudiment. Never-
theless, the occurrence of such rudiments is as diffi-
cult to explain on the belief of the sejmrate 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 ; namely, on parts acquired by one sex

Chap. I. THE DESCENT OF MAX. 31

having been partially transmitted to the other. Here
I will onlv srive some instances of such rudiments. It
is well known that in the males of all mammals, in-
cluding man, rudimentary mammae exist. These in
several instances have become well developed, and have
yielded a copious supply of milk. Their essential iden-
tity in the two sexes is likewise shewn by their occa-
sional sympathetic enlargement in both during an
attack of the measles. The vesicula prostatica, which
has been observed in manv male mammals, is now uni-
versally 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 bifur-
cates, for in the males of these the vesicula likewise
bifurcates.42 Some additional rudimentary structures
belonging to the reproductive system might here have
been adduced.43

The bearing of the three great classes of facts now
given is unmistakeable. But it would be superfluous here
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 simi-
larity of pattern between the hand of a man or monkey,
the foot of a horse, the flipper of a seal, the wing of

42 Lenckart, in Todd's ' Cyclop, of Anat.' 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 a^ well
as in other characters.

43 See, on this subject, Owen, ' Anatomv of Vertebrates,' vol. iii. pp.
675, 676, 706.

32 THE DESCENT OF MAX. Part I.

a bat, &c., is utterly inexplicable. 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 varia-
tions 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 embryo of a
man, dog, seal, bat, reptile, &c, 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 super-
fluous part, aided by the other means previously indi-
cated.

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 struc-
ture 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 de-
rived from their affinities or classification, their geo-
graphical 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

Chap. I.

THE DESCENT OF MAN.

33

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.

VOL. I.

D

S± THE DESCENT OF MAN. Part T.

CHAPTEK II.

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 — Imitation — Attention — Memory — Ima-
gination — Reason — Progressive improvement — Tools and
weapons used by animals — Language — Self-consciousness —
Sense of beauty — Belief in God, spiritual agencies, superstitions.

We have seen in the last chapter that man bears in his
bodily structure clear traces of his descent from some
lower form ; but it may be urged that, as man differs so
greatly in his mental power from all other animals, there
must be some error in this conclusion. No doubt the
difference in this respect is enormous, even if we com-
pare the mind of one of the lowest savages, who has no
words to express any number higher than four, and who
uses no abstract terms for the commonest objects or
affections,1 with that of the most highly organised ape.
The difference would, no doubt, still remain immense,
even if one of the higher apes had been improved or
civilised as much as a dog has been in comparison with
its parent-form, the wolf or jackal. The Fuegians rank
amongst the lowest barbarians ; but I was continually
struck with surprise how closely the three natives on
board H.M.S. " Beagle," who had lived some years in
England and could talk a little English, resembled us
in disposition and in most of our mental faculties. If no

1 See the evidence on these points, as given by Lubbock, ' Prehistoric
Times,' p. 354, &c.

Chap. II. MENTAL POWEES. 35

organic being excepting man had possessed any mental
power, or if his powers had been of a wholly different
nature from those of the lower animals, then we should
never have been able to convince ourselves that our
high faculties had been gradually developed. But it
can be clearly shewn that there is no fundamental
difference of this kind. We must also admit that
there is a much wider interval in mental power be-
tween one of the lowest fishes, as a lamprey or lancelet,
and one of the higher apes, than between an ape and
man ; yet this immense interval is filled up by number-
less gradations.

Nor is the difference slight in moral disposition
between a barbarian, such as the man described by the
old navigator Byron, who dashed his child on the rocks
for dropping a basket of sea-urchins, and a Howard or
Clarkson ; and in intellect, between a savage who does
not use any abstract terms, and a Newton or Shakspeare.
Differences of this kind between the highest men of the
highest races and the lowest savages, are connected by
the finest gradations. Therefore it is possible that they
might pass and be developed into each other. ■

My object in this chapter is solely to shew that there
is no fundamental difference between man and the
higher mammals in their mental faculties. Each divi-
sion of the subject might have been extended into a
separate essay, but must here be treated briefly. As
no classification of the mental powers has been univer-
sally accepted, I shall arrange my remarks in the order
most convenient for my purpose ; and will select those
facts which have most struck me, with the hope that
they may produce some effect on the reader.

With respect to animals very low in the scale, I shall
have to give some additional facts under Sexual Selec-
tion, shewing that their mental powers are higher than

d 2

36 THE DESCENT OF MAN. Part I.

might have been expected. The variability of the facul-
ties in the individuals of the same species is an im-
portant point for us, and some few illustrations will here
be given. But it would be superfluous to enter into
many details on this head, for I have found on frequent
enquiry, that it is the unanimous opinion of all those
who have long attended to animals of many kinds,
including birds, that the individuals differ greatly in
everv mental characteristic. In what manner the mental

at

powrers were first developed in the lowest organisms,
is as hopeless an enquiry as how life first originated.
These are problems for the distant future, if they are
ever to be solved bv man.

As man possesses the same senses with the lower
animals, his fundamental intuitions must be the same.
Man has also some few instincts in common, as that of
self-preservation, sexual love, the love of the mother for
her new-born offspring, the power possessed by the
latter of sucking, and so forth. But man, perhaps, has
somewhat fewer instincts than those possessed by the
animals which come next to him in the series. The
orang in the Eastern islands, and the chimpanzee in
Africa, build platforms on which they sleep ; and, as both
species follow the same habit, it might be argued that
this was due to instinct, but we cannot feel sure that it is
not the result of both animals having similar wants and
possessing similar powers of reasoning. These apes, as
we may assume, avoid the many poisonous fruits of the
tropics, and man has no such knowledge ; but as our
domestic animals, when taken to foreign lands and when
first turned out in the spring, often eat poisonous herbs,
which they afterwards avoid, we cannot feel sure that
the apes do not learn from their own experience or
from that of their parents what fruits to select. It is
however certain, as we shall presently see, that apes have

Chap. II. MENTAL POWERS. 37

an instinctive dread of serpents, and probably of other
dangerous animals.

The fewness and the comparative simplicity of the
instincts in the higher animals are remarkable in con-
trast with those of the lower animals. Cuvier main-
tained that instinct and intelligence stand in an inverse
ratio to each other; and some have thought that the
intellectual faculties of the higher animals have been
gradually developed from their instincts. But Pouch et,
in an interesting essay,2 has shewn that no such inverse
ratio really exists. Those insects which possess the most
wonderful instincts are certainly the most intelligent.
In the vertebrate series, the least intelligent members,
namely fishes and amphibians, do not possess complex
instincts ; and amongst mammals the animal most re-
markable for its instincts, namely the beaver, is highly
intelligent, as will be admitted by every one who has
read Mr. Morgan's excellent account of this animal.3

Although the first dawnings of intelligence, accord-
ing to Mr. Herbert Spencer,4 have been developed
through the multiplication and co-ordination of reflex
actions, and although many of the simpler instincts
graduate into actions of this kind and can hardly be
distinguished from them, as in the case of young animals
sucking, yet the more complex instincts seem to have
originated independently of intelligence. I am, how-
ever, far from wishing to deny that instinctive actions
may lose their fixed and untaught character, and be
replaced by others performed by the aid of the free will.
On the other hand, some intelligent actions — as when
birds on oceanic islands first learn to avoid man — after

2 ' L'Instinct chez les Insectes.' ' Eevue des Deux Mondes,' Feb.
1870, p. 690.

3 ' The American Beaver and his Works,' 1868.

4 ' The Principles of Psychology/ 2nd edit. 1870, pp. 418-443

38 THE DESCENT OF MAN. Part I.

being performed during many generations, become con-
verted into instincts and are inherited. They may then
be said to be degraded in character, for they are no
longer performed through reason or from experience.
But the greater number of the more complex instincts
appear to have been gained in a wholly different man-
ner, through the natural selection of variations of simpler
instinctive actions. Such variations appear to arise from
the same unknown causes acting on the cerebral organ-
isation, which induce slight variations or individual dif-
ferences in other parts of the body ; and these variations,
owing to our ignorance, are often said to arise sponta-
neously. We can, I think, come to no other conclusion
with respect to the origin of the more complex instincts,
when we reflect on the marvellous instincts of sterile
worker-ants and bees, which leave no offspring to inherit
the effects of experience and of modified habits.

Although a high degree of intelligence is certainly
compatible with the existence of complex instincts, as
we see in the insects just named and in the beaver, it is
not improbable that they may to a certain extent inter-
fere with each other's development. Little is known
about the functions of the brain, but we can perceive
that as the intellectual powers become highly developed,
the various parts of the brain must be connected by the
most intricate channels of intercommunication ; and as
a consequence each separate part would perhaps tend to
become less well fitted to answer in a definite and uni-
form, that is instinctive, manner to particular sensations
or associations.

I have thought this digression worth giving, because
we may easily underrate the mental powers of the
higher animals, and especially of man, when we com-
pare their actions founded on the memory of past
events, on foresight, reason, and imagination, with

Chap. II. MENTAL POWEES. 39

exactly similar actions instinctively performed by the
lower animals ; in this latter case the capacity of
performing such actions having been gained, step by
step, through the variability of the mental organs and
natural selection, without any conscious intelligence on
the part of the animal during each successive genera-
tion. No doubt, as Mr. Wallace has argued,5 much of
the intelligent work done by man is due to imitation and
not to reason ; but there is this great difference between
his actions and many of those performed by the lower
animals, namely, that man cannot, on his first trial,
make, for instance, a stone hatchet or a canoe, through
his power of imitation. He has to learn his work by
practice ; a beaver, on the other hand, can make its dam
or canal, and a bird its nest, as well, or nearly as well,
the first time it tries, as when old and experienced.

To return to our immediate subject : the lower
animals, like man, manifestly feel pleasure and pain,
happiness and misery. Happiness is never better exhi-
bited than by young animals, such as puppies, kittens,
lambs, &c„ when playing together, like our own children.
Even insects play together, as has been described by
that excellent observer, P. Huber,6 who saw ants chasing
and pretending to bite each other, like so many puppies.

The fact that the lower animals are excited by the
same emotions as ourselves is so well established, that
it will not be necessary to weary the reader by many
details. Terror acts in the same manner on them as on
us, causing the muscles to tremble, the heart to pal-
pitate, the sphincters to be relaxed, and the hair to
stand on end. Suspicion, the offspring of fear, is emi-
nentlv characteristic of most wild animals. Courage

5 ' Contributions to the Theory of Natural Selection,' 1870, p. 212.

6 ' Kecherches sur les Mceurs des Foumris,' 1810, p. 173.

40 THE DESCENT OF MAN. Part I.

and timidity are extremely variable qualities in the
individuals of the same species, as is plainly seen in
our dogs. Some dogs and horses are ill-tempered and
easily turn sulky ; others are good-tempered ; and
these qualities are certainly inherited. Every one
knows how liable animals are to furious rage, and how
plainly they show it. Many anecdotes, probably true,
have been published on the long-delayed and artful
revenge of various animals. The accurate Kengger and
Brehm7 state that the American and African monkeys
which they kept tame, certainly revenged themselves.
The love of a dog for his master is notorious ; in the
agony of death he has been known to caress his master,
and every one has heard of the dog suffering under
vivisection, who licked the hand of the operator ; this
man, unless he had a heart of stone, must have felt
remorse to the last hour of his life. As Whewell8 has
remarked, "who that reads the touching instances of
" maternal affection, related so often of the women of
•' all nations, and of the females of all animals, can
" doubt that the principle of action is the same in the
" two cases ? "

We see maternal affection exhibited in the most
trifling details ; thus Kengger observed an American
monkey (a Cebus) carefully driving away the flies which
plagued her infant ; and Duvaucel saw a Hylobates
washing the faces of her young ones in a stream. So
intense is the grief of female monkeys for the loss of
their young, that it invariably caused the death of cer-
tain kinds kept under confinement by Brehm in N.

~> All the following statements, given on tlie authority of these two
naturalists, are taken from Kengger's ' Naturges. der Saugethiere
von Paraguay,' 1830, s. 41 -57, and from Brehm's ' Thierleben,' B. i.
s. 10-87. '

8 ' Bridgewater Treatise,' p. 263.

Chap. II. MENTAL POWERS. 41

Africa. Orphan- monkeys were always adopted and
careful] y guarded by the other monkeys, both males
and females. One female baboon had so capacious a
heart that she not only adopted young monkeys of
other species, but stole young dogs and cats, which she
continually carried about. Her kindness, however, did
not go so far as to share her food with her adopted
offspring, at which Brehm was surprised, as his monkeys
always divided everything quite fairly with their own
young ones. An adopted kitten scratched the above-
mentioned affectionate baboon, who certainly had a
fine intellect, for she was much astonished at being
scratched, and immediately examined the kitten's feet,
and without more ado bit off the claws. In the
Zoological Gardens, 1 heard from the keeper that
an old baboon (C. chacma) had adopted a Rhesus
monkey ; but when a young drill and mandrill were
placed in the cage, she seemed to perceive that these
monkeys, though distinct species, were her nearer rela-
tives, for she at once rejected the Rhesus and adopted
both of them. The young Rhesus, as I saw, was greatly
discontented at being thus rejected, and it would, like
a naughty child, annoy and attack the young drill and
mandrill whenever it could do so with safety; this
conduct exciting great indignation in the old baboon.
Monkeys will also, according to Brehm, defend their
master when attacked by any one, as well as dogs to
whom thev are attached, from the attacks of other
dogs. But we here trench on the subject of sympathy,
to which I shall recur. Some of Brehm 's monkeys
took much delight in teasing, in various ingenious
ways, a certain old dog whom they disliked, as well
as other animals.

Most of the more complex emotions are common to
the higher animals and ourselves. Every one has seen

42 THE DESCENT OF MAN. Part I.

how jealous a dog is of his master's affection, if lavished
on any other creature ; and I have observed the same fact
with monkeys. This shews that animals not only love,
but have the desire to be loved. Animals manifestly
feel emulation. They love approbation or praise ; and
a clog carrying a basket for his master exhibits in a high
degree self-complacency or pride. There can, I think,
be no doubt that a dog feels shame, as distinct from
fear, and something very like modesty when begging
too often for food. A great dog scorns the snarling of
a little dog, and this may be called magnanimity.
Several observers have stated that monkeys certainly
dislike being laughed at; and they sometimes invent
imaginary offences. In the Zoological Gardens I saw a
baboon who always got into a furious rage when his
keeper took out a letter or book and read it aloud to
him ; and his rage was so violent that, as I witnessed on
one occasion, he bit his own leg till the blood flowed.

We will now turn to the more intellectual emotions
and faculties, which are very important, as forming the
basis for the development of the higher mental powers.
Animals manifestly enjoy excitement and suffer from
ennui, as may be seen with dogs, and, according to
Rengger, with monkeys. All animals feel Wonder,
and many exhibit Curiosity. They sometimes suffer
from this latter quality, as when the hunter plays antics
and thus attracts them ; I have witnessed this with
deer, and so it is with the wary chamois, and with some
kinds of wild-ducks. Brehm gives a curious account of
the instinctive dread which his monkeys exhibited
towards snakes ; but their curiosity was so great that
they could not desist from occasionally satiating their
horror in a most human fashion, by lifting up the lid
of the box in which the snakes were kept. I was so
much surprised at his account, that I took a stuffed and

Chap. II. MENTAL POWERS. 43

coiled-up snake into the monkey-house at the Zoo-
logical Gardens, and the excitement thus caused was
one of the most curious spectacles which I ever beheld.
Three species of Cercopithecus were the most alarmed ;
they dashed about their cages and uttered sharp signal-
cries of danger, which were understood by the other
monkeys. A few young monkeys and one old Anubis
baboon alone took no notice of the snake. I then
placed the stuffed specimen on the ground in one of
the larger compartments. After a time all the monkeys
collected round it in a large circle, and staring in-
tently, presented a most ludicrous appearance. They
became extremely nervous ; so that when a wooden ball,
with which they were familiar as a plaything, was acci-
dently moved in the straw, under which it was partly
hidden, they all instantly started away. These monkeys
behaved very differently when a dead fish, a mouse, and
some other new objects were placed in their cages ; for
though at first frightened, they soon approached, handled
and examined them. I then placed a live snake in a
paper bag, with the mouth loosely closed, in one of the
larger compartments. One of the monkeys immediately
approached, cautiously opened the bag a little, peeped
in, and instantly dashed away. Then I witnessed what
Brehm has described, for monkey after monkey, with
head raised high and turned on one side, could not
resist taking momentary peeps into the upright bag,
at the dreadful object lying quiet at the bottom. It
would almost appear as if monkeys had some notion of
zoological affinities, for those kept by Brehm exhibited
a strange, though mistaken, instinctive dread of inno-
cent lizards and frogs. An orang, also, has been known
to be much alarmed at the first sight of a turtle.9

9 W. C. L. Martin, ' Nat. Hist, of Mammalia,' 1841, p. 405.

44 THE DESCENT OF MAN. Part I.

The principle of Imitation is strong in man, and
especially in man in a barbarous state. Desor10 has
remarked that no animal voluntarily imitates an action
performed by man, until in the ascending scale we
come to monkeys, which are well-known to be ridiculous
mockers. Animals, however, sometimes imitate each
others' actions : thus two species of wolves, which had
been reared by dogs, learned to bark, as does some-
times the jackal,11 but whether this can be called volun-
tary imitation is another question. From one account
which I have read, there is reason to believe that puppies
nursed by cats sometimes learn to lick their feet and
thus to clean their faces : it is at least certain, as I hear
from a perfectly trustworthy friend, that some dogs
behave in this manner. Birds imitate the songs of their
parents, and sometimes those of other birds ; and par-
rots are notorious imitators of any sound which they
often hear.

Hardly any faculty is more important for the intel-
lectual progress of man than the power of Attention.
Animals clearly manifest this power, as when a cat
watches by a hole and prepares to spring on its prey.
Wild animals sometimes become so absorbed when thus
engaged, that they may be easily approached. Mr.
Bartlett has given me a curious proof how variable this
faculty is in monkeys. A man who trains monkeys to
act used to purchase common kinds from the Zoological
Society at the price of five pounds for each ; but he
offered to give double the price, if he might keep three
or four of them for a few days, in order to select one.
When asked how he could possibly so soon learn whether

10 Quoted by Vogt, ' Memoire sur les Microce'pkales,' 1867, p. 168.

11 ' The Variation of Animals and Plants under Domestication,' vol.
i. p. 27.

Chap. II. MENTAL POWERS. 45

a particular monkey would turn out a good actor, he
answered that it all depended on their power of atten-
tion. If when he was talking and explaining anything
to a monkey, its attention was easily distracted, as by
a fly on the wall or other trifling object, the case was
hopeless. If he tried by punishment to make an in-
attentive monkey act, it turned sulky. On the other
hand, a monkey which carefully attended to him could
always be trained.

It is almost superfluous to state that animals have
excellent Memories for persons and places. A baboon
at the Cape of Good Hope, as I have been informed by
Sir Andrew Smith, recognised him with joy after an
absence of nine months. I had a dog who was savage
and averse to all strangers, and I purposely tried his
memory after an absence of five years and two days. I
went near the stable where he lived, and shouted to
him in my old manner ; he showed no joy, but in-
stantly followed me out walking and obeyed me,
exactly as if I had parted with him only half-an-hour
before. A train of old associations, dormant during five
years, had thus been instantaneously awakened in his
mind. Even ants, as P. Huber 12 has clearly shewn,
recognised their fellow-ants belonging to the same com-
munity after a separation of four months. Animals
can certainly by some means judge of the intervals of
time between recurrent events.

The Imagination is one of the highest prerogatives
of man. By this faculty he unites, independently of
the will, former images and ideas, and thus creates bril-
liant and novel results. A poet, as Jean Paul Richter
remarks,13 " who must reflect whether he shall make a

12 'Les Moeurs des Fourmis,' 1810, p. 150.

13 Quoted in Dr. Maudsley's ' Physiology and Pathology of Mind,'
186S, pp. 19, 220.

46 THE DESCENT OF MAN. Part I.

" character say yes or no — to the devil with him ; he is
" only a stupid corpse." Dreaming gives us the best
notion of this power ; as Jean Paul again says, " The
" dream is an involuntary art of poetry." The value of
the products of our imagination depends of course on
the number, accuracy, and clearness of our impressions ;
on our judgment and taste in selecting or rejecting the
involuntary combinations, and to a certain extent on
our power of voluntarily combining them. As dogs,
cats, horses, and probably all the higher animals, even
birds, as is stated on good authority,14 have vivid dreams,
and this is shewn by their movements and voice, we must
admit that they possess some power of imagination.

Of all the faculties of the human mind, it will, I
presume, be admitted that Beason stands at the summit.
Few persons any longer dispute that animals possess
some power of reasoning. Animals may constantly be
seen to pause, deliberate, and resolve. It is a significant
fact, that the more the habits of any particular animal
are studied by a naturalist, the more he attributes to
reason and the less to unlearnt instincts.15 In future
chapters we shall see that some animals extremely low in
the scale apparently display a certain amount of reason.
No doubt it is often difficult to distinguish between the
power of reason and that of instinct. Thus Dr. Hayes,
in his work on * The Open Polar Sea,' repeatedly re-
marks that his dogs, instead of continuing to draw the
sledges in a compact body, diverged and separated when
they came to thin ice, so that their weight might be
more evenly distributed. This was often the first warn-

14 Dr. Jerdon, ' Birds of India,' vol. i. 1862, p. xxi.

15 Mr. L. H. Morgan's work on ' The American Beaver,' 1868, offers
a good illustration of this remark. I cannot, however, avoid thinking
that he goes too far in underrating the power of Instinct.

Chap. II. MENTAL POWEES. 47

ing and notice which the travellers received that the ice
was becoming thin and dangerous. Now, did the dogs
act thus from the experience of each individual, or from
the example of the older and wiser dogs, or from an
inherited habit, that is from an instinct ? This instinct
might possibly have arisen since the time, long ago,
when dogs were first employed by the natives in draw-
ing their sledges ; or the Arctic wolves, the parent-stock
of the Esquimaux dog, may have acquired this instinct,
impelling them not to attack their prey in a close pack
when on thin ice. Questions of this kind are most
difficult to answer.

So many facts have been recorded in various works
shewing that animals possess some degree of reason,
that I will here give only two or three instances, authen-
ticated by Kengger, and relating to American monkeys,
which stand low in their order. He states that when
he first gave eggs to his monkeys, they smashed them
and thus lost much of their contents ; afterwards thev
gently hit one end against some hard body, and picked
off the bits of shell with their fingers. After cutting
themselves only once with any sharp tool, they would
not touch it again, or would handle it with the greatest
care. Lumps of sugar were often given them wrapped
up in paper ; and Eengger sometimes put a live wasp
in the paper, so that in hastily unfolding it they got
stung ; after this had once happened, they always first
held the packet to their ears to detect any movement
within. Any one who is not convinced by such facts as
these, and by what he may observe with his own dogs,
that animals can reason, would not be convinced by
anything that I could add. Nevertheless I will give
one case with respect to dogs, as it rests on two distinct
observers, and can hardly depend on the modification
of any instinct.

48 THE DESCENT OF MAN. Part I.

Mr. Colquhoun 16 winged two wild-ducks, which fell
on the opposite side of a stream ; his retriever tried to
bring over both at once, but could not succeed ; she
then, though never before known to ruffle a feather,
deliberately killed one, brought over the other, and re-
turned for the dead bird. Col. Hutchinson relates that
two partridges were shot at once, one being killed, the
other wounded ; the latter ran away, and was caught by
the retriever, who on her return came across the dead
bird ; " she stopped, evidently greatly puzzled, and
" after one or two trials, finding she could not take it up
" without permitting the escape of the winged bird, she
" considered a moment, then deliberately murdered it
" by giving it a severe crunch, and afterwards brought
" away both together. This was the only known in-
" stance of her ever having wilfully injured any game."
Here we have reason, though not quite perfect, for the
retriever might have brought the wounded bird first
and then returned for the dead one, as in the case of
the two wild-ducks.

The muleteers in S. America say, "I will not give
" you the mule whose step is easiest, but la mas rational,
" — the one that reasons best ; ,: and Humboldt 17 adds,
"this popular expression, dictated by long experience,
" combats the system of animated machines, better per-
" haps than all the arguments of speculative j)hilosophy."

It has, I think, now been shewn that man and the
higher animals, especially the Primates, have some few
instincts in common. All have the same senses, intui-
tions and sensations — similar passions, affections, and
emotions, even the more complex ones ; they feel

16 'The Moor and the Loch,' p. 45. Col. Hutchinson on ' Dog
Breaking,' 1850, p. 46.

17 'Personal Narrative,' Eng. translat., vol. iii. p. 106.

Chap. II. MENTAL POWERS. 49

wonder and curiosity ; they possess the same faculties
of imitation, attention, memory, imagination, and reason,
though in very different degrees. Nevertheless many
authors have insisted that man is separated through his
mental faculties by an impassable barrier from all the
lower animals. I formerly made a collection of above
a score of such aphorisms, but they are not worth
giving, as their wide difference and number prove the
difficulty, if not the impossibility, of the attempt. It
has been asserted that man alone is capable of progres-
sive improvement ; that he alone makes use of tools or
fire, domesticates other animals, possesses property, or
employs language ; that no other animal is self-con-
scious, comprehends itself, has the power of abstraction,
or possesses general ideas ; that man alone has a sense
of beauty, is liable to caprice, has the feeling of grati-
tude, mystery, &c. ; believes in God, or is endowed with
a conscience. I will hazard a few remarks on the more
important and interesting of these points.

Archbishop Sumner formerly maintained 18 that man
alone is capable of progressive improvement. With
animals, looking first to the individual, every one who
has had any experience in setting traps knows that
young animals can be caught much more easily than
old ones ; and they can be much more easily approached
by an enemy. Even with respect to old animals, it is
impossible to catch many in the same place and in the
same kind of trap, or to destroy them by the same kind
of poison ; yet it is improbable that all should have
partaken of the poison, and impossible that all should
have been caught in the trap. They must learn caution
by seeing their brethren caught or poisoned. In North
America, where the fur-bearing animals have long been

18 Quoted by Sir C. Lyell, 'Antiquity of Man,' p. 497.
VOL. I. E

50 THE DESCENT OF MAN. Part I.

pursued, they exhibit, according to the unanimous tes-
timony of all observers, an almost incredible amount
of sagacity, caution, and cunning ; but trapping has
been there so long carried on that inheritance may have
come into play.

If we look to successive generations, or to the race,
there is no doubt that birds and other animals gradually
both acquire and lose caution in relation to man or
other enemies ; 19 and this caution is certainly in chief
part an inherited habit or instinct, but in part the result
of individual experience. A good observer, Leroy,20
states that in districts where foxes are much hunted,
the vouno* when thev first leave their burrows are in-
contestably much more wary than the old ones in dis-
tricts where they are not much disturbed.

Our domestic dogs are descended from wolves and
jackals,21 and though they may not have gained in
cunning, and may have lost in waryness and suspicion,
yet they have progressed in certain moral qualities,
such as in affection, trust-worthiness, temper, and pro-
bably in general intelligence. The common rat has
conquered and beaten several other species through-
out Europe, in parts of North America, New Zealand,
and recently in Formosa, as well as on the mainland of
China. Mr. Swinhoe,22 who describes these latter cases,
attributes the victory of the common rat over the large
Mus coninga to its 'superior cunning; and this latter
quality may be attributed to the habitual exercise of
all its faculties in avoiding extirpation by man, as well

19 ' Journal of Researches during the Voyage of the " Beagle," ' 184f>,
p. 398. ' Origin of Species,' 5th edit. p. 260.

20 ' Lettres Phil, sur l'lntelligence des Animaux,' nouvelle edit.
1802, p. 86.

21 See the evidence on this head in chap. i. vol. i. ' On the Variation
of Animals and Plants under Domestication.'

22 ' Proc. Zoolog. Soc.' 1864, p. 186.

Chap. II. MENTAL POWEKS. 51

as to nearly all the less cunning or weak-minded rats
having been successively destroyed by him. To main-
tain, independently of any direct evidence, that no
animal during the course of ages has progressed in
intellect or other mental faculties, is to beg the question
of the evolution of species. Hereafter we shall see that,
according to Lartet, existing mammals belonging to
several orders have larger brains than their ancient
tertiary prototypes.

It lias often been said that no animal uses any tool ;
but the chimpanzee in a state of nature cracks a native
fruit, somewhat like a walnut, with a stone.23 Bengger24
easily taught an American monkey thus to break open
hard palm-nuts, and afterwards of its own accord it
used stones to open other kinds of nuts, as well as
boxes. It thus also removed the soft rind of fruit that
had a disagreeable flavour. Another monkey was taught
to open the lid of a large box with a stick, and after-
wards it used the stick as a lever to move heavv bodies ;
and I have myself seen a young orang put a stick into
a crevice, slip his hand to the other end, and use it in
the proper manner as a lever. In the cases just men-
tioned stones and sticks were employed as implements ;
but they are likewise used as weapons. Brehm 25 states,
on the authority of the well-known traveller Schimper,
that in Abyssinia when the baboons belonging to one
species (C. gelada) descend in troops from the moun-
tains to plunder the fields, they sometimes encounter
troops of another species (C. hamadryas), and then a
fight ensues. The Geladas roll down great stones, which
the Hamadryas try to avoid, and then both species,

23 Savage and Wyman in 'Boston Journal of Nat. Hist.' vol. iv.
]f43-44, p. 383.

24 ' Siiugethiere von Paraguay,' 1830, s. 51-56.

25 ' Thierleben,' B. i. s. 79, 82.

E 2

52 THE DESCENT OF MAN. Pabt I.

making a great uproar, rush furiously against each
other. Brehm, when accompanying the Duke of Coburg-
Gotha, aided in an attack with fire-arms on a troop of
baboons in the pass of Mensa in Abyssinia. The baboons
in return rolled so many stones down the mountain,
some as large as a man's head, that the attackers had
to beat a hasty retreat ; and the pass was actually for
a time closed against tlie caravan. It deserves notice
that these baboons thus acted in concert. Mr. Wal-
lace26 on three occasions saw female orangs, accom-
panied by their young, " breaking off branches and
" the great spiny fruit of the Durian tree, with every
" appearance of rage ; causing such a shower of missiles
" as effectually kept us from approaching too near the
" tree."

In the Zoological Gardens a monkev which had weak
teeth used to break open nuts with a stone ; and I was
assured by the keepers that this animal, after using the
stone, hid it in the straw, and would not let any other
monkey touch it. Here, then, we have the idea of
property ; but this idea is common to every dog with a
bone, and to most or all birds with their nests.

The Duke of Argyll 27 remarks, that the fashioning of
an implement for a special purpose is absolutely peculiar
to man ; and he considers that this forms an immeasur-
able gulf between him and the brutes. It is no doubt
a very important distinction, but there appears to me
much truth in Sir J. Lubbock's suggestion,28 that when
primeval man first used flint-stones for any purpose, he
would have accidentally splintered them, and would
then have used the sharp fragments. From this step
it would be a small one to intentionally break the

26 ' The Malay Archipelago,' vol. i. 1869, p. 87.

27 ' Primeval Man,' 1869, pp. 145, 147.

28 ' Prehistoric Times,' 1865, p. 473, &c.

Chap. II. MENTAL POWERS. 53

flints, and not a very wide step to rudely fashion them.
This latter advance, however, may have taken long
ages, if we may judge by the immense interval of time
which elapsed before the men of the neolithic period
took to grinding and polishing their stone tools. In
breaking the flints, as Sir J. Lubbock likewise remarks,
sparks would have been emitted, and in grinding them
heat would have been evolved : " thus the two usual
" methods of obtaining fire may have originated." The
nature of fire would have been known in the many
volcanic regions where lava occasionally flows through
forests. The anthropomorphous apes, guided probably
by instinct, build for themselves temporary platforms ;
but as many instincts are largely controlled by reason,
the simpler ones, such as this of building a platform,
might readily pass into a voluntary and conscious act.
The orang is known to cover itself at night with the
leaves of the Pandanus ; and Brehm states that one of
his baboons used to protect itself from the heat of the
sun by throwing a straw-mat over its head. In these
latter habits, we probably see the first steps towards
some of the simpler arts ; namely rude architecture
and dress, as they arose amongst the early progenitors
of man.

Language. — This faculty has justly been considered as
one of the chief distinctions between man and the lower
animals. But man, as a highly competent judge, Arch-
bishop Whately remarks, " is not the only animal that
" can make use of language to express what is passing in
" his mind, and can understand, more or less, what is so
" expressed by another." 29 In Paraguay the Cebus azarse
when excited utters at least six distinct sounds, which

29 Quoted in ' Anthropological Review,' 18G4, p. 158.

5-4 THE DESCENT OF MAN. Part I.

excite in other monkeys similar emotions.30 The move-
ments of the features and gestures of monkeys are un-
derstood by us, and they partly understand ours, as
Eeno-o-er and others declare. It is a more remark-

DO

able fact that the dog, since being domesticated, has
learnt to bark 31 in at least four or five distinct tones.
Although barking is a new art, no doubt the wild spe-
cies, the parents of the dog, expressed their feelings
by cries of various kinds. With the domesticated
dog we have the bark of eagerness, as in the chase ;
that of anger ; the yelping or howling bark of despair,
as when shut up ; that of joy, as when starting on a
walk with his master ; and the very distinct one of
demand or supplication, as when wishing for a door or
window to be opened.

Articulate language is, however, peculiar to man ;
but he uses in common with the lower animals inarti-
culate cries to express his meaning, aided by gestures
and the movements of the muscles of the face.32 This
especially holds good with the more simple and vivid
feelings, which are but little connected with our higher
intelligence. Our cries of pain, fear, surprise, anger, to-
gether with their appropriate actions, and the murmur
of a mother to her beloved child, are more expressive
than any words. It is not the mere power of articula-
tion that distinguishes man from other animals, for as
every one knows, parrots can talk ; but it is his large
power of connecting definite sounds with definite ideas ;
and this obviously depends on the development of the
mental faculties.

30 Kenggcr, ibid. s. 45.

31 See my ' Variation of Animals and Plants under Domestication,'
vol. i. p. 27.

32 See a discussion on this subject in Mr. E. B. Tylor's very interest-
ing work, 'Eesearches into the Early History of Mankind,' 1865, chaps,
ii. to iv.

Chap. II. MENTAL POWERS. 55

As Home Tooke, one of the founders of the noble
science of philology, observes, language is an art, like
brewing or baking ; but writing would have been a
much more appropriate simile. It certainly is not a
true instinct, as every language has to be learnt. It
differs, however, widely from all ordinary arts, for man
has an instinctive tendency to speak, as we see in the
babble of our young children ; whilst no child has an
instinctive tendency to brew, bake, or write. Moreover,
no philologist now supposes that any language has
been deliberately invented ; each has been slowly and
unconsciously developed by many steps. The sounds
uttered by birds offer in several respects the nearest
analogy to language, for all the members of the same
species utter the same instinctive cries expressive of
their emotions ; and all the kinds that have the power
of singing exert this power instinctively ; but the actual
song, and even the call-notes, are learnt from their
parents or foster-parents. These sounds, as Daines
Barrington 33 has proved, " are no more innate than
" language is in man." The first attempts to sing
" may be compared to the imperfect endeavour in a
" child to babble." The young males continue prac-
tising, or, as the bird-catchers say, recording, for ten
or eleven months. Their first essays show hardly a
rudiment of the future song; but as they grow older
we can perceive what they are aiming at ; and at last
they are said " to sing their song round." Nestlings
which have learnt the song of a distinct species, as
with the canary-birds educated in the Tyrol, teacli and
transmit their new song to their offspring. The slight
natural differences of song in the same species inha-

33 Hon. Daines Barrington in ' Philosoph. Transactions,' 1773, p.
262. See also Dureau de la Malle, in ' Ann. des Sc. Nat.' 3rd series,
Zoolog. torn. x. p. 119.

56 THE DESCENT OF MAN. Part I.

biting different districts may be appositely compared,
as Barrington remarks, " to provincial dialects ;" and
the songs of allied, though distinct species may be com-
pared with the languages of distinct races of man. I
have given the foregoing details to shew that an in-
stinctive tendency to acquire an art is not a peculiarity
confined to man.

With respect to the origin of articulate language,
after having read on the one side the highly interesting
works of Mr. Hensleigh Wedgwood, the Kev. F. Farrar,
and Prof. Schleicher,34 and the celebrated lectures of
Prof. Max Miiller on the other side, I cannot doubt that
language owes its origin to the imitation and mo-
dification, aided by signs and gestures, of various
natural sounds, the voices of other animals, and man's
own instinctive cries. When we treat of sexual selec-
tion we shall see that primeval man, or rather some
early progenitor of man, probably used his voice largely,
as does one of the gibbon-apes at the present day, in
producing true musical cadences, that is in singing ;
we may conclude from a widely -spread analogy that
this power would have been especially exerted during
the courtship of the sexes, serving to express various
emotions, as love, jealousy, triumph, and serving as a
challenge to their rivals. The imitation by articulate
sounds of musical cries might have given rise to
words expressive of various complex emotions. As
bearing on the subject of imitation, the strong tendency
in our nearest allies, the monkeys, in microcephalous

34 ' On the Origin of Language/ by H. Wedgwood, 1866. ' Chapters
on Language,' by the Kev. F. W. Farrar, 1865. These works are most
interesting. See also ' De la Phys. et de Parole,' par Albert Lemoine,
1865, p. 190. The work on this subject, by the late Prof. Aug. Schlei-
cher, has been translated by Dr. Bikkers into English, under the title
of ' Darwinism tested by the Science of Language/ 1869.

Chap. II. MENTAL POWEES. 57

idiots,35 and in the barbarous races of mankind, to imi-
tate whatever they hear deserves notice. As monkeys
certainly understand much that is said to them by man,
and as in a state of nature they utter signal-cries of
danger to their fellows,36 it does not appear altogether
incredible, that some unusually wise ape-like animal
should have thought of imitating the growl of a beast
of prey, so as to indicate to his fellow monkeys the
nature of the expected danger. And this would have
been a first step in the formation of a language.

As the voice was used more and more, the vocal
organs would have been strengthened and perfected
through the principle of the inherited effects of use ;
and this would have reacted on the power of speech.
But the relation between the continued use of language
and the development of the brain has no doubt been far
more important. The mental powers in some early pro-
genitor of man must have been more highly developed
than in any existing ape, before even the most imperfect
form of speech could have come into use ; but we may
confidently believe that the continued use and .advance-
ment of this power would have reacted on the mind by
enabling and encouraging it to carry on long trains of
thought. A long and complex train of thought can no
more be carried on wdthout the aid of words, whether
spoken or silent, than a long calculation without the
use of figures or algebra. It appears, also, that even
ordinary trains of thought almost require some form of
language, for the dumb, deaf, and^ blind girl, Laura
Bridgman, was observed to use her fingers whilst dream-

35 Vogt, 'Memoire sur les Microcephales,' 1867, p. 169. With
respect to savages, I have given some facts in my ' Journal of
Eesearches,' &c, 1845, p. 206.

36 See clear evidence on this head in the two works so often quoted,
by Brehm and Kengger.

58 THE DESCENT OF MAN. Part I.

ing.37 Nevertheless a long succession of vivid and con-
nected ideas, may pass through the mind without the
aid of any form of language, as we may infer from the
prolonged dreams of dogs. We have, also, seen that
retriever-dogs are able to reason to a certain extent ;
and this they manifestly do without the aid of language.
The intimate connection between the brain, as it is
now developed in us, and the faculty of speech, is well
shewn by those curious cases of brain-disease, in which
speech is specially affected, as when the power to re-
member substantives is lost, whilst other words can be
correctly used.38 There is no more improbability in
the effects of the continued use of the vocal and mental
organs being inherited, than in the case of hand-
writing, which depends partly on the structure of the
hand and partly on the disposition of the mind ; and
hand-writing is certainly inherited.39

Why the organs now used for speech should have
been originally perfected for this purpose, rather than
any other organs, it is not difficult to see. Ants have
considerable powers of intercommunication by means
of their antennae, as shewn by Huber, who devotes a
whole chapter to their language. We might have used
our fingers as efficient instruments, for a person wTith
practice can report to a deaf man every word of a speech
rapidly delivered at a public meeting ; but the loss of
our hands, whilst thus employed, would have been
a serious inconvenience. As all the higher mammals
possess vocal organs constructed on the same general

37 See remarks on this head hy Dr. Maudsley, 'The Physiology
and Pathology of Mind,' 2nd edit. 1868, p. 199.

38 Many curious cases have been recorded. See, for instance,
' Inquiries Concerning the Intellectual Powers/ by Dr. Aberciombie,
1838, p. 150.

39 ' The Variation of Animals and Plants under Domestication,' vol.
ii. p. 6.

Chap. II. MENTAL POWERS. 59

plan with ours, and which are used as a means of commu-
nication, it was obviously probable, if the power of com-
munication had to be improved, that these same organs
would have been still further developed ; and this has
been effected by the aid of adjoining and well-adapted
parts, namely the tongue and lips.40 The fact of the
higher apes not using their vocal organs for speech, 1:0
doubt depends on their intelligence not having been
sufficiently advanced. The possession by them of organs,
which with long-continued practice might have been
used for speech, although not thus used, is paralleled by
the case of many birds which possess organs fitted for
singing, though they never sing. Thus, the nightingale
and crow have vocal organs similarly constructed, these
being used by the former for diversified song, and by
the latter merely for croaking.41

The formation of different languages and of dis-
tinct species, and the proofs that both have been de-
veloped through a gradual process, are curiously the
same.42 But we can trace the origin of many words
further back than in the case of species, for we can
perceive how they have actually arisen from the
imitation of various sounds. We find in distinct
languages striking homologies due to community of
descent, and analogies due to a similar process of

40 See some good remarks to this effect by Dr. Maudsley, ' The
Physiology and Pathology of Mind,' 1868, p. 199.

41 Macgillivray, 'Hist, of British Birds,' vol. ii. 1839, p. 29. An
excellent observer, Mr. Blackwall, remarks that the magpie learns to
pronounce single words, and even short sentences, more readily than
ulmost any other British bird ; yet, as he adds, after long and closely
investigating its habits, he has never known it, in a state of nature,
display any unusual capacity for imitation. ' Kesearches in Zoology,'
1834, p. 158.

42 See the very interesting parallelism between the deveh
species and languages, given by Sir C. Lyell in ' The GeoloafT J^v)«en^is ^ > ^
of the Antiquity of Man,' 1 803, chap, xxiii. /^ O <

f£j i* ^*^ -5

LIBRARY

60 THE DESCENT OF MAN. Part I.

formation. The manner in which certain letters or
sounds change when others change is very like corre-
lated growth. We have in both cases the reduplication
of parts, the effects of long-continued use, aud so forth.
The frequent presence of rudiments, both in languages
and in species, is still more remarkable. The letter m
in the word am, means I ; so that in the expression lam,
a superfluous and useless rudiment has been retained.
In the spelling also of words, letters often remain
as the rudiments of ancient forms of pronunciation.
Languages, like organic beings, can be classed in groups
under groups ; and they can be classed either naturally
according to descent, or artificially by other characters.
Dominant languages and dialects spread widely and
lead to the gradual extinction of other tongues. A lan-
guage, like a species, when once extinct, never, as Sir
C. Lyell remarks, reappears. The same language never
has two birth-places. Distinct languages may be crossed
or blended together.43 We see variability in every
tongue, and new words are continually cropping up ; but
as there is a limit to the powers of the memory, single
words, like whole languages, gradually become extinct.
As Max Muller44 has well remarked: — " A struggle for
" life is constantly going on amongst the words and gram-
"matical forms in each language. The better, the
" shorter, the easier forms are constantly gaining the
" upper hand, and they owe their success to their own
" inherent virtue." To these more important causes of
the survival of certain words, mere novelty may, I
think, be added ; for there is in the mind of man a strong
love for slight changes in all things. The survival or

43 See remarks to this effect by the Rev. F. W. Farrar, in an interest-
ing article, entitled " Philology and Darwinism " in ' Nature,' March
24th, 1870, p. 528.

44 ' Nature,' Jan. 6th, 1870, p. 257.

Chap. II. MENTAL POWERS. Gl

preservation of certain favoured words in the struggle
for existence is natural selection.

The perfectly regular and wonderfully complex con-
struction of the languages of many barbarous nations
has often been advanced as a proof, either of the divine
origin of these languages, or of the high art and former
civilisation of their founders. Thus F. von Schlegel
writes : " In those languages which appear to be at the
" lowest grade of intellectual culture, we frequently ob-
" serve a very high and elaborate degree of art in their
" grammatical structure. This is especially the case with
" the Basque and the Lapponian, and many of the Ame-
" rican languages." 45 But it is assuredly an error to speak
of any language as an art in the sense of its having
been elaborately and methodically formed. Philolo-
gists now admit that conjugations, declensions, &c, ori-
ginally existed as distinct words, since joined together ;
and as such words express the most obvious relations
between objects and persons, it is not surprising that
they should have been used by the men of most races
during the earliest ages. With respect to perfection,
the following illustration will best shew how easily we
may err : a Crinoid sometimes consists of no less
than 150,000 pieces of shell,40 all arranged with per-
fect symmetry in radiating lines ; but a naturalist does
not consider an animal of this kind as more perfect
than a bilateral one with comparatively few parts,
and with none of these alike, excepting on the opposite
sides of the body. He iustlv considers the differen-
tiation and specialisation of organs^ as the test of per-
fection. So with languages, the most symmetrical and
complex ought not to be ranked above irregular, abbre-

45 Quoted by C. S. Wake, ' Chapters on Man,' 1868, p. 101.

46 Buckland, ' Bridgewater Treatise,' p. 411.

62 THE DESCENT OF MAN. Part I.

viated, and bastardised languages, which have borrowed
expressive words and useful forms of construction from
various conquering, or conquered, or immigrant races.

From these few and imperfect remarks I conclude
that the extremely complex and regular construction of
many barbarous languages, is no proof that they owe
their origin to a special act of creation.47 Nor, as
we have seen, does the faculty of articulate speech in
itself offer any insuperable objection to the belief that
man has been developed from some lower form.

Self -consciousness, Individuality, Abstraction, General
Ideas, &c. — It would be useless to attempt discussing
these high faculties, which, according to several recent
writers, make the sole and complete distinction between
man and the brutes, for hardly two authors agree in their
definitions. Such faculties could not have been fully
developed in man until his mental powers had advanced
to a high standard, and this implies the use of a perfect
language. No one supposes that one of the lower ani-
mals reflects whence he comes or whither he goes, —
what is death or what is life, and so forth. But can
we feel sure that an old dog with an excellent memory
and some power of imagination, as shewn by his
dreams, never reflects on his past pleasures in the
chase ? and this would be a form of self-consciousness.
On the other hand, as Biiehner48 has remarked, how
little can the hard-worked wife of a degraded Australian
savage, who uses hardly any abstract words and cannot
count above four, exert her self-consciousness, or reflect
on the nature of her own existence.

47 See some good remarks on the simplification of languages, by Sir
J. Lubbock, ' Origin of Civilisation,' 1870, p. 278.

48 ' Conferences sur la The'orie Darwinienne,' French translat., 1869,
p. 132.

Chap. II. MENTAL POWERS. 63

That animals retain their mental individuality is
unquestionable. When my voice awakened a train of
old associations in the mind of the above-mentioned
dog. he must have retained his mental individuality,
although every atom of his brain had probably under-
gone change more than once during the interval of five
years. This dog might have brought forward the
argument lately advanced to crush all evolutionists,
and said, "I abide amid all mental moods and all
" material changes. . . . The teaching that atoms leave
" their impressions as legacies to other atoms falling
" into the places they have vacated is contradictory of
" the utterance of consciousness, and is therefore false ;
" but it is the teaching necessitated by evolutionism,
" consequently the hypothesis is a false one." 4ft

Sense of Beauty. — This sense has been declared to
be peculiar to man. Bat when we behold male birds
elaborately displaying their plumes and splendid colours
before the females, whilst other birds not thus deco-
rated make no such display, it is impossible to doubt
that the females admire the beauty of their male part-
ners. As women everywhere deck themselves with
these plumes, the beauty of such ornaments cannot be
disputed. The Bower-birds by tastefully ornamenting
their playing-passages with gaily-coloured objects, as
do certain humming-birds their nests, offer additional
evidence that they possess a sense of beauty. So with
the song of birds, the sweet strains poured forth by the
males during the season of love are "certainly admired
by the females, of which fact evidence will hereafter be
given. If female birds had been incapable of appre-
ciating the beautiful colours, the ornaments, and voices

49

The Rev. Dr. J. M'Cann, ' Anti-Darwinism,' 1869, p. 13.

64 THE DESCENT OF MAN. Part I.

of their male partners, all the labour and anxiety exhi-
bited by them in displaying their charms before the
females would have been thrown away ; and this it is
impossible to admit. Why certain bright colours and
certain sounds should excite pleasure, when in harmony,
cannot, I presume, be explained any more than why
certain flavours and scents are agreeable ; but assuredly
the same colours and the same sounds are admired by
us and by many of the lower animals.

The taste for the beautiful, at least as far as female
beauty is concerned, is not of a special nature in the
human mind ; for it differs widely in the different
races of man, as will hereafter be shewn, and is not
quite the same even in the different nations of the
same race. Judging from the hideous ornaments and
the equally hideous music admired by most savages, it
might be urged that their aesthetic faculty was not so
highly developed as in certain animals, for instance, in
birds. Obviously no animal would be capable of ad-
miring such scenes as the heavens at night, a beautiful
landscape, or refined music; but such high tastes, de-
pending as they do on culture and complex associa-
tions, are not enjoyed by barbarians or by uneducated
persons.

Many of the faculties, which have been of inesti-
mable service to man for his progressive advance-
ment, such as the powers of the imagination, wonder,
curiosity, an undefined sense of beauty, a tendency
to imitation, and the love of excitement or novelt ,
could not fail to have led to the most capricious
changes of customs and fashions. I have alluded to
this point, because a recent writer50 has oddly fixed
on Caprice " as one of the most remarkable a d

50 • The Spectator,' Dec. 4th, 1869, p. 1430.

Chap. II. MENTAL POWERS. 65

" typical differences between savages and brutes." But
not only can we perceive how it is that man is capri-
cious, but the lower animals are, as we shall hereafter
see, capricious in their affections, aversions, and sense
of beauty. There is also good reason to suspect that
they love novelty, for its own sake.

Belief in God — Religion. — There is no evidence that
man was aboriginally endowed with the ennobling
belief in the existence of an Omnipotent God. On the
contrary there is ample evidence, derived not from hasty
travellers, but from men who have long resided with
savages, that numerous races have existed and still
exist, who have no idea of one or more gods, and who
have no words in their languages to express such an
idea.51 The question is of course wholly distinct from
that higher one, whether there exists a Creator and
Ruler of the universe ; and this has been answered in
the affirmative bv the highest intellects that have ever
lived.

If, however, we include under the term " religion " the
belief in unseen or spiritual agencies, the case is wholly
different; for this belief seems to be almost universal
with the less civilised races. Nor is it difficult to
comprehend how it arose. As soon as the important
faculties of the imagination, wonder, and curiosity,
together with some power of reasoning, had become
partially developed, man would naturally have craved
to understand what was passing around him, and
have vaguely speculated on his own existence. As

51 See an excellent article on this subject by the Eev. F W. Farrar,
in the ' Anthropological Review,' Aug. 1864, p. ccxvii. For further
facts see Sir J. Lubbock, ' Prehistoric Times,' 2nd edit. 1869, p. 564 ;
and especially the chapters on Religion in his ' Origin of Civilisation.'
1870.

VOL. I. F

66 THE DESCENT OF MAN. Part I.

Mr. M'Lennan 52 has remarked, " Some explanation of
" the phenomena of life, a man must feign for himself ;
" and to judge from the universality of it, the simplest
" hypothesis, and the first to occur to men, seems to have
" been that natural phenomena are ascribable to the pre-
" sence in animals, plants, and things, and in the forces
" of nature, of such spirits prompting to action as men
" are conscious they themselves possess." It is probable,
as Mr. Tylor has clearly shewn, that dreams may have
first given rise to the notion of spirits ; for savages do
not readily distinguish between subjective and objective
impressions. When a savage dreams, the figures which
appear before him are believed to have come from a
distance and to stand over him; or "the soul of the
" dreamer goes out on its travels, and comes home with
" a remembrance of what it has seen." 53 But until the
above-named faculties of imagination, curiosity, reason,
&c, had been fairly well developed in the mind of man,
his dreams would not have led him to believe in spirits,
any more than in the case of a dog.

32 The Worship of Animals and Plants, in the ' Fortnightly Keview,'
Oct. 1, 1869, p. 422.

53 Tylor, 'Early History of Mankind,' 1865, p. 6. See also the three
striking chapters on the Development of Eeligion, in Lubbock's ' Origin
of Civilisation,' 1870. In a like manner Mr. Herbert Spencer, in his
ingenious essay in the ' Fortnightly Review' (May 1st, 1870, p. 535),
accounts for the earliest forms of religious belief throughout the world,
by man being led through dreams, shadows, and other causes, to look
at himself as a double essence, corporeal and spiritual. As the spiritual
being is supposed to exist after death and to be powerful, it is propi-
tiated by various gifts and ceremonies, and its aid invoked. He then
further shews that names or nicknames given from some animal or
other object to the early progenitors or founders of a tribe, are sup-
posed after a long interval to represent the real progenitor of the tribe ;
and such animal or object is then naturally believed still to exi^t as
a spirit, is held sacred, and worshipped as a god. Nevertheless I cannot
but suspect that there is a still earlier and ruder stage, when anything
which manifests power or movement is thought to be endowed with
some form of life, and with mental faculties analogous to our own.

Chap. II. MENTAL POWERS. 67

The tendency in savages to imagine that natural
objects and agencies are animated by spiritual or living
essences, is perhaps illustrated by a little fact which I
once noticed : my dog, a full-grown and very sensible
animal, was lying on the lawn during a hot and still
day ; but at a little distance a slight breeze occasionally
moved an open parasol, which would have been wholly
disregarded by the dog, had any one stood near it. As
it was, every time that the parasol slightly moved, the
dog growled fiercely and barked. He must, I think,
have reasoned to himself in a rapid and unconscious
manner, that movement without any apparent cause
indicated the presence of some strange living agent, and
no stranger had a right to be on his territory.

The belief in spiritual agencies would easily pass into
the belief in the existence of one or more gods. For
savages would naturally attribute to spirits the same
passions, the same love of vengeance or simplest form
of justice, and the same affections which they themselves
experienced. The Fuegians appear to be in this respect
in an intermediate condition, for when the surgeon on
board the " Beagle ': shot some young ducklings as
specimens, York Minster declared in the most solemn
manner, " Oh ! Mr. Bynoe, much rain, much snow, blow
" much ;" and this was evidently a retributive punish-
ment for wasting human food. So again he related
how, when his brother killed a " wild man," storms long
raged, much rain and snow fell. Yet we could never
discover that the Fuegians believed in what we should
call a God, or practised any religious rites ; and Jemmy
Button, with justifiable pride, stoutly maintained that
there was no devil in his land. This latter assertion is
the more remarkable, as with savages the belief in
bad spirits is far more common than the belief in good
spirits.

f 2

68 THE DESCENT OF MAN. Part I.

The feeling of religious devotion is a highly com-
plex one, consisting of love, complete submission to
an exalted and mysterious superior, a strong sense of
dependence,54 fear, reverence, gratitude, hope for the
future, and perhaps other elements. No being could
experience so complex an emotion until advanced in
his intellectual and moral faculties to at least a mode-
rately high level. Nevertheless we see some distant
approach to this state of mind, in the deep love of a
dog for his master, associated with complete submission,
some fear, and perhaps other feelings. The behaviour
of a dog when returning to his master after an ab-
sence, and, as I may add, of a monkey to his beloved
keeper, is widely different from that towards their
fellows. In the latter case the transports of joy
appear to be somewhat less, and the sense of equality
is shewn in every action. Professor Braubach55 goes
so far as to maintain that a dog looks on his master as
on a god.

The same high mental faculties which first led man
to believe in unseen spiritual agencies, then in fetish-
ism, polytheism, and ultimately in monotheism, would
infallibly lead him, as long as his reasoning powers
remained poorly developed, to various strange super-
stitions and customs. Many of these are terrible to
think of — such as the sacrifice of human beings to a
blood-loving god ; the trial of innocent persons by the
ordeal of poison or fire ; witchcraft, &c. — yet it is well
occasionally to reflect on these superstitions, for they
shew us what an infinite debt of gratitude we owe to
the improvement of our reason, to science, and our

54 See an able article on the Psychical Elements of Keligion, by
Mr. L. Owen Pike, in ' Anthropolog. Keview/ April, 1870, p. lxiii.
" ' Keligion, Moral, &c, der Darwin'schen Art-Lehre.' 1869, s. 53.

Chap. II. MENTAL POWERS. 69

accumulated knowledge.56 As Sir J. Lubbock has well
observed, "it is not too much to say that the horrible
" dread of unknown evil hansrs like a thick cloud over
" savage life, and embitters every pleasure." These
miserable and indirect consequences of our highest
faculties may be compared with the incidental and
occasional mistakes of the instincts of the lower animals.

56 'Prehistoric Times,' 2nd edit. p. 571. In this work (at p. 553)
there will be found an excellent account of the many strange and
capricious customs of savages.

70 THE DESCENT OF MAN. Part I.

CHAPTER III.

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 in-
stincts— 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 — Trans-
mission of moral tendencies — Summary.

I fully subscribe to the judgment of those writers1
who maintain that of all the differences between man
and the lower animals, the moral sense or conscience
is by far the most important. This sense, as Mack-
intosh2 remarks, "has a rightful supremacy over every
" other principle of human action ; " it is summed up
in that short but imperious word ought, so full of high
significance. It is the most noble of all the attributes
of man, leading him without a moment's hesitation
to risk his life for that of a fellow- creature; or after
due deliberation, impelled simply by the deep feeling
of right or duty, to sacrifice it in some great cause.
Immanuel Kant exclaims, " Duty ! Wondrous thought,
" that workest neither by fond insinuation, flattery, nor
" by any threat, but merely by holding up thy naked
" law in the soul, and so extorting for thyself always

1 See, for instance, on this subject, Quatrefages, ' Unite de PEspece
Humaine,' 1861, p. 21, &c.

* ' Dissertation on Ethical Philosophy,' 1837, p. 231, &c.

Chap. III. MOIIAL SENSE. 71

" reverence, if not always obedience ; before whom all
" appetites are dumb, however secretly they rebel ;
" whence thy original ? " 3

This great question has been discussed by many
writers4 of consummate ability ; and my sole excuse
for touching on it is the impossibility of here passing
it over, and because, as far as I know, no one has ap-
proached it exclusively from the side of natural history.
The investigation possesses, also, some independent in-
terest, as an attempt to see how far the study of the
lower animals can throw light on one of the highest
psychical faculties of man.

The following proposition seems to me in a high
degree probable — namely, that any animal whatever,
endowed with well-marked social instincts,5 would inevi-
tably acquire a moral sense or conscience, as soon as

3 ' Metaphysics of Ethics,' translated by J. W. Semple, Edinburgh,
1836, p. 136.

4 Mr. Bain gives a list (' Mental and Moral Science,' 1868, p. 543-
725) of twenty-six British authors who have written on this subject,
and whose names are familiar to every reader ; to these, Mr, Bain's own
name, and those of Mr. Lecky, Mr. Shadworth Hodgson, and Sir J.
Lubbock, as well as of others, may be added.

5 Sir B. Brodie, after observing that man is a social animal (' Psy-
chological Enquiries,' 1854, p. 192), asks the pregnant question,
" ought not this to settle the disputed question as to the existence of a
" moral sense ?" Similar ideas have probably occurred to many persons,
as they did long ago to Marcus Aurelius. Mr. J. S. Mill speaks, in
his celebrated work, ' Utilitarianism,' (1864, p. 46), of the social feelings
as a " powerful natural sentiment," and as " the natural basis of
" sentiment for utilitarian morality ; " but on the previous page he
says, " if, as is my own belief, the moral feelings are not innate, but
" acquired, they are not for that reason less natural." It is with hesita-
tion that I venture to differ from so profound a thinker, but it can
hardly be disputed that the social feelings are instinctive or innate
in the lower animals ; and why should they not be so in man ?
Mr. Bain (see, for instance, ' The Emotions and the Will,' 1865, p. 481)
and others believe that the moral sense is acquired by each individual
during his lifetime. On the general theory of evolution this is at
least extremely improbable.

72 THE DESCENT OF MAN. Part I.

its intellectual powers had become as well developed, or
nearly as well developed, as in man. For, firstly, the
social instincts lead an animal to take pleasure in
the society of its fellows, to feel a certain amount of
sympathy with them, and to perform various services
for them. The services may be of a definite and evi-
dently instinctive nature ; or there may be only a wish
and readiness, as with most of the higher social animals,
to aid their fellows in certain general ways. But these
feelings and services are by no means extended to all
the individuals of the same species, only to those of
the same association. Secondly, as soon as the mental
faculties had become highly developed, images of all
past actions and motives would be incessantly passing
through the brain of each individual ; and that feeling
of dissatisfaction which invariably results, as we shall
hereafter see, from any unsatisfied instinct, would arise,
as often as it was perceived that the enduring and
always present social instinct had yielded to some other
instinct, at the time stronger, but neither enduring in
its nature, nor leaving behind it a very vivid impres-
sion. It is clear that many instinctive desires, such as
that of hunger, are in their nature of short duration ;
and after being satisfied are not readily or vividly re-
called. Thirdly, after the power of language had been
acquired and the wishes of the members of the same
community could be distinctly expressed, the common
opinion how each member ought to act for the public
good, would naturally become to a large extent the guide
to action. But the social instincts would still give the
impulse to act for the good of the community, this im-
pulse being strengthened, directed, and sometimes even
deflected by public opinion, the power of which rests, as
we shall presently see, on instinctive sympathy. Lastly,
habit in the individual would ultimately play a very

Chap. III. MORAL SENSE. 73

important part in guiding the conduct of each member ;
for the social instincts and impulses, like all other in-
stincts, would be greatly strengthened by habit, as
would obedience to the wishes and judgment of the com-
munity. These several subordinate propositions must
now be discussed ; and some of them at considerable
length.

It may be well first to premise that 1 do not wish to
maintain that any strictly social animal, if its intellec-
tual faculties were to become as active and as highly
developed as in man, would acquire exactly the same
moral sense as ours. In the same manner as various
animals have some sense of beauty, though they admire
widely different objects, so they might have a sense of
right and wrong, though led by it to follow widely dif-
ferent lines of conduct. If, for instance, to take an ex-
treme case, men were reared under precisely the same
conditions as hive-bees, there can hardly be a doubt
that oar unmarried females would, like the worker-
bees, think it a sacred duty to kill their brothers, and
mothers would strive to kill their fertile daughters ;
and no one would think of interfering. Nevertheless
the bee, or any other social animal, would in our sup-
posed case gain, as it appears to me, some feeling of
right and wrong, or a conscience. For each individual
would have an inward sense of possessing certain
stronger or more enduring instincts, and others less
strong or enduring ; so that there would often be a
struggle which impulse should be followed; and satis-
faction or dissatisfaction would be felt, as past impres-
sions were compared during their incessant passage
through the mind. In this case an inward monitor
would tell the animal that it would have been better to
have followed the one impulse rather than the other.
The one course ought to have been followed : the one

74 THE DESCENT OF MAN. Part I.

would have been right and the other wrong; but to
these terms I shall have to recur.

Sociability. — Animals of many kinds are social ; we
find even distinct species living together, as with some
American inoDkeys, and with the united flocks of rooks,
jackdaws, and starlings. Man shows the same feeling
in his strong love for the dog, which the dog returns
with interest. Every one must have noticed how mise-
rable horses, dogs, sheep, &c. are when separated from
their companions ; and what affection at least the two
former kinds show on their reunion. It is curious to
speculate on the feelings of a dog, who will rest peace-
fully for hours in a room with his master or any of the
family, without the least notice being taken of him ; but
if left for a short time by himself, barks or howls dis-
mally. We will confine our attention to the higher
social animals, excluding insects, although these aid
each other in many important ways. The most common
service which the higher animals perform for each other,
is the warning each other of danger by means of the
united senses of alL Every sportsman knows, as Dr.
Jaeger remarks,6 how difficult it is to approach animals
in a herd or troop. Wild horses and cattle do not, I
believe, make any danger-signal ; but the attitude of
any one who first discovers an enemy, warns the others.
Kabbits stamp loudly on the ground with their hind-feet
as a signal : sheep and chamois do the same, but with
their fore-feet, uttering likewise a whistle. Many birds
and some mammals post sentinels, which in the case of
seals are said 7 generally to be the females. The leader
of a troop of monkeys acts as the sentinel, and utters
cries expressive both of danger and of safety.8 Social

6 ' Die Darwin'sche Theorie,' s. 101.

■ Mr. R. Brown in ' Proc. Zoolog. Soc/ 1868, p. 409.

8 Brehm, ' Thierleben,' B. i. 1864, s. 52, 79. For the case of the

Chap. III. MOEAL SENSE. 75

animals perform many little services for each other:
horses nibble, and cows lick each other, on any spot
which itches : monkeys search for each other's external
parasites ; and Brehm states that after a troop of the
Cercopithecus griseo-viridis has rushed through a thorny
brake, each monkey stretches itself on a branch, and
another monkey sitting by " conscientiously " examines
its fur and extracts every thorn or burr.

Animals also render more important services to each
other : thus wolves and some other beasts of prey hunt
in packs, and aid each other in attacking their victims.
Pelicans fish in concert. The Hamadryas baboons turn
over stones to find insects, &c. ; and when they come to
a large one, as many as can stand round, turn it over
together and share the booty. Social animals mutually
defend each other. The males of some ruminants come
to the front when there is danger and defend the herd
with their horns. I shall also in a future chapter give
cases of two young wild bulls attacking an old one in
concert, and of two stallions together trying to drive
away a third stallion from a troop of mares. Brehm
encountered in Abyssinia a great troop of baboons which
were crossing a valley : some had already ascended the
opposite mountain, and some were still in the valley :
the latter were attacked by the dogs, but the old males
immediately hurried down from the rocks, and with
mouths widely opened roared so fearfully, that the dogs
precipitately retreated. They were again encouraged
to the attack ; but by this time all the baboons had re-
ascended the heights, excepting a young one, about six

monkeys extracting thorns from each other, see s. 54. With respect to
the Hamadryas turning over stones, the fact is given (s. 76) on the
evidence of Alvarez, whose observations Brehm thinks quite trust-
worthy. For the cases of the old male baboons attacking the dogs, see
s. 79 ; and with respect to the eagle, s. 56.

76 THE DESCENT OF MAN. Part I.

months old, who, loudly calling for aid, climbed on a
block of rock and was surrounded. Now one of the
largest males, a true hero, came down again from the
mountain, slowly went to the young one, coaxed him,
and triumphantly led him away — the dogs being too
much astonished to make an attack. I cannot resist
giving another scene which was witnessed by this same
naturalist ; an eagle seized a young Cercopithecus,
which, by clinging to a branch, was not at once carried
off; it cried loudly for assistance, upon which the other
membeis of the troop with much uproar rushed to the
rescue, surrounded the eagle, and pulled out so many
feathers, that he no longer thought of his prey, but only
how to escape. This eagle, as Brehm remarks, assuredly
would never again attack a monkey in a troop.

It is certain that associated animals have a feeling of
love for each other which is not felt by adult and non-
social animals. How far in most cases they actually
sympathise with each other's pains and pleasures is
more doubtful, especially with respect to the latter.
Mr. Buxton, however, who had excellent means of
observation,9 states that his macaws, which lived free in
Norfolk, took "an extravagant interest" in a pair
with a nest, and whenever the female left it, she was
surrounded by a troop " screaming horrible accla-
" mations in her honour." It is often difficult to judge
whether animals have any feeling for each other's
sufferings. Who can say what cows fee], when they
surround and stare intently on a dying or dead
companion? That animals sometimes are far from
feeling any sympathy is too certain ; for they will expel
a wounded animal from the herd, or gore or worry
it to death. This is almost the blackest fact in natural

9 « Annals and Mag. of Nat. Hist.' Novcml er, 1868, p. 382.

Chap. III. MORAL SENSE. 77

history, unless indeed the explanation which has been
suggested is true, that their instinct or reason leads them
to expel an injured companion, lest beasts of prey,
including man, should be tempted to follow the troop.
In this case their conduct is not much worse than that
of the North American Indians who leave their feeble
comrades to perish on the plains, or the Feegeans, who,
when their parents get old or fall ill, bury them alive.10

Many animals, however, certainly sympathise with
each other's distress or clanger. This is the case even
with birds ; Capt. Stansbury n found on a salt lake in
Utah an old and completely blind pelican, which was
very fat, and must have been long and well fed by his
companions. Mr. Blyth, as he informs me, saw Indian
crows feeding two or three of their companions which
were blind ; and I have heard of an analogous case
with the domestic cock. We may, if we choose, call
these actions instinctive ; but such cases are much too
rare for the development of any special instinct.12 I
have myself seen a dog, who never passed a great
friend of his, a cat which lay sick in a basket, with-
out giving her a few licks with his tongue, the surest
sign of kind feeling in a dog.

It must be called sympathy that leads a courageous
dog to fly at any one who strikes his master, as he
certainly will. I saw a person pretending to beat a
lady who had a very timid little dog on her lap, and
the trial had never before been made. The little crea-

10 Sir J. Lubbock, ' Prehistoric Times,' 2nd edit. p. 416.

11 As quoted by Mr. L. H. Morgan, * The American Beaver,' 18GS,
p. 272. Capt. Stansbury also gives an interesting account of the man-
ner in which a very young pelican, carried away by a strong stream,
was guided and encouraged in its attempts to reach the shore by half a
dozen old birds.

12 As Mr. Bain states, " effective aid to a sufferer springs from syra-
" pathy proper:" ' Mental and Moral Science,' 1868, p. 245.

78 THE DESCENT OF MAN. Part I.

ture instantly jumped away, but after the pretended
beating was over, it was really pathetic to see how per-
se veringly he tried to lick his mistress's face and com-
fort her. Brehm 13 states that when a baboon in con-
finement was pursued to be punished, the others tried
to protect him. It must have been sympathy in the
cases above given which led the baboons and Cercopi-
theci to defend their young comrades from the dogs
and the eagle. I will give only one other instance of
sympathetic and heroic conduct in a little American
monkey. Several years ago a keeper at the Zoological
Gardens, showed me some deep and scarcely healed
wounds on the nape of his neck, inflicted on him whilst
kneeling on the floor by a fierce baboon. The little
American monkey, who was a warm friend of this
keeper, lived in the same large compartment, and was
dreadfully afraid of the great baboon. Nevertheless, as
soon as he saw his friend the keeper in peril, he rushed
to the rescue, and by screams and bites so distracted the
baboon that the man was able to escape, after running
great risk, as the surgeon who attended him thought,
of his life.

Besides love and sympathy, animals exhibit other
qualities which in us would be called moral ; and I agree
with Agassiz14 that dogs possess something very like a
conscience. They certainly possess some power of self-
command, and this does not appear to be wholly the
result of fear. As Braubach 15 remarks, a dog will
refrain from stealing food in the absence of his master.
Dogs have long been accepted as the very type of
fidelity and obedience. All animals living in a body
which defend each other or attack their enemies

13 ' Thierleben,' B. i. s. 85.

14 ' De l'Espece et de la Class.' 1869, p. 97.

15 ' Der Darwin'scken Art-Lehre,' 1869, s. 54.

Chap. III. MORAL SENSE. 79

in concert, must be in some degree faithful to each
other; and those that follow a leader must be in
some degree obedient. When the baboons in Abys-
sinia16 plunder a garden, they silently follow their
leader; and if an imprudent young animal makes a
noise, he receives a slap from the others to teach him
silence and obedience ; but as soon as they are sure
that there is no danger, all show their joy by much
clamour.

With respect to the impulse which leads certain
animals to associate together, and to aid each other in
many ways, we may infer that in most cases they are
impelled by the same sense of satisfaction or pleasure
which they experience in performing other instinctive
actions; or by the same sense of dissatisfaction, as in
other cases of prevented instinctive actions. We see
this in innumerable instances, and it is illustrated in
a striking manner by the acquired instincts of our
domesticated animals ; thus a young shepherd-dog
delights in driving and running round a flock of sheep,
but not in worrying them ; a young foxhound delights
in hunting a fox, whilst some other kinds of dogs
as I have witnessed, utterly disregard foxes. What a
strong feeling of inward satisfaction must impel a bird,
so full of activity, to brood day after day over her eggs.
Migratory birds are miserable if prevented from migrat-
ing, and perhaps they enjoy starting on their long
flight. Some few instincts are determined solely by
painful feelings, as by fear, which leads to self-preser-
vation, or is specially directed against certain enemies.
No one, I presume, can analyse the sensations of
pleasure or pain. In many cases, however, it is pro-
bable that instincts are persistently followed from the

18 Brelim, ■ Thierleben,' B. i. s. 76.

80 THE DESCENT OF MAN. Part I.

mere force of inheritance, without the stimulus of either
pleasure or pain. A young pointer, when it first scents
game, apparently cannot help pointing. A squirrel in
a cage who pats the nuts which it cannot eat, as if to
bury them in the ground, can hardly be thought to act
thus either from pleasure or pain. Hence the common
assumption that men must be impelled to every action
by experiencing some pleasure or pain may be erro-
neous. Although a habit may be blindly and implicitly
followed, independently of any pleasure or pain felt at
the moment, yet if it be forcibly and abruptly checked,
a vague sense of dissatisfaction is generally expe-
rienced ; and this is especially true in regard to persons
of feeble intellect.

It has often been assumed that animals were in the
first place rendered social, and that they feel as a con-
sequence uncomfortable when separated from each other,
and comfortable whilst together ; but it is a more pro-
bable view that these sensations were first developed, in
order that those animals which would profit by living
in society, should be induced to live together. In the
same manner as the sense of hunger and the pleasure of
eating were, no doubt, first acquired in order to induce
animals to eat. The feeling of pleasure from society
is probably an extension of the parental or filial affec-
tions ; and this extension may be in chief part attributed
to natural selection, but perhaps in part to mere habit.
For with those animals which were benefited by living
in close association, the individuals which took the
greatest pleasure in society would best escape various
dangers ; whilst those that cared least for their com-
rades and lived solitary would perish in greater numbers.
With respect to the origin of the parental and filial
affections, which apparently lie at the basis of the
social affections, it is hopeless to speculate ; but we

Chap. III. MORAL SENSE. 81

may infer that they have been to a large extent gained
through natural selection. So it has almost certainly
been with the unusual and opposite feeling of hatred
between the nearest relations, as with the worker-bees
which kill their brother-drones, and with the queen-bees
which kill their daughter-queens ; the desire to destroy,
instead of loving, their nearest relations having been
here of service to the community.

The all-important emotion of sympathy is distinct
from that of love. A mother may passionately love her
sleeping and passive infant, but she can then hardly be
said to feel sympathy for it. The love of a man for
his dog is distinct from sympathy, and so is that of a
dog for his master. Adam Smith formerly argued, as
has Mr. Bain recently, that the basis of sympathy lies
in our strong retentiveness of former states of pain or
pleasure. Hence, " the sight of another person enduring
" hunger, cold, fatigue, revives in us some recollection
" of these states, which are painful even in idea." We
are thus impelled to relieve the sufferings of another,
in order that our own painful feelings may be at the
same time relieved. In like manner we are led to
participate in the pleasures of others.17 But I cannot
see how this view explains the fact that sympathy
is excited in an immeasurably stronger degree by a
beloved than by an indifferent person. The mere

17 See the first and striking chapter in Adam Smith's ' Theory of
Moral Sentiments.' Also Mr. Bain's ' Mental .and Moral Science,'
18G8, p. 244, and 275-282. Mr. Bain states, that "sympathy is,
' indirectly, a source of pleasure to the sympathiser ; " and he accounts
for this through reciprocity. He remarks that " the person benefited,
" or others in his stead, may make up, by sympathy and good offices
" returned, for all the sacrifice." But if, as appears to be the case,
sympathy is strictly an instinct, its exercise would give direct pleasure,
in the same manner as the exercise, as before remarked, of almost every
other ins'.inct.

vol. i. a

82 THE DESCENT OF MAN. Part I.

sight of suffering, independently of love, would suffice
to call up in us vivid recollections and associations.
Sympathy may at first have originated in the manner
above suggested; but it seems now to have become
an instinct, which is especially directed towards be-
loved objects, in the same manner as fear with ani-
mals is especially directed against certain enemies. As
sympathy is thus directed, the mutual love of the
members of the same community will extend its limits.
No doubt a tiger or lion feels sympathy for the suffer-
ings of its own young, but not for any other animal.
With strictly social animals the feeling will be more
or less extended to all the associated members, as we
know to be the case. With mankind selfishness, expe-
rience, and imitation probably add, as Mr. Bain has
shewn, to the power of sympathy ; for we are led
by the hope of receiving good in return to perform
acts of sympathetic kindness to others ; and there can
be no doubt that the feeling of sympathy is much
strengthened by habit. In however complex a manner
this feeling may have originated, as it is one of high
importance to all those animals which aid and defend
each other, it will have been increased, through natural
selection ; for those communities, which included the
greatest number of the most sympathetic members,
would flourish best and rear the greatest number of
offspring.

In many cases it is impossible to decide whether
certain social instincts have been acquired through
natural selection, or are the indirect result of other
instincts and faculties, such as sympathy, reason, expe-
rience, and a tendency to imitation ; or again, whether
they are simply the result of long-continued habit.
So remarkable an instinct as the placing sentinels to
warn the community of danger, can hardly have been

Chap. III. MORAL SENSE. 83

the indirect result of any other faculty ; it must there-
fore have been directly acquired. On the other hand,
the habit followed by the males of some social animals,
of defending the community and of attacking their
enemies or their prey in concert, may perhaps have
originated from mutual sympathy; but courage, and
in most cases strength, must have been previously
acquired, probably through natural selection.

Of the various instincts and habits, some are much
stronger than others, that is, some either give more
pleasure in their performance and more distress in their
prevention than others ; or, which is probably quite as
important, they are more persistently followed through
inheritance without exciting any special feeling of plea-
sure or pain. We are ourselves conscious that some
habits are much more difficult to cure or change than
others. Hence a struggle mav often be observed in
animals between different instincts, or between an
instinct and some habitual disposition ; as when a dog
rushes after a hare, is rebuked, pauses, hesitates, pursues
again or returns ashamed to his master ; or as between
the love of a female dog for her yoimg puppies and for
her master, for she may be seen to slink away to them,
as if half ashamed of not accompanying her master.
But the most curious instance known to me of one
instinct conquering another, is the migratory instinct
conquering the maternal instinct. The former is won-
derfully strong ; a confined bird will at the proper
season beat her breast against the wires of her cage, until
it is bare and bloody. It causes young salmon to leap
out of the fresh water, where they could still continue to
live, and thus unintentionally to commit suicide. Every
one knows how strong the maternal instinct is, leading
even timid birds to face great danger, though with
hesitation and in opposition to the instinct of self-

G 2

84 THE DESCENT OF MAN. Part I.

preservation. Nevertheless trie migratory instinct is so
powerful that late in the autumn swallows and house-
martins frequently desert their tender young, leaving
them to perish miserably in their nests.18

We can perceive that an instinctive impulse, if it be in
any way more beneficial to a species than some other or
opposed instinct, would be rendered the more potent of
the two through natural selection ; for the individuals
which had it most strongly developed would survive in
larger numbers. Whether this is the case with the
migratory in comparison with the maternal instinct,
may well be doubted. The great persistence or steady
action of the former at certain seasons of the year
during the whole day, may give it for a time para-
mount force.

Man a social animal. — Most persons admit that man
is a social being. We see this in his dislike of solitude,
and in his wish for society beyond that of his own
family. Solitary confinement is one of the severest
punishments which can be inflicted. Some authors sup-
pose that man primevally lived in single families ; but
at the present day, though single families, or only two
or three together, roam the solitudes of some savage
lands, they are always, as far as I can discover, friendly
with other families inhabiting the same district. Such
families occasionally meet in council, and they unite

18 This fact, the Eev. L. Jenyns states (see his edition of ' White's
Nat. Hist, of Selborne,' 1853, p. 204) was first recorded by the illus-
trious Jenner, in ' Phil. Transact.' 1824, and has since been confirmed
by several observers, especially by Mr. Blackwall. This latter careful
observer examined, late in the autumn, during two years, thirty-six
nests ; he found that twelve contained young dead birds, five contained
eggs on the point of being hatched, and three eggs not nearly hatched.
Many birds not yet old enough for a prolonged flight are likewise
deserted and left behind. See Blackwall, 'Researches in Zoology,'
1834, pp. 108, 118. For some additional evidence, although this is nut
wanted, see Leroy, 4 Lettres Phil.' 1802, p. 217.

Chap. III. MORAL SENSE. 85

for tlieir common defence. It is no argument against
savage man being a social animal, that the tribes in-
habiting adjacent districts are almost always at war
with each other ; for the social instincts never extend
to all the individuals of the same species. Judging
from the analogy of the greater number of the Quad-
rumana, it is probable that the early ape-like pro-
genitors of man were likewise social ; but this is not of
much importance for us. Although man, as he now
exists, has few special instincts, having lost any which
his early progenitors may have possessed, this is no
reason why he should not have retained from an ex-
tremely remote period some degree of instinctive love
and sympathy for his fellows. We are indeed all con-
scious that we do possess such sympathetic feelings ; 19
but our consciousness does not tell us whether they are
instinctive, having originated long ago in the same
manner as with the lower animals, or whether they have
been acquired by each of us during our early years.
As man is a social animal, it is also probable that he
would inherit a tendency to be faithful to his comrades,
for this quality is common to most social animals. He
would in like manner possess some capacity for self-
command, and perhaps of obedience to the leader of
the community. He would from an inherited tendency
still be willing to defend, in concert with others, his
fellow-men, and would be ready to aid them in any
way which did not too greatly interfere with his own
welfare or his own strong desires.

The social animals which stand at the bottom of the

19 Hume remarks ('An Enquiry Concerning the Principles of Morals,'
edit, of 1751, p. 132), " there seems a necessity for confessing that the
" happiness and misery of others are not spectacles altogether iu-
" different to us, but that the view of the former . . . communicates a
" secret joy; the appearance of the latter . . . throws a melancholy
" damp over the imagination." //-" ' ^

ILtB

8G THE DESCENT OF MAN. Part I.

scale are guided almost exclusively, and those which
stand higher in the scale are largely guided, in the aid
which they give to the members of the same community,
by special instincts ; but they are likewise in part im-
pelled by mutual love and sympathy, assisted appa-
rently by some amount of reason. Although man, as just
remarked, has no special instincts to tell him how to aid
his fellow-men, he still has the impulse, and with his
improved intellectual faculties would naturally be much
guided in this respect by reason and experience. In-
stinctive sympathy would, also, cause him to value highly
the approbation of his fellow-men ; for, as Mr. Bain has
clearly shewn,20 the love of praise and the strong feeling
of glory, and the still stronger horror of scorn and in-
famy, " are due to the workings of sympathy." Conse-
quently man would be greatly influenced by the wishes,
approbation, and blame of his fellow-men, as expressed
by their gestures and language. Thus the social in-
stincts, which must have been acquired by man in a
very rude state, and probably even by his early ape-like
progenitors, still give the impulse to many of his best
actions ; but his actions are largely determined by the
expressed wishes and judgment of his fellow-men, and
unfortunately still oftener by his own strong, selfish
desires. But as the feelings of love and sympathy and
the power of self-command become strengthened by
habit, and as the power of reasoning becomes clearer so
that man can appreciate the justice of the judgments of
his fellow-men, he will feel himself impelled, independ-
ently of any pleasure or pain felt at the moment, to
certain lines of conduct. He may then say, I am the
supreme judge of my own conduct, and in the words of
Kant, I will not in my own person violate the dignity
of humanity.

20 ' Mental and Moral Science/ 1868, p. 254.

Chap. III. MORAL SENSE. 87

The more enduring Social Instincts conquer the less
Persistent Instincts. — We have, however, not as yet con-
sidered the main point, on which the whole question of
the moral sense hinges. Why should a man feel that
he ought to obey one instinctive desire rather than
another ? Why does he bitterly regret if he has yielded
to the strong sense of self-preservation, and has not
risked his life to save that of a fellow-creature ; or why
does he regret having stolen food from severe hunger?

It is evident in the first place, that with mankind the
instinctive impulses have different degrees of strength ;
a young and timid mother urged by the maternal in-
stinct will, without a moment's hesitation, run the
greatest danger for her infant, but not for a mere fel-
low-creature. Many a man, or even boy, who never
before risked his life for another, but in whom courage
and sympathy were well developed, has, disregarding the
instinct of self-preservation, instantaneously plunged
into a torrent to save a drowning fellow-creature. In
this case man is impelled by the same instinctive mo-
tive, which caused the heroic little American monkey,
formerly described, to attack the great and dreaded
baboon, to save his keeper. Such actions as the above
appear to be the simple result of the greater strength of
the social or maternal instincts than of any other instinct
or motive ; for they are performed too instantaneously
for reflection, or for the sensation of pleasure or pain ;
though if prevented distress would be caused.

I am aware that some persons maintain that actions
performed impulsively, as in the above cases, do not
come under the dominion of the moral sense, and
cannot be called moral. They confine this term to
actions done deliberately, after a victory over opposing
desires, or to actions prompted by some lofty motive.
But it appears scarcely possible to draw any clear line

88 THE DESCENT OF MAN. Part I.

of distinction of this kind ; though the distinction may
be real. As far as exalted motives are concerned, many
instances have been recorded of barbarians, destitute of
any feeling of general benevolence towards mankind,
and not guided by any religious motive, who have deli-
berately as prisoners sacrificed their lives,21 rather than
betray their comrades ; and surely their conduct ought
to be considered as moral. As far as deliberation and
the victory over opposing motives are concerned, ani-
mals may be seen doubting between opposed instincts,
as in rescuing their offspring or comrades from dan-
ger ; yet their actions, though done for the good of
others, are not called moral. Moreover, an action
repeatedly performed by us, will at last be done with-
out deliberation or hesitation, and can then hardly be
distinguished from an instinct ; yet surely no one will
pretend that an action thus done ceases to be moral.
On the contrary, we all feel that an act cannot be
considered as perfect, or as performed in the most
noble manner, unless it be done impulsively, without
deliberation or effort, in the same manner as by a man
in whom the requisite qualities are innate. He
who is forced to overcome his fear or want of sym-
pathy before he acts, deserves, however, in one way
higher credit than the man whose innate disposition
leads him to a good act without effort. As we cannot
distinguish between motives, we rank all actions of a
certain class as moral, when they are performed by
a moral being. A moral being is one who is capable of
comparing his past and future actions or motives, and
of approving or disapproving of them. We have no
reason to suppose that any of the lower animals have

21 I have given one such case, namely of three Patagonian Indians
who preferred being shot, one after the other, to betraying the plans of
their companions in war (, Journal of Researches,' 1845,. p. 103).

Chap. III. MORAL SENSE. 89

this capacity ; therefore when a monkey faces danger
to rescue its comrade, or takes charge of an orphan-
monkey, we do not call its conduct moral. But in the
case of man, who alone can with certainty be ranked as
a moral being, actions of a certain class are called moral,
whether performed deliberately after a struggle with
opposing motives, or from the effects of slowly-gained
habit, or impulsively through instinct.

But to return to our more immediate subject ; al-
though some instincts are more powerful than others,
thus leading to corresponding actions, yet it cannot
be maintained that the social instincts are ordinarily
stronger in man, or have become stronger through
long-continued habit, than the instincts, for instance,
of self-preservation, hunger, lust, vengeance, &c. Why
then does man regret, even though he may endeavour
to banish any such regret, that he has followed the
one natural impulse, rather than the other; and why
does he further feel that he ought to regret his conduct ?
Man in this respect differs profoundly from the lower
animals. Nevertheless we can, I think, see with some
degree of clearness the reason of this difference.

Man, from the activity of his mental faculties, cannot
avoid reflection : past impressions and images are in-
cessantly passing through his mind with distinctness.
Now with those animals which live permanently in a
body, the social instincts are ever present and per-
sistent. Such animals are always ready to utter the
danger-signal, to defend the community, and to give
aid to their fellows in accordance with their habits;
they feel at all times, without the stimulus of any
special passion or desire, some degree of love and
sympathy for them ; they are unhappy if long separated
from them, and always happy to be in their company.
So it is with ourselves. A man who possessed no trace

90 THE DESCENT OF MAN. Part I.

of such feelings would be an unnatural monster. On
the other hand, the desire to satisfy hunger, or any
passion, such as vengeance, is in its nature temporary,
and can for a time be fully satisfied. Nor is it easy,
perhaps hardly possible, to call up with complete vivid-
ness the feeling, for instance, of himger ; nor indeed, as
has often been remarked, of any suffering. The in-
stinct of self-preservation is not felt except in the pre-
sence of danger ; and many a coward has thought him-
self brave until he has met his enemy face to face.
The wish for another man's property is perhaps as
persistent a desire as any that can be named ; but even
in this case the satisfaction of actual possession is gene-
rally a weaker feeling than the desire : many a thief, if
not an habitual one, after success has wondered why he
stole some article.

Thus, as man cannot prevent old impressions con-
tinually repassing through his mind, he will be com-
pelled to compare the weaker impressions of, for in-
stance, past hunger, or of vengeance satisfied or danger
avoided at the cost of other men, with the instinct of
sympathy and good-will to his fellows, which is still pre-
sent and ever in some degree active in his mind. He
will then feel in his imagination that a stronger instinct
has yielded to one which now seems comparatively
weak ; and then that sense of dissatisfaction will in-
evitably be felt with which man is endowed, like every
other animal, in order that his instincts may be obeyed.
The case before given, of the swallow, affords an illus-
tration, though of a reversed nature, of a temporary
though for the time strongly persistent instinct con-
quering another instinct which is usually dominant over
all others. At the proper season these birds seem all
day long to be impressed with the desire to migrate ;
their habits change ; they become restless, are noisy,

Chap. III. MORAL SENSE. 91

and congregate in flocks. AYhilst the mother-bird is
feeding or brooding over her nestlings, the maternal
instinct is probably stronger than the migratory ; but
the instinct which is more persistent gains the victory,
and at last, at a moment when her young ones are not
in sight, she takes flight and deserts them. When
arrived at the end of her long journey, and the migra-
tory instinct ceases to act, what an agony of remorse
each bird wrould feel, if, from being endowed with great
mental activity, she could not prevent the image con-
tinually passing before her mind of her young ones
perishing in the bleak north from cold and hunger.

At the moment of action, mam will no doubt be apt
to follow the stronger impulse ; and though this may
occasionally prompt him to the noblest deeds, it will
far more commonly lead him to gratify his own desires
at the expense of other men. But after their grati-
fication, when past and weaker impressions are con-
trasted with the ever-enduring social instincts, retri-
bution will surely come. Man will then feel dissatis-
fied with himself, and will resolve with more or less
force to act differently for the future. This is con-
science ; for conscience looks backwards and judges past
actions, inducing that kind of dissatisfaction, which if
weak we call regret, and if severe remorse.

These sensations are, no doubt, different from those
experienced when other instincts or desires are left
unsatisfied; but every unsatisfied instinct has its own
proper prompting sensation, as we recognise with hunger,
thirst, &c. Man thus prompted, "will through long
habit acquire such perfect self-command, that his desires
and passions will at last instantly yield to his social
sympathies, and there will no longer be a struggle
between them. The still hungry, or the still revengeful
man will not think of stealing food, or of wreaking his

92 THE DESCENT OF MAN. Part I.

vengeance. It is possible, or, as we shall hereafter
see, even probable, that the habit of self-command
may, like other habits, be inherited. Thus at last
man comes to feel, through acquired and perhaps in-
herited habit, that it is best for him to obey his more
persistent instincts. The imperious word ought seems
merely to imply the consciousness of the existence of a
persistent instinct, either innate or partly acquired,
serving him as a guide, though liable to be disobeyed.
We hardly use the word ought in a metaphorical sense,
when we say hounds ought to hunt, pointers to point,
and retrievers to retrieve their game. If they fail thus
to act, they fail in their duty and act wrongly.

If any desire or instinct, leading to an action opposed
to the good of others, still appears to a man, when re-
called to mind, as strong as, or stronger than, his social
instinct, he will feel no keen regret at having followed
it ; but he will be conscious that if his conduct were
known to his fellows, it would meet with their disap-
probation ; and few are so destitute of sympathy as not
to feel discomfort when this is realised. If he has no
such sympathy, and if his desires leading to bad actions
are at the time strong, and when recalled are not over-
mastered by the persistent social instincts, then he is
essentially a bad man ; ffl and the sole restraining motive
left is the fear of punishment, and the conviction that
in the long run it would be best for his own selfish
interests to regard the good of others rather than his
own.

It is obvious that every one may with an easy con-
science gratify his own desires, if they do not interfere

27 Dr. Prosper Despine, in his 'Psychologic Naturelle,' ]868 (torn,
i. p. 243 ; torn. ii. p. 169) gives many curious cases of the worst criminals,
who apparently have been entirely destitute of conscience.

Chap. III. MORAL SENSE. 93

with his social instincts, that is with the good of others ;
but in order to be quite free from self-reproach, or at
least of anxiety, it is almost necessary for him to avoid
the disapprobation, whether reasonable or not, of his
fellow men. Nor must he break through the fixed habits
of his life, especially if these are supported by reason ;
for if he does, he will assuredly feel dissatisfaction.
He must likewise avoid the reprobation of the one
God or gods, in whom according to his knowledge or
superstition he may believe ; but in this case the addi-
tional fear of divine punishment often supervenes.

The strictly Social Virtues at first alone regarded. —
The above view of the first origin and nature of the moral
sense, which tells us what we ought to do, and of
the conscience which reproves us if we disobey it,
accords well with what we see of the early and un-
developed condition of this faculty in mankind. The
virtues which must be practised, at least generally, by
rude men, so that they may associate in a body, are
those which are still recognised as the most important.
But they are practised almost exclusively in relation to
the men of the same tribe ; and their opposites are not
regarded as crimes in relation to the men of other tribes.
No tribe could hold together if murder, robbery, trea-
chery, &c, were common ; consequently such crimes
within the limits of the same tribe "are branded
" with everlasting infamy ; " ffl but excite no such senti-
ment beyond these limits. A North-American Indian
is well pleased with himself, and is honoured by others,
when he scalps a man of another tribe ; and a Dyak

23 Sec an able article in the ' North British Review,' 1867, p. 395.
See also Mr. W. Bagehot's articles on the Importance of Obedience
and Coherence to Primitive Man, in the ' Fortnightly Review,' 1867,
p. 529, and 1868, p. 457, &c.

94 THE DESCENT OF MAN. Pakt I.

cuts off the head of an unoffending person and dries it
as a trophy. The murder of infants has prevailed on
the largest scale throughout the world,24 and has met
with no reproach ; but infanticide, especially of females,
has been thought to be good for the tribe, or at least
not injurious. Suicide during former times was not
generally considered as a crime,25 but rather from the
courage displayed as an honourable act ; and it is still
largely practised by some semi-civilised nations without
reproach, for the loss to a nation of a single individual
is not felt : whatever the explanation may be, suicide
is rare amongst barbarians ; the negroes on the west
coast of Africa offering, however, as I hear from Mr.
W. Keade, in this respect an exception. It has been
recorded that an Indian Thug conscientiously regretted
that he had not strangled and robbed as many travellers
as did his father before him. In a rude state of civilisa-
tion the robbery of strangers is, indeed, generally con-
sidered as honourable.

The great sin of Slavery has been almost universal, and
slaves have often been treated in an infamous manner.
As barbarians do not regard the opinion of their women,
wives are commonly treated like slaves. Most savages
are utterly indifferent to the sufferings of strangers, or
even delight in witnessing them. It is well known that
the women and children of the North- American Indians
aided in torturing their enemies. Some savages take a
horrid pleasure in cruelty to animals,26 and humanity
with them is an unknown virtue. Nevertheless, feelings
of sympathy and kindness are common, especially

24 The fullest account which I have met with is by Dr. Gerland, in
his ' Ueber das Aussterben der Naturvolker,' 1868 ; but I shall have to
recur to the subject of infanticide in a future chapter.

25 See the very interesting discussion on Suicide in Lecky's ' History
of European Morals,' vol. i. 1869, p. 223.

26 See, for instance, Mr. Hamilton's account of the Kaffirs, ' Anthro-
pological Review/ 1870, p. xv.

Chap. III. MORAL SENSE. 95

during sickness, between the members of the same tribe,
and are sometimes extended beyond the limits of the
tribe. Mungo Park's touching account of the kindness
of the negro women of the interior to him is well known.
Many instances could be given of the noble fidelity of
savages towards each other, but not to strangers;
common experience justifies the maxim of the Spaniard,
"Never, never trust an Indian." There cannot be
fidelity without truth ; and this fundamental virtue is
not rare between the members of the same tribe : thus
Mungo Park heard the negro women teaching their
young children to love the truth. This, again, is one of
the virtues which becomes so deeply rooted in the mind
that it is sometimes practised by savages even at a high
cost, towards strangers ; but to lie to your enemy has
rarely been thought a sin, as the history of modern
diplomacy too plainly shews. As soon as a tribe has
a recognised leader, disobedience becomes a crime, and
even abject submission is looked at as a sacred virtue.

As during rude times no man can be useful or faithful
to his tribe without courage, this quality has universally
been placed in the highest rank; and although, in
civilised countries, a good, yet timid, man may be far
more useful to the community than a brave one, we
cannot help instinctively honouring the latter above
a coward, however benevolent. Prudence, on the other
hand, which does not concern the welfare of others,
though a very useful virtue, has never been highly
esteemed. As no man can practise the virtues necessary
for the welfare of his tribe without ^self-sacrifice, self-
command, and the power of endurance, these qualities
have been at all times highly and most justly valued.
The American savage voluntarily submits without
a groan to the most horrid tortures to prove and
strengthen his fortitude and courage ; and we cannot

96 THE DESCENT OF MAN. Part I.

help admiring him, or even an Indian Fakir, who, from
a foolish religious motive, swings suspended by a hook
buried in his flesh.

The other self-regarding virtues, which do not ob-
viously, though they may really, affect the welfare of
the tribe, have never been esteemed by savages, though
now highly appreciated by civilised nations. The
greatest intemperance with savages is no reproach.
Their utter licentiousness, not to mention unnatural
crimes, is something astounding.27 As soon, however, as
marriage, whether polygamous or monogamous, becomes
common, jealousy will lead to the inculcation of female
virtue ; and this being honoured will tend to spread to the
unmarried females. How slowly it spreads to the male
sex we see at the present day. Chastity eminently
requires self-command ; therefore it has been honoured
from a very early period in the moral history of civilised
man. As a consequence of this, the senseless practice
of celibacy has been ranked from a remote period as
a virtue.28 The hatred of indecency, which appears to
us so natural as to be thought innate, and which is so
valuable an aid to chastity, is a modern virtue, apper-
taining exclusively, as Sir G. Staunton remarks,29 to
civilised life. This is shewn by the ancient religious
rites of various nations, by the drawings on the walls of
Pompeii, and by the practices of many savages.

We have now seen that actions are regarded by
savages, and were probably so regarded by primeval
man, as good or bad, solely as they affect in an obvious
manner the welfare of the tribe, — not that of the
species, nor that of man as an individual member of the

27 Mr. M'Lennan has given ('Primitive Marriage,' 1865, p. 176) a
good collection of facts on this head.

Lecky, ' History of European Morals,' vol. i. 1869, p. 109.
29 'Embassy to China,' vol. ii. p. 348.

28

Chap. III. MOKAL SENSE. 97

tribe. This conclusion agrees well with the belief that
the so-called moral sense is aboriginally derived from
the social instincts, for both relate at first exclusively to
the community. The chief causes of the low morality
of savages, as judged by our standard, are, firstly, the
confinement of sympathy to the same tribe. Secondly,
insufficient powers of reasoning, so that the bearing of
many virtues, especially of the self-regarding virtues, on
the general welfare of the tribe is not recognised.
Savages, for instance, fail to trace the multiplied evils
consequent on a want of temperance, chastity, &c.
And, thirdly, weak power of self-command; for this
power has not been strengthened through long-con-
tinued, perhaps inherited, habit, instruction and religion.
I have entered into the above details on the immor-
ality of savages,30 because some authors have recently
taken a high view of their moral nature, or have attri-
buted most of their crimes to mistaken benevolence.31
These authors appear to rest their conclusion on savages
possessing, as they undoubtedly do possess, and often
in a high degree, those virtues which are serviceable,
or even necessary, for the existence of a tribal com-
munity.

Concluding Remarks. — Philosophers of the derivative32
school of morals formerly assumed that the foundation
of morality lay in a form of Selfishness ; but more
recently in the " Greatest Happiness principle." Ac-
cording to the view given above, the moral sense is

30 See on this subject copious evidence in Chap. vii. of Sir J.
Lubbock, ' Origin of Civilisation,' 1870.

31 For instance Lecky, ' Hist. European Morals,' vol. i. p. 124.

32 This term is used in an able article in the ' Westminister Eeview,'
Oct. 1S69, p. 498. For the Greatest Happiness principle, see J. S.
Mill, ' Utilitarianism,' p. 17.

VOL. I. H

98 THE DESCENT OF MAN. Part I-

fundamentally identical with the social instincts; and
in the case of the lower animals it would be absurd to
speak of these instincts as having been developed from
selfishness, or for the happiness of the community.
They have, however, certainly been developed for the
general good of the community. The term, general
good, may be defined as the means by which the great-
est possible number of individuals can be reared in
full vigour and. health, with all their faculties perfect,
under the conditions to which they are exposed. As
the social instincts both of man and the lower animals
have no doubt been developed by the same steps, it
would be advisable, if found practicable, to use the
same definition in both cases, and to take as the test
of morality, the general good or welfare of the com-
munity, rather than the general happiness ; but this
definition would perhaps require some limitation on
account of political ethics.

When a man risks his life to save that of a fellow-
creature, it seems more appropriate to say that he acts
for the general good or welfare, rather than for the
general happiness of mankind. No doubt the welfare
and the happiness of the individual usually coincide ;
and a contented, happy tribe will flourish better than
one that is discontented and unhappy. We have seen
that at an early period in the history of man, the ex-
pressed wishes of the community will have naturally
influenced to a large extent the conduct of each mem-
ber ; and as all wish for happiness, the " greatest happi-
ness principle" will have become a most important
secondary guide and object ; the social instincts, includ-
ing sympathy, always serving as the primary impulse
and guide. Thus the reproach of laying the foundation
of the most noble part of our nature in the base prin-
ciple of selfishness is removed ; unless indeed the satis-

Chap. III. MORAL SENSE. 99

faction which every animal feels when it follows its
proper instincts, and the dissatisfaction felt when pre-
vented, be called selfish.

The expression of the wishes and judgment of the
members of the same community, at first by oral and
afterwards by written language, serves, as just re-
marked, as a most important secondary guide of
conduct, in aid of the social instincts, but sometimes
in opposition to them. This latter fact is well exem-
plified by the Law of Honour, that is the law of the
opinion of our equals, and not of all our country-
men. The breach of this law, even when the breach
is known to be strictly accordant with true mo-
rality, has caused many a man more agony than a real
crime. We recognise the same influence in the burn-
ing sense of shame which most of us have felt even
after the interval of years, when calling to mind some
accidental breach of a trifling though fixed rule of eti-
quette. The judgment of the community will generally
be guided by some rude experience of what is best in
the long run for all the members ; but this judgment
will not rarely err from ignorance and from weak powers
of reasoning. Hence the strangest customs and super-
stitions, in complete opposition to the true welfare and
happiness of mankind, have become all-powerful through-
out the world. We see this in the horror felt by a
Hindoo who breaks his caste, in the shame of a Maho-
metan woman who exposes her face, and in innumerable
other instances. It would be difficult to distinguish
between the remorse felt by a Hindoo who has eaten
unclean food, from that felt after committing a theft;
but the former would probably be the more severe.

How so many absurd rules of conduct, as well as so
many absurd religious beliefs, have originated we do
not know ; nor how it is that they have become, in all

h 2

100 THE DESCENT OF MAN. Part I.

quarters of the world, so deeply impressed on the mind
of men ; but it is worthy of remark that a belief con-
stantly inculcated during the early years of life, whilst
the brain is impressible, appears to acquire almost the
nature of an instinct ; and the very essence of an in-
stinct is that it is followed independently of reason.
Neither can Ave say why certain admirable virtues, such
as the love of truth, are much more highly appre-
ciated by some savage tribes than by others ; ^ nor,
again, why similar differences prevail even amongst
civilised nations. Knowing how firmly fixed many
strange customs and superstitions have become, we
need feel no surprise that the self-regarding virtues
should now appear to us so natural, supported as they
are by reason, as to be thought innate, although they
were not valued by man in his early condition.

Notwithstanding many sources of doubt, man can
generally and readily distinguish between the higher
and lower moral rules. The higher are founded on the
social instincts, and relate to the welfare of others.
They are supported by the approbation of our fellow-
men and by reason. The lower rules, though some of
them when implying self-sacrifice hardly deserve to be
called lower, relate chiefly to self, and' owe their origin
to public opinion, when matured by experience and
cultivated ; for they are not practised by rude tribes.

As man advances in civilisation, and small tribes
are united into larger communities, the simplest reason
would tell each individual that he ought to extend his
social instincts and sympathies to all the members of
the same nation, though personally unknown to him.
This point being once reached, there is only an arti-

83 Good instances are given by Mr. Wallace in ' Scientific Opinion,'
Sept. 15, 1869 ; and more fully in his ' Contributions to the Theory of
Natural Selection,' 1S70, p. 353.

Chap. III. MORAL SENSE. 101

fieial barrier to prevent his sympathies extending to the
men of all nations and races. If, indeed, such men are
separated from him by great differences in appearance
or habits, experience unfortunately shews us how long
it is before we look at them as our fellow-creatures.
Sympathy beyond the confines of man, that is humanity
to the lower animals, seems to be one of the latest
moral acquisitions. It is apparently unfelt by savages,
except towards their pets. How little the old Romans
knew of it is shewn by their abhorrent gladiatorial
exhibitions. The very idea of humanity, as far as I
could observe, was new to most of the G-auchos of the
Pampas. This virtue, one of the noblest with which
man is endowed, seems to arise incidentally from our
sympathies becoming more tender and more widely dif-
fused, until they are extended to all sentient beings.
As soon as this virtue is honoured and practised by some
few men, it spreads through instruction and example to
the young, and eventually through public opinion.

The highest stage in moral culture at which we can
arrive, is when we recognise that we ought to control
our thoughts, and " not even in inmost thought to think
" again the sins that made the past so pleasant to us."34
Whatever makes any bad action familiar to the mind,
renders its performance by so much the easier. As
Marcus Aurelius long ago said, " Such as are thy habi-
" tual thoughts, such also will be the character of thy
" mind; for the soul is dyed by the thoughts."35

Our great philosopher, Herbert Spencer, has recently
explained his views on the moral sense. He says,36 " I

34 Tennyson, ■ Idylls of the King,' p. 244. /CW ^

35 'The Thoughts of the Emperor M. Aurelius Anton h^u^Enp S hf(
translat., 2nd edit., 186'.', p. 112. Marcus Aurelius was boynQS). fl£l .

36 Letter to Mr. Mill in Bain's 'Mental and Moral S/igte^lSD^
P. 722. 'UBR<-

&\ N4» ■* -i

102 THE DESCENT OF MAN. Part I.

" believe that the experiences of utility organised and
" consolidated through all past generations of the human
" race, have been producing corresponding modifications,
" which, by continued transmission and accumulation,
" have become in us certain faculties of moral intuition —
" certain emotions responding to right and wrong con-
" duct, which have no apparent basis in the individual
" experiences of utility." There is not the least inhe-
rent improbability, as it seems to me, in virtuous ten-
dencies being more or less strongly inherited ; for, not
to mention the various dispositions and habits trans-
mitted by many of our domestic animals, I have heard
of cases in which a desire to steal and a tendency to lie
appeared to run in families of the upper ranks; and
as stealing is so rare a crime in the wealthy classes,
we can hardly account by accidental coincidence for the
tendency occurring in two or three members of the
same family. If bad tendencies are transmitted, it is
probable that good ones are likewise transmitted. Ex-
cepting through the principle of the transmission of
moral tendencies, we cannot understand the differences
believed to exist in this respect between the various
races of mankind. We have, however, as yet, hardly
sufficient evidence on this head.

Even the partial transmission of virtuous tendencies
would be an immense assistance to the primary impulse
derived directly from the social instincts, and indirectly
from the approbation of our fellow-men. Admitting
for the moment that virtuous tendencies are inherited,
it appears probable, at least in such cases as chastity,
temperance, humanity to animals, &c, that they become
first impressed on the mental organisation through
habit, instruction, and example, continued during several
generations in the same family, and in a quite subor-
dinate degree, or not at all, by the individuals pos-

Chap. III. MORAL SENSE. 103

sessing such virtues, having succeeded best in the
struggle for life. My chief source of doubt with respect
to any such inheritance, is that senseless customs, super-
stitions, and tastes, such as the horror of a Hindoo for
unclean food, ought on the same principle to be trans-
mitted. Although this in itself is perhaps not less pro-
bable than that animals should acquire inherited tastes
for certain kinds of food or fear of certain foes, I have
not met with any evidence in support of the trans-
mission of superstitious customs or senseless habits.

Finally, the social instincts which no doubt were
acquired by man, as by the lower animals, for the good
of the community, will from the first have given to him
some wish to aid his fellows, and some feeling of sym-
pathy. Such impulses will have served him at a very
early period as a rude rule of right and wrong. But as
man gradually advanced in intellectual power and was
enabled to trace the more remote consequences of his
actions; as he acquired sufficient knowledge to reject
baneful customs and superstitions ; as he regarded
more and more not only the welfare but the happi-
ness of his fellow-men ; as from habit, following on
beneficial experience, instruction, and example, his
sympathies became more tender and widely diffused,
so as to extend to the men of all races, to the im-
becile, the maimed, and other useless members of
society, and finally to the lower animals, — so would the
standard of his morality rise higher and higher. And
it is admitted by moralists of the derivative school and
by some intuitionists, that the standard of morality has
risen since an early period in the history of man.37

As a struggle may sometimes be seen going on

37 A writer in the 'North British Keview' (July, 1869, p. 531), well
capable of forming a sound judgment, expresses himself strongly to this

104 THE DESCENT OF MAN. Part I.

between the various instincts of the lower animals, it is
not surprising that there should be a struggle in man
between his social instincts, with their derived virtues,
and his lower, though at the moment, stronger impulses
or desires. This, as Mr. Galton 38 has remarked, is all
the less surprising, as man has emerged from a state of
barbarism within a comparatively recent period. After
having yielded to some temptation we feel a sense of
dissatisfaction, analogous to that felt from other un-
satisfied instincts, called in this case conscience ; for we
cannot prevent past images and impressions continually
passing through our minds, and these in their weakened
state we compare with the ever-present social instincts,
or with habits gained in early youth and strengthened
during our whole lives, perhaps inherited, so that they
are at last rendered almost as strong as instincts.
Looking to future generations, there is no cause to fear
that the social instincts will grow weaker, and we may
expect that virtuous habits will grow stronger, becoming
perhaps fixed by inheritance. In this case the struggle
between our higher and lower impulses will be less
severe, and virtue will be triumphant.

Summary of the two last Chapters. — There can be no
doubt that the difference between the mind of the lowest
man and that of the highest animal is immense. An
anthropomorphous ape, if he could take a dispassionate
view of his own case, would admit that though he could
form an artful plan to plunder a garden — though he
could use stones for fighting or for breaking open nuts,

effect. Mr. Lecky (' Hist, of Morals,' vol. i. p. 143) seems to a certain
extent to coincide.

38 See his remarkable work on ' Hereditary Genius,' 1SG9, p. 349.
The Duke of Argyll ('Primeval Man,' 1869, p. 188) lias some good
remarks on the contest in man's nature between right and wrong.

Chap. III. SUMMARY. 105

yet that the thought of fashioning a stone into a tool
was quite beyond his scope. Still less, as he would
admit, could he follow out a train of metaphysical
reasouing, or solve a mathematical problem, or reflect
on God, or admire a grand natural scene. Some apes,
however, would probably declare that they could and
did admire the beauty of the coloured skin and fur of
their partners in marriage. They would admit, that
though they could make other apes understand by cries
some of their perceptions and simpler wants, the notion
of expressing definite ideas by definite sounds had
never crossed their minds. They might insist that they
were ready to aid their fellow-apes of the same troop in
many ways, to risk their lives for them, and to take
charge of their orphans ; but they would be forced to
acknowledge that disinterested love for all living crea-
tures, the most noble attribute of man, was quite be-
yond their comprehension.

Nevertheless the difference in mind between man
and the higher animals, great as it is, is certainly one
of degree and not of kind. We have seen that the
senses and intuitions, the various emotions and faculties,
such as love, memory, attention, curiosity, imitation,
reason, &c, of which man boasts, may be found in an
incipient, or even sometimes in a well-developed con-
dition, in the lower animals. They are also capable of
some inherited improvement, as we see in the domestic
dog compared with the wolf or jackal. If it be main-
tained that certain powers, such as self-consciousness,
abstraction, &c, are peculiar to man, it may well be
that these are the incidental results of other highly-
advanced intellectual faculties; and these again are
mainly the result of the continued use of a highly
developed language. At what age does the new-born
infant possess the power of abstraction, or become self-

106 THE DESCENT OF MAN. Part I.

conscious and reflect on its own existence ? We cannot
answer ; nor can we answer in regard to the ascending
organic scale. The half-art and half-instinct of lan-
guage still bears the stamp of its gradual evolution.
The ennobling belief in God is not universal with man ;
and the belief in active spiritual agencies naturally fol-
lows from his other mental powers. The moral sense
perhaps affords the best and highest distinction between
man and the lower animals ; but I need not say any-
thing on this head, as I have so lately endeavoured
to shew that the social instincts, — the prime principle
of man's moral constitution39 — with the aid of active
intellectual powers and the effects of habit, naturally
lead to the golden rule, " As ye would that men should
" do to you, do ye to them likewise ; " and this lies at
the foundation of morality.

In a future chapter I shall make some few remarks
on the probable steps and means by which the several
mental and moral faculties of man have been gradually
evolved. That this at least is possible ought not
to be denied, when we daily see their development in
every infant ; and when we may trace a perfect grada-
tion from the mind of an utter idiot, lower than that of
the lowest animal, to the mind of a Newton.

39 < The Thoughts of Marcus Aurelius,' &c, p. 139.

Chap. IV. MANNER OF DEVELOPMENT. 107

CHAPTEE IV.

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 — Re-
version— 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 — AbseDce of
a tail — Defenceless condition of man.

We have seen in the first chapter that the home-logical
structure of man, his embryological development and
the rudiments which he still retains, all declare in the
plainest manner that he is descended from some lower
form. The possession of exalted mental powers is no
insuperable objection to this conclusion. In order that
an ape-like creature should have been transformed into
man, it is necessary that this early form, as well as
many successive links, should all have varied in mind
and body. It is impossible to obtain direct evidence on
this head; but if it can.be shewn that man now varies
— that his variations are induced by the same general
causes, and obey the same general laws, as in the case
of the lower animals — there can be little doubt that
the preceding intermediate links varied in a like
maimer. The variations at each successive stage of
descent must, also, have been in some manner accumu-
lated and fixed.

108 THE DESCENT OF MAN. Part I.

The facts and conclusions to be given in this chapter
relate almost exclusively to the probable means by
which the transformation of man has been effected,
as far as his bodily structure is concerned. The fol-
lowing chapter will be devoted to the development of
his intellectual and moral faculties. But the present
discussion likewise bears on the origin of the different
races or species of mankind, whichever term may be
preferred.

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, arid
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 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 An emi-
nent 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

1 ' Investigations in Military and Anthropolog. Statistics of American
Soldiers,' by B. A. Gould. 1869, p. 256.

2 With respect to the " Cranial forms of the American aborigines,"
see Dr. Aitktn Meigs in ' Proc. Acad. Nat. Sci.' Philadelphia, May,
1866. On the Australians, see Huxley, in Lyell's ' Anticpiiity of Man,'
1863, p. 87. On the Sandwich Islanders, Prof. J. Wyinan, ' Observa-
tions on Crania,' Boston, 1868, p. 18.

Chap. IV. MANNEE OF DEVELOPMENT. 1 09

from 12,000 corpses how often each course prevails.3
The muscles are eminently variable : thus those of the
foot were found by Prof. Turner4 not to be strictly
alike in any two out of fifty bodies ; and in some the
deviations were considerable. Prof. Turner 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 the occurrence
of 295 muscular variations in thirty-six subjects, and in
another set of the same number no less than 558 varia-
tions, reckoning both sides of the body as one. In the
last set, not one body out of the thirty-six was " found
" totally wanting in departures from the standard de-
" scriptions of the muscular system given in anatomical
" text-books." A single body presented the extraordi-
nary number of twenty-five distinct abnormalities. The
same muscle sometimes varies in many ways : thus
Prof. Macalister describes 6 no less than twenty distinct
variations in the palmaris accessorius.

The famous old anatomist, Wolff,7 insists that the
internal viscera are more variable than the external
parts: Nulla particula est quw non aliter et aliier 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, &c, 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

3 ' Anatomy of the Arteries,' by E. Quain.

4 ' Transact. Eoyal Soc.' Edinburgh, vol. xxiv. p. 1.75, 189.

5 * Proc, Eoyal Soc.' 1867, p. 544 ; also 1868, p. 483, 524. There is
a previous paper, 1866, p. 229.

6 ' Proc. E. Irish Academy,' vol. x. 1868, p. 141.

7 ' Act. Acad.,' St. Petersburg, 1778, part ii. p. 217.

110 THE DESCENT OF MAN. Part I.

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, as has been illustrated by
a few examples in the last chapter. 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 under confinement 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 remark-
able for intelligence. Eengger, also, insists on the di-
versity 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

I have elsewhere 9 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, &c, are certainly transmitted. With man
we see similar facts in almost every famil}r ; and we

8 Brehm, ' Thierleben,' B. i. s. 58, 87. Piengger, ' S'augethiere von

Paraguay,' s. 57.

9 k Variation of Animals and Plants under Domestication,' vol. ii.
chap. xii.

Chap. IV. MANNEK OF DEVELOPMENT. Ill

now know through the admirable labours of Mr. G-alton 10
that genius, which implies a wonderfully complex com-
bination 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 the same
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 staud in some relation with
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 appa-
rently due to the diversified and changing nature of
their conditions. The different races of man resemble
in this respect 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,
the members of which belong to different grades of rank
and follow different occupations, presenting a greater
range of character than 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 It is nevertheless an error to speak of man,
even if we look only to the conditions to which he
has been subjected, as " far more domesticated " 12 than

10 ' Hereditary Genius : an Inquiry into its Laws and Consequences,'
1869.

11 Mr. Bates remarks (' The Naturalist on the Amazons,' 18G3, vol. ii.
p. 159 , with respect to the Indians of the same S. 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,
" aud obliquity of eyes."

12 Blumenbach, ' Treatises on Anthropolog.' Eng. translat., 1865,
p. 205.

112 THE DESCENT OF MAN. Part I.

any other animal. Some savage races, such as the
Australians, are not exposed to more diversified con-
ditions than are many species which have very wide
ranges. In another and much more important re-
spect, man differs widely from any strictly domesti-
cated animal ; for his breeding has not been controlled,
either through methodical or unconscious selection. No
race or body of men has been so completely subjugated
by other men, that certain individuals have been pre-
served and thus unconsciously selected, from being in
some way more useful 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 me-
thodical selection ; for it is asserted that many tall men
were reared in the villages inhabited by the grenadiers
with their tall wives.

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 re-
stricted 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 knver animals by the same general causes,
but in both the same characters 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.13 Monstrosities, wrhich gra-

13 Godron, ' De l'Espece,' 1859, tom. ii. livre 3. Quatrefages, ' Unite
de l'Espece Humaine,' 1861. Also Lectures on Anthropology, given
in the ' Eevue des Cours Scientifiques,' 18G6-1SG8.

Chap. IV. MANNER OF DEVELOPMENT. 113

duate into slight variations, are likewise so similar in
man and the lower animals, that the same classifica-
tion and the same terms can be used for both, as may
be seen in Isidore Geoffroy St.-Hilaire's great work.14
This is a necessary consequence of the same laws of
change prevailing throughout the animal kingdom. 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 shewn 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 vari-
ability of multiple parts. Compensation of growth ;
but of this law I have found no good instances 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 ; 15 but several are so im-
portant for us, that they must be treated at consider-
able length.

The direct and definite action of changed conditions. —
This is a most perplexing subject. It- cannot be denied

14 ' Hist. Gen. et Part, des Anomalies de 1' Organisation,' in three
volumes, torn. i. 1832.

15 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 ' Do l'lnfluence
des Milieux, &c.' He lays much stress on the nature of the soil.

VOL I. I

114 THE DESCENT OF MAN. Part I.

that changed conditions produce some effect, and occa-
sionally 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 condi-
tions 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 recorded.16 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 band, it is certain that with
sailors, their manner of life delays growth, as shewn " by
" the great difference between the statures of soldiers and
" sailors at the ages of 17 and 18 years." Mr. B. A. Gould
endeavoured to ascertain the nature of the influences
which thus act on stature ; but he arrived only at nega-
tive results, namely, that they did not relate to climate,
the elevation of the land, soil, nor even " in any con-
u trolling degree " to the abundance or need of the com-

1(5 ' Investigations in Military and Anthrop. Statistics,' &c. 1869, by
B. A. Gould, p. 93, 107, 126, 131, 131.

CiiAP. IV. MANNER OF DEVELOPMENT. 115

forts of life. This latter conclusion is directly opposed
to that arrived at by Yillerme 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,17 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 in-
fluence 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."18

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, as the lungs and kidneys are brought
into fuller activity under a low temperature, and the
liver and skin under a high one.19 It was former! v
thought that the colour of the skin and the character

17 For the Polynesians, see Pricharcl's ' Physical Hist, of Mankind,'
vol. v. 1847, p. 145, 283. Also Godron, < De "l'Espece,' torn. ii. p. 289.
There is also a remarkable difference in appearance between the closely-
allied Hindoos inhabiting the Upper Ganges and Bengal ; see Elphin-
stone's ' History of India,' vol. i. p. 324.

18 * Memoirs, Anthropolog. Soc.' vol. id. 1867-69, p. 561, 565, 567.

10 Dr. Brakenridge, ' Theory of Diathesis,' ' Medical Times,' June 19
and July 17, 1869.

I 2

116 THE DESCENT OF MAN. Part I .

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 Ave treat of the different races of man-
kind. 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 evi-
dence 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 acting 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.20 Different
occupations habitually followed lead to changed pro-
portions in various parts of the body. Thus it was
clearly ascertained by the United States Commission21
that the legs of the sailors employed in the late war
were longer by 0*217 of an inch than those of the sol-
diers, 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

20 I have given authorities for these several statements in my 'Varia-
tion of Animals under Domestication,' vol. ii. p. 297-300. Dr. Jaeger,
" Ueber das Langenwachsthum der Knochen," ' Jenaischen Zeitschrift,'
B. v. Heft i.

21 ' Investigations,' &c. By B. A. Gould, 1869, p. 288.

Chap. IV. MANNER OF DEVELOPMENT. 117

their lesser height. This shortness of the arms is
apparently due to their greater use, and is an un-
expected result; but sailors chieily use their arms in
pulling and not in supporting weights. The girth of
the neck and the depth of the instep are greater, whilst
the circumference of the chest, waist, and hips is less in
sailors than in soldiers.

Whether the several foresfoins: modifications would
become hereditary, if the same habits of life were fol-
lowed during many generations, is not known, but is
probable. Rengger22 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 other analogous cases.
According to Cranz,23 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 him-
" self 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.24 From the correlation which
exists, at least in some cases,25 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,

22 * S'augetkiere yon Paraguay,' 1830, s. 4.

23 ' History of Greenland,' Eng. translat. 1767, vol. i. p. 230.

24 ' Intermarriage.' By Alex. Walker, 1838, p. 377.

25 ' The Variation of Animals under Domestication,' vol. i. p. 173.

118 THE DESCENT OF MAN. Part I.

26

is certain. But with savages, as Mr. Herbert Spencer
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;27 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 en-
gravers are liable to be short-sighted, whilst men living
much out of doors, and especially savages, are generally
long-sighted. Short-sight and long-sight certainly tend
to be inherited.28 The inferiority of Europeans, in com-
parison with savages, in eye-sight and in the other
senses, is no doubt the accumulated and transmitted
effect of lessened use during many generations; for
Eengger29 states that he has repeatedly observed Euro-
peans, 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
no doubt indicates a corresponding difference in the
dimensions of the organs themselves. Blumenbach has
also remarked on the large size of the nasal cavities

26 ( Principles of Biology,' vol. i. p. 455.

27 Paget, ' Lectures on Surgical Pathology,' vol. ii. 1853, p. 209.
2S ' The Variation of Animals under Domestication,' vol. i. p. 8.

29 ' Saugethiere von Paraguay,' s. 8, 10. I have had good oppor-
tunities for observing the extraordinary power of eyesight in the
Fuegians. See also Lawrence (' Lectures on Physiology,' &c, 1822, p.
404) on this same subject. M. Giraud-Teulon has recently collected
(' Ptevue 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.'

Chap. IV. MANNER OF DEVELOPMENT. 119

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.30

The Quechua Indians inhabit the lofty plateaux of
Peru, and Alcide d'Orbismv states31 that from con-
tinually breathing a highly rarefied atmosphere they
have acquired chests and lungs of extraordinary dimen-
sions. 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 ten and fifteen thousand feet ; and
he informs me 32 that they differ conspicuously from the
men of all other races seen by him, in the circum-
ference 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 pecu-
liarity, 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

30 Prichard, 'Phys. Hist, of Mankind,' on the authority of Blurnen-
bach, vol. i. 1851, p. 311 ; for the statement by Pallas, vol. iv. 1844, p.
407.

31 Quoted by Prichard, ' Eesearches into the Phys. Hist, of Man-
kind,' vol. v. p. 463.

32 Mr. Forbes' valuable paper is now published in the ' Journal of
the Ethnological Soc. of London,' new series, vol. ii. 1870, p. 193.

120 THE DESCENT OF MAN". Part I.

time, the femora to the tibiae were as 244 to 230 ; and
in three Negroes as 258 to 241. The humerus is like-
wise 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 sur-
vived 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 measure-
ment their bodies were found not to be so much elon-
gated as those of the men on the high plateau ; whilst
their femora had become somewhat lengthened, as had
their tibiae but in a less degree. The actual measure-
ments may be seen by consulting Mr. Forbes' memoir.
From these valuable 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.33

Although man may not have been much modified
during the latter stages of his existence through the

33 Dr. Wilckens (' Landwirthschaft. Wochenblatt,' No. 10, 1S69)
has lately published an interesting essay shewing how domestic
animals, which live in mountainous regions, have their frames
modified.

Chap IV. MANNER OF DEVELOPMENT. 121

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
bodv.

Arrests of Development — Arrested development differs
from arrested growth, as parts in the former state con-
tinue to grow whilst still retaining their early condition.
Various monstrosities come under this head, and some
are known to be occasionally inherited, as a cleft-palate.
It will suffice for our purpose to refer to the arrested
brain-development of microcephalous idiots, as described
in Vogt's great memoir.34 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 " ejfrayant " 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 gamboling and jumping
about, and making grimaces. They .often ascend stairs
on all-fours ; and are curiously fond of climbing tip
furniture or trees. We are thus reminded of the delight

34 'Mernoire sur les Microcephales,' 18(37, p. 50, 125, l^lYl^.isi-
198. /^ i* -*>'

i! LI B

<p\

~ Cs.

322 THE DESCENT OF MAX. Part I.

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.

Reversion. — Many of the cases to be here given
might have been introduced under the last heading.
Whenever 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, we may in one sense consider it as a case
of reversion. The lower members in a group give us
some idea how the common progenitor of the group was
probably constructed ; and it is hardly credible that a
part arrested at an early phase of embryonic develop-
ment should be enabled to continue growing so as ulti-
mately to perform its proper function, unless it had
acquired this power of continued growth 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. There are other cases which come more
strictly under our present heading 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 present in the normal human
embryo, they become developed in an abnormal manner,
though this manner of development is proper to the
lowrer members of the same 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 twro passages,
as in the marsupials, into a single organ, showing no
signs of doubleness except a slight internal fold, as in

Chap. IV. MANNER OF DEVELOPMENT. 12

o

the higher apes and man. The rodents exhibit a per-
fect 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. Farce "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 un-
" united. 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 so high in the scale as
in the lower apes, and their allies the lemurs.

Now in women anomalous cases are not very infre-
quent, 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 con-
" centrative 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.35 No such stage is passed
through during the ordinary development of the embryo,
and it is difficult to believe, though perhaps not im-
possible, that the two simple, minute, primitive tubes
could know how (if such an expression may be used) to

35 See Dr. A. Farre's well-known article in the ' Cyclop, of Anat.
and Phys.' vol. v. 1859, p. 642. Owen ' Anatomy of Vertebrates,' vol.
iii. 1868. p. 687. Prof. Turner in ' Edinburgh Medical Journal,' Feb.
1865.

124 THE DESCENT OF MAN. Part I.

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 develop-
ment, 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 long-
lost dormant structures are called back into existence,
might serve as the guide for the full development of
the organ, even after the lapse of an enormous interval
of time.

Professor Canestrini,36 after discussing the foregoing
and various analogous cases, arrives at the same con-
clusion as that just given. He adduces, as another
instance, the malar bone, which, in some of the Quad-
rumana and other mammals, normally consists of two
portions. This is its condition in the two-months-old
human foetus ; and thus it sometimes remains, through
arrested development, in man when adult, more especially
in the lower prognathous races. Hence Canestrini con-
cludes that some ancient progenitor of man must have
possessed this bone normally divided into two portions,
which subsequently 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 dis-
tinct suture. This suture occasionally persists, more
or less distinctly, in man after maturity, and more fre-

36 'Armuario della Soc. dei Naturalisti in 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 connexion of the two malar bones in several
human subjects and in certain apes, he cannot consider this disposition
of the parts as simply accidental.

Chap. IV. MANNEK OF DEVELOPMENT. 125

qnently 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 ana-
logons 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 cha-
racters 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 analo-
gous with the foregoing, have been advanced by dif-
ferent authors37 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 struc-
tures normally present.38

37 A whole series of cases is given by Isid. Geoffroy St.-Hilaire,
' Hist, des Anomalies,' torn. iii. p. 437.

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 ano.ther
woman with supernumerary mammas. But Prof. Preyer (' Der Kampf
nm das Dasein.' 1869, s. 45 ) states that mammas erratics have been
known to occur in other situations, even on the back ; so that the force
of my argument is greatly weakened or perhaps quite destroyed.

With much hesitation I, in the same work (vol. ii. p. 12), attributed
the frequent cases of polydactylism in men to reversion. I was partly
led to this through Prof. Owen's statement, that some of the Ichthy-
opterygia possess more than five digits, and therefore, as I supposed, had
retained a primordial condition ; but after reading Prof. Gegenbaur's
paper (' Jenaischen Zeitschrift,' B. v. Heft 3, s. 341 ), who is the highest
authority in Europe on such a point, and who disputes Owen's con-
clusion, I see that it is extremely doubtful whether supernumerary
digits can thus be accounted for. It was the fact that such digits not
only frequently occur and are strongly inherited, but have the power
of regrowth after amputation, like the normal digits of the lower verte-

126 THE DESCENT OF MAX. Part I.

In man the canine teeth are perfectly efficient instru-
ments for mastication. But their true canine character,
as Owen39 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 con-
cerned, be considered as rudimentary. In every large
collection of human skulls some may be found, as
Hackel40 observes, with the canine teeth projecting con-
siderably beyond the others in the same manner, but
in a less degree, as in the anthropomorphous apes. In
these cases, open spaces between the teeth in the one
jaw are left for the reception of the canines belonging
to the opposite jaw. An interspace of this kind in a
Kaffir skull, figured by Wagner, is surprisingly wide.41
Considering how few ancient skulls have been examined
in comparison with recent skulls, it is an interesting
fact that in at least three cases the canines project
largely ; and in the Naulette jaw they are spoken of as

enormous.42

brata, that chiefly led me to the above conclusion. This extraordinary
fact of their regrowth remains inexplicable, if the belief in reversion to
some extremely remote progenitor must be rejected. I cannot, how-
ever, follow Prof. Gegenbaur in supposing that additional digits could
not reappear through reversion, without at the same time other parts of
the skeleton being simultaneously and similarly modified ; for single
chaiacters often reappear through reversion.

39 ' Anatomy of Vertebrates,' vol. iii. 18G8, p. 323.

40 ' Generelle Morphologic,' 1866, B. ii. s. civ.

41 Carl Vogt's 'Lectures on Man,' Eng. translat. 1864, p. 151.

42 C. Carter Blake, on a jaw from La Nanlette, ' Anthiopolog.
Review,' 1867, p. 295. Schaaffhausen, ibid. 18C8, p. 426.

Chap. IV. MANNER OF DEVELOPMENT. 127

The males alone of the anthropomorphous apes have
their canines fully developed ; but in the female gorilla,
and in a less degree in the female orang, these teeth
project considerably beyond the others; therefore the
fact that women sometimes have, as I have been assured,
considerably projecting canines, is no serious objection
to the belief that their occasional great development in
man is a case of reversion to an ape-like progenitor.
He who rejects with scorn the belief that the shape of
his own canines, and their occasional great development
in other men, are due to our early progenitors having
been provided with these formidable weapons, will pro-
bably reveal by sneering the line of his descent. For
though he no longer intends, nor has the power, to use
these teeth as weapons, he will unconsciously retract his
" snarling muscles ,!l (thus named by Sir C. Bell) 43 so as
to expose them readv for action, like a dog prepared to
fight.

Many muscles are occasionally developed in man,
which are proper to the Quadrumana or other mam-
mals. Professor Vlacovich44 examined fortv male sub-
jects, and found a muscle, called by him the ischio-
pubic, in nineteen of them ; in three others there was
a ligament which represented this muscle ; and in the
remaining eighteen no trace of it. Out of thirty female
subjects this muscle was developed on both sides in only
two, but in three others the rudimentary ligament was
present. This muscle, therefore, appears to be much
more common in the male than in the female sex ; and
on the principle of the descent of mairfrom some lower
form, its presence can be understood ; for it has been
detected in several of the lower animals, and in all of

43 I

The Anatomy of Expression,' 1844, p. 110, 131.
44 Quoted by Prof. Canestrini iu the ' Annuario,' &e., 1 SG7, p. 90.

128 THE DESCENT OF MAN. Part 1.

these it serves exclusively to aid the male in the act
of reproduction.

Mr. J. Wood, in his valuable series of papers,45 has
minutely described a vast number of muscular varia-
tions in man, which resemble normal structures in the
lower animals. Looking only to the muscles which
closely resemble those regularly present in our nearest
allies, the Quaclrumana, they are too numerous to be
here even specified. In a single male subject, having
a strong bodily frame and well-formed skull, no less
than seven muscular variations were observed, all of
which plainly represented muscles proper to various
kinds of apes. This man, for instance, had on both
sides of his neck a true and powerful " levator clavi-
cular" such as is found in all kinds of apes, and which
is said to occur in about one out of sixty human sub-
jects.46 Again, this man had "a special abductor of
" the metatarsal bone of the fifth digit, such as Pro-
" fessor Huxley and Mr. Flower have shewn to exist
" uniformly in the higher and lower apes." The hands
and arms of man are eminently characteristic structures,
but their muscles are extremely liable to vary, so as to
resemble the corresponding muscles in the lower ani-
mals.47 Such resemblances are either complete and per-

45 These papers deserve careful study by any one who desires to learn
how frequently our muscles vary, and in varying come to resemble those
of the Quadrumana. The following references relate to the few points
touched on in my text : Proc. Eoyal Soc. vol. xiv. 1805, p. 379-384 ;
vol. xv. 1866, p. 241, 242 ; vol. xv. 1867, p. 544 ; vol. xvi. 1868, p. 524. I
may here add that Dr. Murie and Mr. St. George Mivart have shewn
in their Memoir on the Lemuroidea (' Transact. Zoolog. Soc' vol. vii.
1869, p. 96), how extraordinarily variable some of the muscles are in
these animals, the lowest members of the Primates. Gradations, also,
in the muscles leading to structures found in animals still lower in
the scale, are numerous in the Lemuroidea.

46 Prof. Macalister in 'Proc. R. Irish Academy,' vol. x. 1868, p. 124.

47 Prof. Macalister (ibid. p. 121) has tabulated his observations, and
finds that muscular abnormalities are most frequent in the forearms,
secondly in the face, thirdly in the foot, &c.

Chap. IV. MANNER OF DEVELOPMENT. 129

feet or imperfect, yet in tins latter case manifestly of
a transitional nature. Certain variations are more com-
mon in man, and others in woman, without our being
able to assign any reason. Mr. Wood, after describing
numerous cases, makes the following pregnant remark :
" Notable departures from the ordinary type of the
" muscular structures run in grooves or directions, which
" must be taken to indicate some unknown factor, of
" much importance to a comprehensive knowledge of
" general and scientific anatomy." 48

That this unknown factor is reversion to a former
state of existence may be admitted as in the highest
degree probable. It is quite incredible that a man
should through mere accident abnormally resemble, in
no less than seven of his muscles, certain apes, if there
had been no genetic connection between them. On the
other hand, if man is descended from some ape-like
creature, no valid reason can be assigned why certain
muscles should not suddenly reappear after an interval of
many thousand generations, in the same manner as with
horses, asses, and mules, dark-coloured stripes suddenly
reappear on the legs and shoulders, after an interval of
hundreds, or more probably thousands, of generations.

These various cases of reversion are so closely related

48 The Eev. Dr. Hatighton, after giving (' Proc. R. Irish Academy/
June 27, 1864, p. 715) a remarkable case of variation in the human
flexor pollicis longus, adds, " This remarkable example shews that man
" may sometimes possess the arrangement of tendons of thumb and
" fingers characteristic of the macaque ; but whether such a case should
" be regarded as a macaque passing upwards -into a man, or a man
" passing downwards into a macaque, or as a congenital freak of
" nature, I cannot undertake to say." It is satisfactory to hear so
capable an anatomist, and so embittered an opponent of evolutionism,
admitting even the possibility of either of his first propositions. Prof.
Macalister has also described (' Proc. R. Irish Acad.' vol. x. 1864, p.
138) variations in the flexor pollicis longus, remarkable from their rela-
tions to the same musole in the Quadrumana.

VOL. I. K

130 THE DESCENT OF MAN. Part I.

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

Correlated Variation. — In man, as in the lower ani-
mals, many structures are so intimately related, that
when one part varies so does another, without our being
able, in most cases, to assign any reason. We cannot say
whether the one part governs the other, or whether both
are governed by some earlier developed part. Various
monstrosities, as I. Geoffroy repeatedly insists, are thus
intimately connected. Homologous structures are par-
ticularly liable to change together, as we see on the
opposite sides of the body, and in the upper and lower
extremities. Meckel long ago remarked that when the
muscles of the arm depart from their proper type, they
almost always imitate those of the leg ; and so conversely
with the muscles of the legs. The organs of sight and
hearing, the teeth and hair, the colour of the skin and
hair, colour and constitution, are more or less correlated.49
Professor Schaaffhausen first drew attention to the rela-

49 The authorities for these several statements are given in my
Variation of Animals under Domestication,' vol. ii. p. 320-335.

Chap. IV. MANNEK OF DEVELOPMENT. 131

tion apparently existing between a muscular frame and
strongly-pronounced supra-orbital ridges, which are so
characteristic of the lower races of man.

Besides the variations which can be grouped with
more or less probability under the foregoing heads,
there is a large class of variations which may be provi-
sionally called spontaneous, for they appear, owing to our
ignorance, to arise without any exciting cause. It can,
however, be shewn that such variations, whether con-
sisting of slight individual differences, or of strongly-
marked and abrupt deviations of structure, depend
much more on the constitution of the organism than
on the nature of the conditions to which it has been
subjected.5

50

Piate of Increase. — Civilised populations have been
known under favourable conditions, as in the United
States, to double their number in twenty -five years ;
and according to a calculation by Euler, this might
occur in a little over twelve years.51 At the former rate
the present population of the United States, namely,
thirty millions, would in 657 years cover the whole
terraqueous globe so thickly, that four men would have
to stand on each square yard of surface. The primary
or fundamental check to the continued increase of man
is the difficulty of gaining subsistence and of living in
comfort. We may infer that this is the case from what
we see, for instance, in the United States, where subsist-
ence is easy and there is plenty of room. If such means
were suddenly doubled in Great Britain, our number
would be quickly doubled. With civilised nations the

50 This whole subject has been discussed in chap, xxiii. vol. ii. of
my ' Variation of Animals and Plants under Domestication.'

51 See the ever memorable ' Essay on the Principle of Population.'
by the Eev. T. Malthus, vol. i. 1826, p. 6, 517.

K 2

132 THE DESCENT OF MAN. Part I.

above primary check acts chiefly by restraining mar-
riages. The greater death-rate of infants in the poorest
classes is also very important; as well as the greater
mortality at all ages, and from various diseases, of the
inhabitants of crowded and miserable houses. The
effects of severe epidemics and wars are soon counter-
balanced, and more than counterbalanced, in nations
placed under favourable conditions. Emigration also
comes in aid as a temporary check, but not to any
great extent with the extremely poor classes.

There is reason to suspect, as Malthus has remarked,
that the reproductive power is actually less in barbarous
than in civilised races. We know nothing positively on
this head, for with savages no census has been taken ;
but from the concurrent testimony of missionaries, and
of others who have long resided with such people, it
appears that their families are usually small, and large
ones rare. This may be partly accounted for, as it is
believed, by the women suckling their infants for a pro-
longed period; but it is highly probable that savages,
who often suffer much hardship, and who do not obtain so
much nutritious food as civilised men, would be actually
less prolific. I have shewn in a former work,52 that
all our domesticated quadrupeds and birds, and all
our cultivated plants, are more fertile than the corre-
sponding species in a state of nature. It is no valid
objection to this conclusion that animals suddenly
supplied with an excess of food, or when rendered very
fat, and that most plants when suddenly removed from
very poor to very rich soil, are rendered more or less
sterile. We might, therefore, expect that civilised
men, who in one sense are hifrhlv domesticated, would

52 ' Variation of Animals and Plants under Domestication,' vol. ii.
p. 111-113, 163.

Chap. IV. MANNER OF DEVELOPMENT. 133

be more prolific than wild men. Tt is also probable
that the increased fertility of civilised nations would
become, as with our domestic animals, an inherited
character: it is at least known that with mankind a
tendency to produce twins runs in families.53

Notwithstanding- that savages appear to be less pro-
lific than civilised people, they would no doubt rapidly
increase if their numbers were not by some means
rigidly kept down. The Santali, or hill-tribes of India,
have recently afforded a good illustration of this fact ;
for they have increased, as shewn by Mr. Hunter,54
at an extraordinary rate since vaccination has been
introduced, other pestilences mitigated, and war sternly
repressed. This increase, however, would not have been
possible had not these rude people spread into the
adjoining districts and worked for hire. Savages almost
always marry; yet there is some prudential restraint,
for they do not commonly marry at the earliest possible
age. The young men are often required to show that
they can support a wife, and they generally have first
to earn the price with which to purchase her from her
parents. With savages the difficulty of obtaining sub-
sistence occasionally limits their number in a much
more direct manner than with civilised people, for all
tribes periodically suffer from severe famines. At such
times savages are forced to devour much bad food,
and their health can hardly fail to be injured. Many
accounts have been published of their protruding sto-
machs and emaciated limbs after and during famines.
They are then, also, compelled to wander much about,
and their infants, as I was assured in Australia, perish

53 Mr. Sedgwick, ' British and Foreign Medico-Clrirurg. Review, '
July, 18G3, p. 170.

54 ' The Annals of Rural Bengal,' by W. W. Hunter, 1868, p. 259.

134 THE DESCENT OF MAN. Part I.

in large numbers. As famines are periodical, depending
chiefly on extreme seasons, all tribes must fluctuate in
number. They cannot steadily and regularly increase,
as there is no artificial increase in the supply of food.
Savages when hardly pressed encroach on each other's
territories, and war is the result ; but they are indeed
almost always at war with their neighbours. They are
liable to many accidents on land and water in their search
for food ; and in some countries they must suffer much
from the larger beasts of prey. Even in India, districts
have been depopulated by the ravages of tigers.

Malthus has discussed these several checks, but he
does not lay stress enough on what is probably the most
important of all, namely infanticide, especially of female
infants, and the habit of procuring abortion. These
practices now prevail in many quarters of the world,
and infanticide seems formerly to have prevailed, as
Mr. M'Lennan55 has shewn, on a still more extensive
scale. These practices appear to have originated in
savages recognising the difficulty, or rather the impos-
sibility of supporting all the infants that are born.
Licentiousness may also be added to the foregoing
checks ; but this does not follow from failing means of
subsistence ; though there is reason to believe that in
some cases (as in Japan) it has been intentionally
encouraged as a means of keeping down the population.

If we look back to an extremely remote epoch, before
man had arrived at the dignity of manhood, he would
have been guided more by instinct and less by reason
than are savages at the present time. Our early semi-
human progenitors would not have practised infanticide,
for the instincts of the lower animals are never so per-
verted as to lead them regularly to destroy their own

55

' Primitive Marriage/ 1865.

Chap. IV. MANNER OF DEVELOPMENT. 135

offspring. There would have been no prudential re-
straint from marriage, and. the sexes would have freely
united at an early age. Hence the progenitors of
man would have tended to increase rapidly, but checks
of some kind, either periodical or constant, must have
kept down their numbers, even more severely than with
existing savages. What the precise nature of these
checks may have been, we cannot say, any more than
with most other animals. We know that horses and
cattle, which are not highly prolific animals, when first
turned loose in South America, increased at an enormous
rate. The slowest breeder of all known animals, namely
the elephant, would in a few thousand years stock the
whole world. The increase of every species of monkey
must be checked by some means ; but not, as Brehm
remarks, by the attacks of beasts of prey. No one
will assume that the actual power of reproduction in
the wild horses and cattle of America, was at first in
any sensible degree increased ; or that, as each district
became fully stocked, this same power was diminished.
No doubt in this case and in all others, many checks
concur, and different checks under different circum-
stances ; periodical dearths, depending on unfavourable
seasons, being probably the most important of all. So
it will have been with the early progenitors of man.

Natural Selection. — We have now seen that man is
variable in boclv and mind; and that the variations
are induced, either directly or indirectly, by the same
general causes, and obey the same general laws, as with
the lower animals. Man has spread widely over the
face of the earth, and must have been exposed, during
his incessant migrations,56 to the most diversified con-

56 See some good remarks to this effect by W. Stanley Jevons, " A
Deduction from Darwin's Theory," ' Nature,' 1869, p. 231.

136 THE DESCENT OF MAN. Part I.

ditions. The inhabitants of Tierra del Fuego, the Cape
of Good Hope, and Tasmania in the one hemisphere,
and of the Arctic regions in the other, must have passed
through many climates and changed their habits many-
times, before they reached their present homes.57 The
early progenitors of man must also have tended, like all
other animals, to have increased beyond their means of
subsistence ; they must therefore occasionally have been
exposed to a struggle for existence, and consequently to
the rigid law of natural selection. Beneficial variations
of all kinds will thus, either occasionally or habitually,
have been preserved, and injurious ones eliminated. I
do not refer to strongly-marked deviations of structure,
which occur only at long intervals of time, but to mere
individual differences. We know, for instance, that the
muscles of our hands and feet, which determine our
powers of movement, are liable, like those of the lower-
animals,58 to incessant variability. If then the ape-like
progenitors of man which inhabited any district, espe-
cially one undergoing some change in its conditions, were
divided into two equal bodies, the one half which in-
cluded all the individuals best adapted by their powers
of movement for gaining subsistence or for defending
themselves, would on an average survive in greater
number and procreate more offspring than the other
and less well endowed half.

Man in the rudest state in which he now exists is
the most dominant animal that has ever appeared
on the earth. He has spread more widely than any

37 Latham, ' Man and his Migrations,' 1851, p. 135.

58 Messrs. Murie and Mivart in their " Anatomy of the Lemuroidea"
(' Transact. Zoolog. Soc.' vol. vii. 1869, p. 96-98) say, " some muscles
" are so irregular in their distribution that they cannot be well classed
" in any of the above groups." These muscles differ even on the oppo-
site sides of the same individual.

Chap. IV. MANNEK OF DEVELOPMENT. 137

other highly organised form ; and all others have
yielded before him. He manifestly owes this immense
superiority to his intellectual faculties, his social habits,
which lead him to aid and defend his fellows, and to
his corporeal structure. The supreme importance of
these characters has been proved by the final arbitra-
ment of the battle for life. Through his powers of in-
tellect, articulate language has been evolved; and on
this his wonderful advancement has mainly depended.
He has invented and is able to use various weapons,
tools, traps, &c, with which he defends himself, kills or
catches prey, and otherwise obtains food. He has made
rafts or canoes on which to fish or cross over to neigh-
bouring fertile islands. He has discovered the art of
making fire, by which hard and stringy roots can be
rendered digestible, and poisonous roots or herbs in-
nocuous. This last discovery, probably the greatest,
excepting language, ever made by man, dates from
before the dawn of history. These several inventions,
by which man in the rudest state has become so pre-
eminent, are the direct result of the development of
his powers of observation, memory, curiosity, imagina-
tion, and reason. I cannot, therefore, understand how
it is that Mr. Wallace 59 maintains, that " natural selec-

59 ' Quarterly Review,' April, 18G9, p. 392. This subject is more
fully discussed in Mr. Wallace's ' Contributions to the Theory of Natural
Selection,' 1870, in which all the essays referred to in this work are
republished. The ' Essay on Man ' has been ably criticised by Prof.
Claparede, one of the most distinguished zoologists in Europe, in an
article published in the ' Bibliotheque Universelle,' June, 1S70. The
remark quoted in my text will surprise every one who has read
Mr. Wallace's celebrated paper on 'The Origin of Human Paces
deduced from the Theory of Natural Selection,' originally publishec
in the ' Anthropological Review,' May, 1864, p. clviii. I cannot h#re^
resist quoting a most just remark by Sir J. Lubbock (' Prehis^*^ O
Times,' 1865, p. 479) in reference to this paper, namely, thay-ifc\x ^j^i
Wallace, "with characteristic unselfishness, ascribes it (i.e. the iilb&or

TuJ LIB I

\t- , ^

138 THE DESCENT OF MAX. Part I.

" tion could only have endowed the savage with a brain
" a little superior to that of an ape."

Although the intellectual powers and social habits of
man are of paramount importance to him, we must not
underrate the importance of his bodily structure, to which
subject the remainder of this chapter will be devoted.
The development of the intellectual and social or moral
faculties will be discussed in the following chapter.

Even to hammer with precision is no easy matter, as
every one who has tried to learn carpentry will admit.
To throw a stone with as true an aim as can a Fueofian in
defending himself, or in killing birds, requires the most
consummate perfection in the correlated action of the
muscles of the hand, arm, and shoulder, not to mention
a fine sense of touch. In throwing a stone or spear, and
in many other actions, a man must stand firmly on his
feet ; and this again demands the perfect coadaptation of
numerous muscles. To chip a flint into the rudest tool,
or to form a barbed spear or hook from a bone, demands
the use of a perfect hand ; for, as a most capable judge,
Mr. Schoolcraft,60 remarks, the shaping fragments of
stone into knives, lances, or arrow-heads, shews " extra-
" ordinary ability and long practice." We have evidence
of this in primeval men having practised a division of
labour ; each man did not manufacture his own flint
tools or rude pottery ; but certain individuals appear to
have devoted themselves to such work, no doubt re-
ceiving in exchange the produce of the chase. Archaeo-
logists are convinced that an enormous interval of time

" natural selection) unreservedly to Mr. Darwin, although, as is well
" known, he struck out the idea independently, and published it,
" though not with the same elaboration, at the same time."

60 Quoted by Mr. Lawson Tait in his " Law of Natural Selection,"
— 'Dublin Quarterly Journal of Medical Science,' Feb. 1869. Dr.
Keller is likewise quoted to the same effect.

Chap. IV. MANNER OF DEVELOPMENT. 139

elapsed before our ancestors thought of grinding chipped
flints into smooth tools. A man-like animal who pos-
sessed a hand and arm sufficiently perfect to throw a
stone with precision or to form a flint into a rude tool,
could, it can hardly be doubted, with sufficient practice
make almost anything, as far as mechanical skill alone
is concerned, which a civilised man can make. The
structure of the hand in this respect may be compared
with that of the vocal organs, which in the apes are
used for uttering various signal-cries, or, as in one spe-
cies, musical cadences ; but in man closely similar vocal
organs have become adapted through the inherited
effects of use for the utterance of articulate language.

Turning now to the nearest allies of man, and there-
fore to the best representatives of our early progenitors,
we find that the hands in the Quadrumana are con-
structed on the same general pattern as in us, but are
far less perfectly adapted for diversified uses. Their
hands do not serve so well as the feet of a dog for loco-
motion ; as may be seen in those monkeys which walk
on the outer margins of the palms, or on the backs of
their bent fingers, as in the chimpanzee and orang.16
Their hands, however, are admirably adapted for climb-
ing trees. Monkeys seize thin branches or ropes, with
the thumb on one side and the fingers and palm on
the other side, in the same manner as we do. They
can thus also carry rather large objects, such as the
neck of a bottle, to their mouths. Baboons turn over
stones and scratch up roots with their hands. They
seize nuts, insects, or other sinalL objects with the
thumb in opposition to the fingers, and no doubt they
thus extract eggs and the young from the nests of
birds. American monkeys beat the wild oranges on the

61 Owen, ' Anatomy of Vertebrates,' vol. iii. p. 71.

140 THE DESCENT OF MAN. Part I.

branches until the rind is cracked, and then tear it off
with the fingers of the two hands. Other monkeys open
mussel-shells with the two thumbs. With their fingers
they pull out thorns and burs, and hunt for each other's
parasites. In a state of nature they break open hard
fruits with the aid of stones. They roll down stones
or throw them at their enemies ; nevertheless, they
perform these various actions clumsily, and they are
quite unable, as I have myself seen, to throw a stone
with precision.

It seems to me far from true that because " objects
" are grasped clumsily " by monkeys, " a much less
" specialised organ of prehension " would have served
them62 as well as their present hands. On the con-
trary, I see no reason to doubt that a more perfectly
constructed hand would have been an advantage to them,
provided, and it is important to note this, that their
hands had not thus been rendered less well adapted for
climbing trees. We may suspect that a perfect hand
would have been disadvantageous for climbing ; as the
most arboreal monkeys in the world, namely Ateles in
America and Hylobates in Asia, either have their thumbs
much reduced in size and even rudimentary, or their
fingers partially coherent, so that their hands are con-
verted into mere grasping-hooks.63

As soon as some ancient member in the great series
of the Primates came, owing to a change in its manner
of procuring subsistence, or to a change in the con-

62 'Quarterly Review,' April, 1869, p. 392.

63 In Hylobates syndactylus, as the name expresses, two of the digits
regularly cohere ; and this, as Mr. Blyth informs me, is occasiorjally
the case with the digits of H. agilis, lar, and leuciscm. In Colobus the
thumb is likewise deficient ; these monkeys are strictly arboreal and
extraordinarily active (Brehni, ' Thierleben,' B. i. s. 50), but whether
they are better climbers or graspers than the species of the allied
genera, I do not know.

Chap. IV. MANNER OF DEVELOPMENT. 141

ditions of its native country, to live somewhat less on
trees and more on the ground, its manner of progres-
sion would have been modified ; and in this case it
would have had to become either more strictly quad-
rupedal or bipedal. Baboons frequent hilly and rocky
districts, and only from necessity climb up high trees ; M
and they have acquired almost the gait of a clog. Man
alone has become a biped ; and we can, I think, partly
see how he has come to assume his erect attitude, which
forms one of the most conspicuous differences between
him and his nearest allies. Man could not have
attained his present dominant position in the world
without the use of his hands which are so admirably
adapted to act in obedience to his will. As Sir C. Bell 65
insists " the hand supplies all instruments, and by its
" correspondence with the intellect gives him universal
" dominion." But the hands and arms could hardly
have become perfect enough to have manufactured
weapons, or to have hurled stones and spears with a
true aim, as long as they were habitually used for
locomotion and for supporting the whole weight of the
body, or as long as they were especially well adapted,
as previously remarked, for climbing trees. Such rough
treatment would also have blunted the sense of touch,
on which their delicate use largely depends. From
these causes alone it would have been an advantage to
man to have become a biped ; but for many actions it is
almost necessary that both arms and the whole upper
part of the body should be free ; and he must for this
end stand firmly on his feet. To gain this great
advantage, the feet have been rendered flat, and the
great toe peculiarly modified, though this has entailed
the loss of the power of prehension. It accords with

64 Brehm, ' Thierleben,' B. i. s. 80.

65 " The Hand," &c. 4 Bridgewater Treatise,' 1S33, p. 38.

142 THE DESCENT OF MAN. Part 1.

the principle of the division of physiological labour,
which prevails throughout the animal kingdom, that
as the hands became perfected for prehension, the
feet should have become perfected for support and
locomotion. With some savages, however, the foot has
not altogether lost its prehensile power, as shewn by
their manner of climbing trees and of using them in
other ways.66

If it be an advantage to man to have his hands and
arms free and to stand firmly on his feet, of which there
can be no doubt from his pre-eminent success in the
battle of life, then I can see no reason why it should
not have been advantageous to the progenitors of man
to have become more and more erect or bipedal. They
would thus have been better able to have defended
themselves with stones or clubs, or to have attacked
their prey, or otherwise obtained food. The best con-
structed individuals would in the long run have succeeded
best, and have survived in larger numbers. If the
gorilla and a few allied forms had become extinct, it
might have been argued with great force and apparent
truth, that an animal could not have been gradually
converted from a quadruped into a biped; as all the
individuals in an intermediate condition would have
been miserably ill-fitted for progression. But we know
(and this is well worthy of reflection) that several kinds
of apes are now actually in this intermediate condition ;
and no one doubts that they are on the whole well
adapted for their conditions of life. Thus the gorilla

66 H'ackel has an excellent discussion on the steps by which man
became a biped : ' Naiiirliche Schopfungsgeschichte,' 1868, s. 507. Dr.
Biichuer (' Conferences sur la The'orie Darwinienne,' 18b'9, p. 135) has
given good cases of the use of the foot as a prehensile organ by man ;
also on the manner of progression of the higher apes to which I allude
in the following paragraph : see also Owen (' Anatomy of Vertebrates,'
vol. iii. p. 71) on this latter subject.

Chap. IV. MANNER OF DEVELOPMENT. 143

runs with a sidelong shambling gait, but more commonly
progresses by resting on its bent hands. The long-
armed apes occasionally use their arms like crutches,
swinging their bodies forward between them, and some
kinds of Hylobates, without having been taught, can
walk or run upright with tolerable quickness ; yet they
move awkwardly, and much less securely than man.
We see, in short, with existing monkeys various grada-
tions between a form of progression strictly like that of
a quadruped and that of a biped or man.

As the progenitors of man became more and more
erect, with their hands and arms more and more modi-
fied for prehension and other purposes, with their feet
and legs at the same time modified for firm support
and progression, endless other changes of structure
would have been necessary. The pelvis would have
had to be made broader, the spine peculiarly curved
and the head fixed in an altered position, and all these
changes have been attained by man. Prof. Schaaff-
hausen67 maintains that " the powerful mastoid processes
of the human skull are the result of his erect position ; "
and these processes are absent in the orangj chim-
panzee, &c, and are smaller in the gorilla than in man.
Various other structures might here have been specified,
which appear connected with man's erect position. It
is very difficult to decide how far all these correlated
modifications are the result of natural selection, and
how far of the inherited effects of the increased use of
certain parts, or of the action of one part on another.
No doubt these means of change act .and react on each
other : thus when certain muscles, and the crests of
bone to which they are attached, become enlarged by

67 "On the Primitive Form of the Skull," translated in 'Anthropo-
logical Review,' Oct. 1868, p. 428. Owen (' Anatomy of Vertebrates,'
vol. ii. 1SUG, p. 551) on the mastoid processes in the higher apes.

144 THE DESCENT OF MAN. Part I.

habitual use, this shews that certain actions are
habitually performed and must be serviceable. Hence
the individuals which performed them best, would tend
to survive in greater numbers.

The free use of the arms and hands, partly the cause
and partly the result of man's erect position, appears to
have led in an indirect manner to other modifications of
structure. The early male progenitors of man were, as
previously stated, probably furnished with great canine
teeth ; but as they gradually acquired the habit of using
stones, clubs, or other weapons, for fighting with their
enemies, they would have used their jaws and teeth less
and less. In this case, the jaws, together with the
teeth, would have become reduced in size, as we may
feel sure from innumerable analogous cases. In a future
chapter we shall meet with a closely-parallel case, in
the reduction or complete disappearance of the canine
teeth in male ruminants, apparently in relation with the
development of their horns ; and in horses, in relation
with their habit of fighting with their incisor teeth and
hoofs.

In the adult male anthropomorphous apes, as Riiti-
meyer,68 and others have insisted, it is precisely the effect
which the jaw-muscles by their great development have
produced on the skull, that causes it to differ so greatly
in many respects from that of man, and has given to
it " a truly frightful physiognomy." Therefore as the
jaws and teeth in the progenitors of man gradually
become reduced in size, the adult skull would have
presented nearly the same characters which it offers in
the young of the anthropomorphous apes, and would
thus have come to resemble more nearly that of existing

68

'Die Grenzen der Thierwelt, eine Betrachtung zu Darwin's

Lehre,' 1868, s. 51.

Chap. IV. MANNER OF DEVELOPMENT. 145

man. A great reduction of the canine teeth in the
males would almost certainly, as we shall hereafter see,
have affected through inheritance the teeth of the
females.

As the various mental faculties were gradually de- •
veloped, the brain would almost certainly have become
larger. No one, I presume, doubts that the large size
of the brain in man, relatively to his body, in compari-
son with that of the gorilla or orang, is closely con-
nected with his higher mental powers. We meet with
closely analogous facts with insects, in which the cerebral
ganglia are of extraordinary dimensions in ants; these
ganglia in all the Hymenoptera being many times larger
than in the less intelligent orders, such as beetles.69
On the other hand, no one supposes that the intellect o
any two animals or of any two men can be accurately
gauged bv the cubic contents of their skulls. It is
certain that there may be extraordinary mental activity
with an extremely small absolute mass of nervous
matter : thus the wonderfully diversified instincts,
mental powers, and affections of ants are generally
known, yet their cerebral ganglia are not so large as the
quarter of a small pin's head. Under this latter point
of view, the brain of an ant is one of the most marvellous
atoms of matter in the world, perhaps more marvellous
than the brain of man.

The belief that there exists in man some close relation
between the size of the brain and the development of
the intellectual faculties is supported by the comparison
of the skulls of savage and civilised races, of ancient and
modern people, and by the analogy of the whole verte-

6D

Dujardin, ' Annales des Sc. Nat.' 3rd series, Zoolog. torn. xiv.
1850, p. 203. See also Mr. Lowne, ' Anatomy and Phys. of the Musca
vomitoria,' 1870, p. 14. My son, Mr. F. Darwin, dissected for me the
cerebral ganglia of the Formica rufa.

VOL. I. l L

146 THE DESCENT OF MAN. Part I.

brate series. Dr. J. Barnard Davis has proved70 by
many careful measurements, that the mean internal
capacity of the skull in Europeans is 92*3 cubic inches ;
in Americans 87*5 ; in Asiatics 87*1 ; and in Australians
only 81*9 inches. Professor Broca71 found that skulls
from graves in Paris of the nineteenth century, were
larger than those from vaults of the twelfth century, in
the proportion of 1484 to 1426 ; and Prichard is per-
suaded that the present inhabitants of Britain have
"much more capacious brain-cases" than the ancient
inhabitants. Nevertheless it must be admitted that
some skulls of very high antiquity, such as the famous
one of Neanderthal, are well developed and capacious.
With respect to the lower animals, M. E. Lartet,72 by com-
paring the crania of tertiary and recent mammals, be-
longing to the same groups, has come to the remarkable
conclusion that the brain is generally larger and the
convolutions more complex in the more recent form.
On the other hand I have shewn 73 that the brains of
domestic rabbits are considerably reduced in bulk, in
comparison with those of the wild rabbit or hare ; and
this may be attributed to their having been closely con-
fined during many generations, so that they have exerted
but little their intellect, instincts, senses, and voluntary
movements.

The gradually increasing weight of the brain and
skull in man must have influenced the development of
the supporting spinal column, more especially whilst
he was becoming erect. As this change of position was

70 ' Philosophical Transactions,' ] 869, p. 513.

71 Quoted in C. Vogt's 'Lectures on Man,' En<r. translat. 1864, p.
88. 90. Prichard, ' Phys. Hist, of Mankind,' vol. i. 1838, p. 305.

72 ' Comptes Eendus des Seances,' &c. June 1, 1868.

73 ' The Variation of Animals and Plants under Domestication,' vo
i. p. 124-129.

Chap. IV. MANNER OF DEVELOPMENT. 147

being brought about, the internal pressure of the brain,
will, also, have influenced the form of the skull; for
many facts shew how easily the skull is thus affected.
Ethnologists believe that it is modified by the kind of
cradle in which infants sleep. Habitual spasms of the
muscles and a cicatrix from a severe burn have per-
manently modified the facial bones. In young persons
whose heads from disease have become fixed either
sideways or backwards, one of the eyes has changed
its position, and the bones of the skull have been
modified ; and this apparently results from the brain
pressing in a new direction.74 I have shewn that with
long-eared rabbits, even so trifling a cause as the lopping
forward of one ear drags forward on that side almost
every bone of the skull ; so that the bones on the oppo-
site sides no longer strictly correspond. Lastly, if any
animal were to increase or diminish much in general
size, without any change in its mental powers ; or if
the mental powers were to be much increased or
diminished without any great change in the size of the
body ; the shape of the skull would almost certainly be
altered. I infer this from my observations on domestic
rabbits, some kinds of which have become very much
larger than the wild animal, whilst others have retained
nearly the same size, but in both cases the brain has
been much reduced relatively to the size of the body.
Now I was at first much surprised by finding that in all
these rabbits the skull had become elongated or dolicho-

74 Schaaff hausen gives from Blumenbach and Busch, the cases of the
spasms and cicatrix, in ' Anthropolog. Review,' Oct. 1868, p. 420. Dr.
Jarrold (' Anthropologia,' 1808, p. 115, 116) adduces from Camper and
from bis own observations, cases of the modification of the skull from
the head being fixed in an unnatural position. He believes that cer-
tain trades, such as that of a shoemaker, by causing the head to be
habitually held forward, makes the forehead more rounded and
prominent.

L _

\&

148 THE DESCENT OF MAN. Part I.

cephalic ; for instance, of two skulls of nearly equal
breadth, the one from a wild rabbit and the other from
a large domestic kind, the former was only 3*15 and the
latter 4*3 inches in length.75 One of the most marked
distinctions in different races of man is that the skull
in some is elongated, and in others rounded ; and here
the explanation suggested by the case of the rabbits
may partially hold good ; for Welcker finds that short
" men incline more to brachycephaly, and tall men to
dolichocephaly ; " 76 and tall men may be compared with
the larger and longer-bodied rabbits, all of which have
elongated skulls, or are dolichocephalic.

From these several facts we can to a certain ex-
tent understand the means through which the great
size and more or less rounded form of the skull has
been acquired by man ; and these are characters emi-
nently distinctive of him in comparison with the lower
animals.

Another most conspicuous difference between man and
the lower animals is the nakedness of his skin. Whales
and dolphins (Cetacea), dugongs (Sirenia) and the hip-
popotamus are naked ; and this may be advantageous
to them for gliding through the water ; nor would it
be injurious to them from the loss of warmth, as the
species which inhabit the colder regions are protected
by a thick layer of blubber, serving the same purpose
as the fur of seals and otters. Elephants and rhino-
ceroses are almost hairless ; and as certain extinct
species which formerly lived under an arctic climate
were covered with long wool or hair, it would almost
appear as if the existing species of both genera had lost

75 'Variation of Animals,' &c, vol. i. p. 117 on the elongation of the
skull; p. 119, on the effect of the lopping of one ear.

76 Quoted by Schaaflhausun, in ' Anthropolog. Review,' Oct. 1868,
p. 419.

Chap. IV. MANNER OF DEVELOPMENT. 149

their hairy covering from exposure to heat. This appears
the more probable, as the elephants in India which live
on elevated and cool districts are more hairy77 than those
on the lowlands. May we then infer that man became
divested of hair from having aboriginally inhabited some
tropical land ? The fact of the hair being chiefly
retained in the male sex on the chest and face, and in
both sexes at the junction of all four limbs with the
trunk, favours this inference, assuming that the hair was
lost before man became erect ; lor the parts which now
retain most hair would then have been most protected
from the heat of the sun. The crown of the head,
however, offers a curious exception, for at all times it
must have been one of the most exposed parts, yet
it is thickly clothed with hair. In this respect man
agrees with the great majority of quadrupeds, which
generally have their upper and exposed surfaces more
thickly clothed than the lower surface. Nevertheless,
the fact that the other members of the order of Pri-
mates, to which man belongs, although inhabiting vari-
ous hot regions, are well clothed with hair, generally
thickest on the upper surface,78 is strongly opposed
to the supposition that man became naked through the
action of the sun. I am inclined to believe, as we
shall see under sexual selection, that man, or rather
primarily woman, became divested of hair for orna-
mental purposes ; and according to this belief it is not

77 Owen, l Anatomy of Vertebrates,' vol. iii. p. 619.

78 Isidore Geoffroy St.-Hilaire remarks (' Hi§t. Nat. Generale,' torn,
ii. 1859, p. 215-217; on the head of man being covered with long hair ;
also on the upper surfaces of monkeys and of other mammals being
more thickly clothed than the lower surfaces. This has likewise been
observed by various authors. Prof. P. Gervais ( ' Hist. Nat. des Mam-
miferes,' torn. i. 1854, p. 28), however, states that in the Gorilla the
hair is thinner on the back, where it is partly rubbed off, than on the
lower surface.

150 THE DESCENT OF MAN. Part I.

surprising that man should differ so greatly in hairi-
ness from all his lower brethren, for characters gained
through sexual selection often differ in closely-related
forms to an extraordinary degree.

According to a popular impression, the absence of a
tail is eminently distinctive of man ; but as those apes
which come nearest to man are destitute of this organ,
its disappearance does not especially concern us. Never-
theless it may be well to own that no explanation, as
far as I am aware, has ever been given of the loss of the
tail by certain apes and man. Its loss, however, is not
surprising, for it sometimes differs remarkably in length
in species of the same genera: thus in some species
of Macacus the tail is longer than the whole body, con-
sisting of twenty-four vertebrae ; in others it consists of a
scarcely visible stump, containing only three or four
vertebrae. In some kinds of baboons there are twenty-
five, whilst in the mandrill there are ten verv small
stunted caudal vertebrae, or, according to Cuvier,79 some-
times only five. This great diversity in the structure and
length of the tail in animals belonging to the same genera,
and following nearly the same habits of life, renders it
probable that the tail is not of much importance to
them ; and if so, we might have expected that it would
sometimes have become more or less rudimentary, in
accordance with what we incessantly see with other struc-
tures. The tail almost always tapers towards the end
whether it be long or short; and this, I presume, re-
sults from the atrophy, through disuse, of the terminal
muscles together with their arteries and nerves, lead-
ing to the atrophy of the terminal bones. With respect

79 Mr. St. George Mivart, ' Proc. Zoolog. Soc' 1865, p. 562, 583.
Dr. J. E. Gray, 'Cat. Brit. Mus. : Skeletons.' Owen, 'Anatomy of
Vertebrates,' vol. ii. p. 517. Isidore Geoffrov, 'Hist. Nat. Ge'n.' tom.
li p. 244.

Chap. IV. MANNER OF DEVELOPMENT. 151

to the os coccyx, which in man and the higher apes
manifestly consists of the few basal and tapering seg-
ments of an ordinary tail, I have heard it asked how
could these have become completely embedded within
the body ; but there is no difficulty in this respect,
for in many monkeys the basal segments of the true
tail are thus embedded. For instance, Mr. Murie in-
forms me that in the skeleton of a not fall-grown
Macacus inomatus, he counted nine or ten caudal ver-
tebrae, which altogether were only 1*8 inch in length.
Of these the three basal ones appeared to have been
embedded ; the remainder forming the free part of the
tail, which was only one inch in length, and half an
inch in diameter. Here, then, the three embedded
caudal vertebras plainly correspond with the /our coal-
esced vertebras of the human os coccyx.

I have now endeavoured to shew that some of the
most distinctive characters of man have in all proba-
bility been acquired, either directly, or more commonly
indirectly, through natural selection. We should bear
in mind that modifications in structure or constitution,
which are of no service to an organism in adapt-
ing it to its habits of life, to the food which it con-
sumes, or passively to the surrounding conditions, can-
not have been thus acquired. We must not, however,
be too confident in deciding what modifications are of
service to each being : we should remember how little
we know about the use of many parts, or what changes
in the blood or tissues may serve to jit an organism for
a new climate or some new kind of food. Nor must
we forget the principle of correlation, by which, as
Isidore Geoffroy has shewn in the case of man, many
strange deviations of structure are tied together. Inde-
pendently of correlation, a change in one part often leads

152 THE DESCENT OF MAN. Part I.

through the increased or decreased use of other parts,
to other changes of a quite unexpected nature. It is
also well to reflect on such facts, as the wonderful
growth of galls on plants caused by the poison of an
insect, and on the remarkable changes of colour in the
plumage of parrots when fed on certain fishes, or in-
oculated with the poison of toads ; 80 for we can thus
see that the fluids of the system, if altered for some
special purpose, might induce other strange changes.
We should especially bear in mind that modifications
acquired and continually used during past ages for
some useful purpose would probably become firmly
fixed and might be long inherited.

Thus a very large yet undefined extension may safely
be given to the direct and indirect results of natural
selection ; but I now admit, after reading the essay by
JSTageli on plants, and the remarks by various authors
with respect to animals, more especially those recently
made by Professor Broca, that in the earlier editions of
my * Origin of Species ' I probably attributed too much
to the action of natural selection or the survival of the
fittest. I have altered the fifth edition of the Origin
so as to confine my remarks to adaptive changes of
structure. I had not formerly sufficiently considered
the existence of many structures which appear to be,
as far as we can judge, neither beneficial nor injurious ;
and this I believe to be one of the greatest oversights as
yet detected in my work. I may be permitted to say
as some excuse, that I had two distinct objects in view,
firstly, to shew that species had not been separately
created, and secondly, that natural selection had been
the chief agent of change, though largely aided by the

80 ' The Variation of Animals and Plants under Domestication.' vol.
ii. p. 280, 282.

Chap. IV. MANNER OF DEVELOPMENT. 153

inherited effects of habit, and slightly by the direct
action of the surrounding conditions. Nevertheless I
was not able to annul • the influence of my former
belief, then widely prevalent, that each species had
been purposely created ; and this led to my tacitly
assuming that every detail of structure, excepting rudi-
ments, was of some special, though unrecognised, ser-
vice. Any one with this assumption in his mind would
naturally extend the action of natural selection, either
during past or present times, too far. Some of those
who admit the principle of evolution, but reject natural
selection, seem to forget, when criticising my book, that
I had the above two objects in view ; hence if I have
erred in giving to natural selection great power, which
I am far from admitting, or in having exaggerated its
power, which is in itself probable, I have at least, as I
hope, done good service in aiding to overthrow the
dogma of separate creations.

That all organic beings, including man, present many
modifications of structure which are of no service to
them at present, nor have been formerly, is, as I can
now see, probable. We know not what produces the
numberless slight differences between the individuals of
each species, for reversion only carries the problem a few
steps backwards ; but each peculiarity must have had
its own efficient cause. If these causes, whatever they
may be, were to act more uniformly and energetically
during a lengthened period (and no reason can be
assigned why this should not sometimes occur), the
result would probably be not mere slight individual
differences, but well-marked, constant modifications.
Modifications which are in no way beneficial cannot
have been kept unifornf through natural selection,
though any which were injurious would have been thus
eliminated. Uniformity of character would, however,

154 THE DESCENT OF MAN. Part I.

naturally follow from the assumed uniformity of the
exciting causes, and likewise from the free intercros-
sing of many individuals. The same organism might
acquire in this manner during successive periods suc-
cessive modifications, and these would be transmitted
in a nearly uniform state as long as the exciting causes
remained the same and there was free intercrossing.
With respect to the exciting causes we can only say, as
when speaking of so-called spontaneous variations, that
they relate much more closely to the constitution of the
varying organism, than to the nature of the conditions
to which it has been subjected.

Conclusion. — In this chapter we have seen that as man
at the present day is liable, like every other animal, to
multiform individual differences or slight variations, so
no doubt were the early progenitors of man ; the varia-
tions being then as now induced by the same general
causes, and governed by the same general and complex
laws. As all animals tend to multiply beyond their
means of subsistence, so it must have been with the
progenitors of man ; and this will inevitably have led
to a struggle for existence and to natural selection.
This latter process will have been greatly aided by
the inherited effects of the increased use of parts;
these two processes incessantly reacting on each other.
It appears, also, as we shall hereafter see, that various
unimportant characters have been acquired by man
through sexual selection. An unexplained residuum
of change, perhaps a large one, must be left to the
assumed uniform action of those unknown agencies,
which occasionally induce strongly-marked and abrupt
deviations of structure in our domestic productions.

Judging from the habits of savages and of the greater
number of the Quadrumana, primeval men, and even

Chap. IV. MANNER OF DEVELOPMENT. 155

the ape-like progenitors of man, probably lived in
society. With strictly social animals, natural selection
sometimes acts indirectly on the individual, through
the preservation of variations which are beneficial only
to the community. A community including a large
number of well-endowed individuals increases in number
and is victorious over other and less well-endowed com-
munities ; although each separate member may gain no
advantage over the other members of tire same com-
munity. With associated insects many remarkable
structures, which are of little or no service to the indi-
vidual or its own offspring, such as the pollen-collecting
apparatus, or the sting of the worker-bee, or the
great jaws of soldier- ants, have been thus acquired.
With the higher social animals, I am not aware that
any structure has been modified solely for the good of
the community, though some are of secondary service
to it. For instance, the horns of ruminants and the
great canine teeth of baboons appear to have been
acquired by the males as weapons for sexual strife, but
they are used in defence of the herd or troop. In
resard to certain mental faculties the case, as we shall
see in the following chapter, is wholly different ; for
these faculties have been chiefly, or even exclusively,
gained for the benefit of the community; the indi-
viduals composing the community being at the same
time indirectly benefited.

It has often been objected to such views as the fore-
going, that man is one of the most helpless and defence-
less creatures in the world ; and that during his early
and less well-developed condition he would have been
still more helpless. The Duke of Argyll, for instance,
insists81 that " the human frame lias diverged from

81 ' Primeval Man,' 1869, p. 60.

156 THE DESCENT OF MAN. Part I.

" the structure of brutes, in the direction of greater
<( physical helplessness and weakness. That is to say,
" it is a divergence which of all others it is most
" impossible to ascribe to mere natural selection." He
adduces the naked and unprotected state of the body,
the absence of great teeth or claws for defence, the
little strength of man, his small speed in running, and
his slight power of smell, by which to discover food or
to avoid danger. To these deficiencies there might
have been added the still more serious loss of the power
of quickly climbing trees, so as to escape from enemies.
Seeing that the unclothed Fuegians can exist under
their wretched, climate, the loss of hair would not
have been a great injury to primeval man, if he inha-
bited a warm country. When we compare defenceless
man with the apes, many of which are provided with
formidable canine teeth, wre must remember that these
in their fully-developed condition are possessed by the
males alone, being chiefly used by them for fighting
with their rivals ; yet the females which are not thus
provided, are able to survive.

In regard to bodily size or strength, we do not know
whether man is descended from some comparatively
small species, like the chimpanzee, or from one as
powerful as the gorilla ; and, therefore, we cannot say
whether man has become larger and stronger, or smaller
and weaker, in comparison with his progenitors. We
should, however, bear in mind that an animal possessing
great size, strength, and ferocity, and which, like the
gorilla, could defend itself from all enemies, would
probably, though not necessarily, have failed to become
social ; and this would most effectually have checked
the acquirement by man of his higher mental quali-
ties, such as sympathy and the love of his fellow-
creatures. Hence it might have been an immense

Chap. IV. MANNER OF DEVELOPMENT. 157

advantage to man to have sprung from some com-
paratively weak creature.

The slight corporeal strength of man, his little speed,
his want of natural weapons, &c, are more than coun-
terbalanced, firstly by his intellectual powers, through
which he has, whilst still remaining in a barbarous state,
formed for himself weapons, tools, &c, and secondly by
his social qualities which lead him to give aid to his
fellow-men and to receive it in return. No country
in the world abounds in a greater degree with dan-
gerous beasts than Southern Africa; no country pre-
sents more fearful physical hardships than the Arctic
regions ; yet one of the puniest races, namely, the
Bushmen, maintain themselves in Southern Africa, as
do the dwarfed Esquimaux in the Arctic regions. The
early progenitors of man were, no doubt, inferior in
intellect, and probably in social disposition, to the
lowest existing savages ; but it is quite conceivable that
they might have existed, or even flourished, if, whilst
they gradually lost their brute-like powers, such as
climbing trees, &c, they at the same time advanced
in intellect. But granting that the progenitors of man
were far more helpless and defenceless than any existing
savages, if they had inhabited some warm continent
or large island, such as Australia or New Guinea, or
Borneo (the latter island being now tenanted by the
orang), they would not have been exposed to any special
danger. In an area as large as one of these islands,
the competition between tribe and tribe would have
been sufficient, under favourable conditions, to have
raised man, through the survival of the fittest, combined
with the inherited effects of habit, to his present high
position in the organic scale.

158 THE DESCENT OF MAN. Part I.

CHAPTEK V.

On the Development of the Intellectual and Moral
Faculties during Primeval and Civilised Times.

The advancement of the intellectual powers through natural selec-
tion— 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.

The subjects to be discussed in this chapter are of
the highest interest, but are treated by me in a most
imperfect and fragmentary manner. Mr. Wallace,
in an admirable paper before referred to,1 argues that
man after he had partially acquired those intellectual
and moral faculties which distinguish him from the
lower animals, would have been but little liable to
have had his bodily structure modified through natural
selection or any other means. For man is enabled
through his mental faculties "to keep with an un-
" changed body in harmony with the changing universe."
He has great power of adapting his habits to new
conditions of life. He invents weapons, tools and
various stratagems, by which he procures food and
defends himself. When he migrates into a colder
climate he uses clothes, builds sheds, and makes fires ;
and, by the aid of fire, cooks food otherwise indigestible.
He aids his fellow-men in many ways, and anticipates
future events. Even at a remote period he practised
some subdivision of labour.

i <

Anthropological Review,' May, 18G4, p. clviii.

Chap. V. INTELLECTUAL FACULTIES. 159

The lower animals, on the other hand, must have
their bodily structure modified in order to survive under
greatly changed conditions. They must be rendered
stronger, or acquire more effective teeth or claws, in
order to defend themselves from new enemies ; or they
must be reduced in size so as to escape detection and
danger. When they migrate into a colder climate they
must become clothed with thicker fur, or have their
constitutions altered. If they fail to be thus modified,
they will cease to exist.

The case, however, is widely different, as Mr. Wal-
lace has with justice insisted, in relation to the intel-
lectual and moral faculties of man. These faculties are
variable ; and we have every reason to believe that the
variations tend to be inherited. Therefore, if they were
formerly of high importance to primeval man and to
his ape-like progenitors, they would have been per-
fected or advanced through natural selection. Of the
high importance of the intellectual faculties there can
be no doubt, for man mainly owes to them his pre-
eminent position in the world. We can see that, in
the rudest state of society, the individuals who were the
most sagacious, who invented and used the best weapons
or traps, and who were best able to defend themselves,
would rear the greatest number of offspring. The tribes
which included the largest number of men thus endowed
would increase in number and supplant other tribes.
Numbers depend primarily on the means of subsistence,
and this, partly on the physical nature of the country,
but in a much higher degree on the arts which are there
practised. As a tribe increases and is victorious, it is
often still further increased by the absorption of other
tribes.2 The stature and strength of the men of a tribe

2 After a time the members or tribes which are absorbed into another
tribe assume, as Mr. Maine remarks (' Ancient Law,' 1SG1, p. 131), that
they are the co-descendants of the same ancestors.

160 THE DESCENT OF MAN. Part I.

are likewise of some importance for its success, and
these depend in part on the nature and amount of the
food which can be obtained. In Europe the men of the
Bronze period were supplanted by a more powerful and,
judging from their sword- handles, larger-handed race ;3
but their success was probably due in a much higher
degree to their superiority in the arts.

All that we know about savages, or may infer from
their traditions and from old monuments, the history
of which is quite forgotten by the present inhabitants,
shew that from the remotest times successful tribes have
supplanted other tribes. Kelics of extinct or forgotten
tribes have been discovered throughout the civilised
regions of the earth, on the wild plains of America, and
on the isolated islands in the Pacific Ocean. At the
present day civilised nations are everywhere supplanting
barbarous nations, excepting where the climate opposes
a deadly barrier ; and they succeed mainly, though not
exclusively, through their arts, which are the products
of the intellect. It is, therefore, highly probable that
with mankind the intellectual faculties have been
gradually perfected through natural selection ; and this
conclusion is sufficient for our purpose. Undoubtedly
it would have been very interesting to have traced the
development of each separate faculty from the state in
which it exists in the lower animals to that in which it
exists in man ; but neither my ability nor knowledge
permit the attempt.

It deserves notice that as soon as the progenitors of
man became social (and this probably occurred at a
very early period), the advancement of the intellectual
faculties will have been aided and modified in an
important manner, of which we see only traces in

3 Morlot, ' Soc. Vaud. Sc Nat.1 i860, p. 294.

Chap. V. INTELLECTUAL FACULTIES. 161

the lower animals, namely, through the principle
of imitation, together with reason and experience.
Apes are much given to imitation, as are the lowest
savages ; and the simple fact previously referred to.
that after a time no animal can be caught in the same
place by the same sort of trap, shews that animals learn
by experience, and imitate each others' caution. Now,
if some one man in a tribe, more sagacious than the
others, invented a new snare or weapon, or other means
of attack or defence, the plainest self-interest, without
the assistance of much reasoning power, would prompt
the other members to imitate him ; and all would thus
profit. The habitual practice of each new art must
likewise in some slight degree strengthen the intellect.
If the new invention were an important one, the tribe
would increase in number, spread, and supplant other
tribes. In a tribe thus rendered more numerous there
would always be a rather better chance of the birth of
other superior and inventive members. If such men
left children to inherit their mental superiority, the
chance of the birth of still more ingenious members
would be somewhat better, and in a very small tribe
decidedly better. Even if they left no children, the
tribe would still include their blood-relations ; and it has
been ascertained by agriculturists4 that by preserving
and breeding from the family of an animal, which
when slaughtered was found to be valuable, the desired
character has been obtained.

Turning now to the social and moral faculties. In
order that primeval men, or the ape-like progenitors
of man, should have become social, thev must have

4 I have given instances in my ' Variation of Animals under Domes-
tication/ vol. ii. p. 196.

VOL. I. M

162 THE DESCENT OF MAN. Part I.

acquired the same instinctive feelings which impel other
animals to live in a body ; and they no doubt exhi-
bited the same general disposition. They would have
felt uneasy when separated from their comrades, for
whom they would have felt some degree of love ; they
would have warned each other of danger, and have
given mutual aid in attack or defence. All this implies
some degree of sympathy, fidelity, and courage. Such
social qualities, the paramount importance of which
to the lower animals is disputed by no one, were no
doubt acquired by the progenitors of man in a similar
manner, namely, through natural selection, aided by
inherited habit. When two tribes of primeval man,
living in the same country, came into competition,
if the one tribe included (other circumstances being
equal) a greater number of courageous, sympathetic,
and faithful members, who were always ready to warn
each other of danger, to aid and defend each other, this
tribe would without doubt succeed best and conquer the
other. Let it be borne in mind how all-important, in
the never-ceasing wars of savages, fidelity and courage
must be. The advantage which disciplined soldiers
have over undisciplined hordes follows chiefly from the
confidence which each man feels in his comrades.
Obedience, as Mr. Bagehot has well shewn,5 is of the
highest value, for any form of government is better
than none. Selfish and contentious people will not
cohere, and without coherence nothing can be effected.
A tribe possessing the above qualities in a high de-
gree would spread and be victorious over other tribes ;
but in the course of time it would, judging from all
past history, be in its turn overcome by some other

5 See a remarkable series of articles on Physics and Politics in the
Fortnightly Review,' Nov. 1SU7 ; April 1, 1SGS; July 1, 1869.

Chap. Y. MOKAL FACULTIES. 16

30

o

and still more highly endowed tribe. Thus the social
and moral qualities would tend slowly to advance and
be diffused throughout the world.

But it may be asked, how within the limits of the
same tribe did a large number of members first become
endowed with these social and moral qualities, and
how was the standard of excellence raised? It is
extremely doubtful whether the offspring of the more
sympathetic and benevolent parents, or of those which
were the most faithful to their comrades, would be
reared in greater number than the children of selfish
and treacherous parents of the same tribe. He who
was ready to sacrifice his life, as many a savage has
been, rather than betray his comrades, would often leave
no offspring to inherit his noble nature. The bravest
men, who were always willing to come to the front in
war, and who freely risked their lives for others, would
on an average perish in larger number than other men.
Therefore it seems scarcely possible (bearing in mind
that we are not here speaking of one tribe being vic-
torious over another) that the number of men gifted
with such virtues, or that the standard of their excel-
lence, could be increased through natural selection, that
is, by the survival of the fittest.

Although the circumstances which lead to an increase
in the number of men thus endowed within the same
tribe are too complex to be clearly followed out, we can
trace some of the probable steps. In the first place, as
the reasoning powers and foresight of the members
became improved, each man would soon learn from
experience that if he aided his fellow-men, he would
commonly receive aid in return. From this low motive
he might acquire the habit of aiding his fellows ; and
the habit of performing benevolent actions certainly
strengthens the feeling of sympathy, which gives the

M 2

164 THE DESCENT OF MAN. Part I.

first impulse to benevolent actions. Habits, moreover,
followed during many generations probably tend to be
inherited.

But there is another and much more powerful sti-
mulus to the development of the social virtues, namely,
the praise and the blame of our fellow-men. The love
of approbation and the dread of infamy, as well as the
bestowal of praise or blame, are primarily clue, as Ave
have seen in the third chapter, to the instinct of sym-
pathy; and this instinct no doubt was originally acquired,
like all the other social instincts, through natural selec-
tion. At how early a period the progenitors of man, in
the course of their development, became capable of feel-
ing and being impelled by the praise or blame of their
fellow-creatures, we cannot, of course, say. But it appears
that even dogs appreciate encouragement, praise, and
blame. The rudest savages feel the sentiment of glory,
as they clearly show by preserving the trophies of their
prowess, by their habit of excessive boasting, and even
by the extreme care which they take of their personal
appearance and decorations ; for unless they regarded
the opinion of their comrades, such habits would be
senseless.

Thev certainlv feel shame at the breach of some of
their lesser rules ; but how far they experience remorse
is doubtful. I was at first surprised that I could not re-
collect any recorded instances of this feeling in savages ;
and Sir J. Lubbock 6 states that he knows of none.
But if we banish from our minds all cases given in
novels and plays and in death-bed confessions made
to priests, I doubt whether many of us have actually
witnessed remorse; though we may have often seen
shame and contrition for smaller offences. Remorse is

' Origin of Civilisation,' 1870, p. 265.

Chai\ V. MOKAL FACULTIES. 165

a deeply hidden feeling. It is incredible that a savage,
who will sacrifice his life rather than betray his tribe,
or one who will deliver himself up as a prisoner rather
than break his parole,7 would not feel remorse in his
inmost soul, though he might conceal it, if he had failed
in a duty which he held sacred.

We may therefore conclude that primeval man, at a
very remote period, would have been influenced by the
praise and blame of his fellows. It is obvious, that the
members of the same tribe wrould approve of conduct
which appeared to them to be for the general good, and
would reprobate that which appeared evil. To do good
unto others — to do unto others as ye would they should
do unto you, — is the foundation-stone of morality. It
is, therefore, hardly possible to exaggerate the impor-
tance during rude times of the love of praise and the
dread of blame. A man who was not impelled by any
deep, instinctive feeling, to sacrifice his life for the good
of others, yet was roused to such actions by a sense
of glory, would by his example excite the same wish
for glory in other men, and would strengthen by exer-
cise the noble feeling of admiration. He might thus
do far more good to his tribe than by begetting offspring
with a tendency to inherit his own high character.

With increased experience and reason, man perceives
the more remote consequences of his actions, and the
self-regarding virtues, such as temperance, chastity, &c,
which during early times are, as we have before seen,
utterly disregarded, come to be highly esteemed or even
held sacred. I need not, however, repeat what I have
said on this head in the third chapter. Ultimately a
highly complex sentiment, having its first origin in the

7 Mr. Wallace gives cases in his ' Contributions to the Theory of
Natural Selection/ 1870, p. 354.

166 THE DESCENT OF MAN. Part I.

social instincts, largely guided by the approbation of
our fellow-men, ruled by reason, self-interest, and in
later times by deep religious feelings, confirmed by
instruction and habit, all combined, constitute our moral
sense or conscience.

It must not be forgotten that although a high stand-
ard of morality gives but a slight or no advantage to
each individual man and his children over the other
men of the same tribe, yet that an advancement in the
standard of morality and an increase in the number
of well-endowed men will certainly give an immense
advantage to one tribe over another. There can be no
doubt that a tribe including many members who, from
possessing in a high degree the spirit of patriotism,
fidelity, obedience, courage, and sympathy, were always
ready to give aid to each other and to sacrifice them-
selves for the common good, would be victorious over
most other tribes ; and this would be natural selection.
At all times throughout the world tribes have sup-
planted other tribes ; and as morality is one element
in their success, the standard of morality and the num-
ber of well-endowed men will thus everywhere tend to
rise and increase.

It is, however, very difficult to form any judgment why
one particular tribe and not another has been successful
and has risen in the scale of civilisation. Many savages
are in the same condition as when first discovered several
centuries ago. As Mr. Bagehot has remarked, we are
apt to look at progress as the normal rule in human
society ; but history refutes this. The ancients did not
even entertain the idea ; nor do the oriental nations at
the present day. According to another high authority,
Mr. Maine,8 "the greatest part of mankind has never

8 'Ancient Law,' 1861, p. 22. For Mr. Bagehot's remarks, 'Fort-
nightly Review,' April 1, 1868, p. 452.

Chap. V. CIVILISED NATIONS. 167

" shewn a particle of desire that its civil institutions
" should be improved." Progress seems to depend on
many concurrent favourable conditions, far too complex
to be followed out. But it has often been remarked, that
a cool climate from leading to industry and the various
arts has been highly favourable, or even indispensable
for this end. The Esquimaux, pressed by hard necessity,
have succeeded in many ingenious inventions, but their
climate has been too severe for continued progress.
Nomadic habits, whether over wide plains, or through
the dense forests of the tropics, or along the shores of
the sea, have in every case been highly detrimental.
Whilst observing the barbarous inhabitants of Tierra
del Fuego, it struck me that the possession of some
property, a fixed abode, and the union of many families
under a chief, were the indispensable requisites for
civilisation. Such habits almost necessitate the culti-
vation of the ground ; and the first steps in cultivation
would probably result, as I have elsewhere shewn,9 from
some such accident as the seeds of a fruit-tree falling
on a heap of refuse and producing an unusually fine
variety. The problem, however, of the first advance of
savages towards civilisation is at present much too diffi-
cult to be solved.

Natural Selection as affecting Civilised Nations. — In
the last and present chapters I have considered the
advancement of man from a former semi-human con-
dition to his present state as a barbarian. But some
remarks on the agencv of natural selection on civilised
nations may be here worth adding. This subject has
been ably discussed by Mr. W. R. Greg,10 and previously

9 ' The Variation of Animals and Plants under Domestication,' vol. i.
p. 309.

10 ' Fraser's Magazine,' Sept. 1868, p. 353. This article seems to
have struck many persons, and has given rise to two remarkable essays

168 THE DESCENT OF MAN. Part I.

by Mr. Wallace and Mr. Galton.11 Most of my remarks
are taken from these three authors. With savages, the
weak in body or mind are soon eliminated ; and those
that survive commonly exhibit a vigorous state of
health. We civilised men, on the other hand, do our
utmost to check the process of elimination ; we build
asylums for the imbecile, the maimed, and the sick ; we
institute poor-laws; and our medical men exert their
utmost skill to save the life of every one to the last
moment. There is reason to believe that vaccination
has preserved thousands, who from a weak constitution
would formerly have succumbed to small-pox. Thus
the weak members of civilised societies propagate their
kind. No one who has attended to the breeding of
domestic animals will doubt that this must be highly
injurious to the race of man. It is surprising how soon
a want of care, or care wrongly directed, leads to the
degeneration of a domestic race ; but excepting in the
case of man himself, hardly any one is so ignorant as
to allow his worst animals to breed.

The aid which we feel impelled to give to the help-
less is mainly an incidental result of the instinct of
sympathy, which was originally acquired as part of
the social instincts, but subsequently rendered, in the
manner previously indicated, more tender and more
widely diffused. Nor could we check our sympathy, if
so urged by hard reason, without deterioration in the

and a rejoinder in the ' Spectator,' Oct. 3rd and I7th 1868. It has
also been discussed in the ' Q. Journal of Science,' 1869, p. 152, and by
Mr. Lawson Tait in the 'Dublin Q. Journal of Medical Science,' Feb.
1869, and by Mr. E. Kay Lankester in his ' Comparative Longevity/
1S70, p. 128. Similar views appeared previously in the 'Australasian,'
July 13, 1867. I have borrowed ideas from several of these writers.

11 For Mr. Wallace, see ' Antbropolog. Review,' as before cited,
Mr. Galton in ' Macmillan's Magazine,' Aug. 1865, p. 318 ; also his
great work, ' Hereditary Genius,' 1870.

Chap. V. CIVILISED NATIONS. 169

noblest part of our nature. The surgeon may harden
himself whilst performing an operation, for he knows
that he is acting for the good of his patient ; but if we
were intentionally to neglect the weak and helpless, it
could only be for a contingent benefit, with a certain
and great present evil. Hence we must bear without
complaining the undoubtedly bad effects of the weak
surviving and propagating their kind ; but there appears
to be at least one check in steady action, namely the
weaker and inferior members of society not marrying
so freely as the sound ; and this check might be inde-
finitely increased, though this is more to be hoped for
than expected, by the weak in body or mind refraining
from marriage.

In all civilised countries man accumulates property
and bequeaths it to his children. So that the children
in the same country do not by any means start fair in
the race for success. But this is far from an unmixed
evil ; for without the accumulation of capital the arts
could not progress ; and it is chiefly through their power
that the civilised races have extended, and are now
everywhere extending, their range, so as to take the place
of the lower races. Nor does the moderate accumulation
of wealth interfere with the process of selection. When
a poor man becomes rich, his children enter trades or
professions in which there is struggle enough, so that
the able in body and mind succeed best. The presence
of a body of well-instructed men, who have not to
labour for their daily bread, is important to a degree
which cannot be over-estimated ; as all high intellectual
work is carried on by them, and on such work material
progress of all kinds mainly depends, not to mention
other and higher acl vantages. No doubt wealth when
very great tends to convert men into useless drones, bu^\\3^
their number is never large ; and some degree of elimi-

170 THE DESCENT OF MAN. Part I.

nation here occurs, as we daily see rich men, who happen
to be fools or profligate, squandering away all their
wealth.

Primogeniture with entailed estates is a more direct
evil, though it may formerly have been a great advan-
tage by the creation of a dominant class, and any
government is better than anarchy. The eldest sons,
though they may be weak in body or mind, generally
marry, whilst the younger sons, however superior in these
respects, do not so generally marry, Nor can worthless
eldest sons with entailed estates squander their wealth.
But here, as elsewhere, the relations of civilised life are
so complex that some compensatory checks intervene.
The men who are rich through primogeniture are able
to select generation after generation the more beautiful
and charming women ; and these must generally be
healthy in body and active in mind. The evil con-
sequences, such as they may be, of the continued pre-
servation of the same line of descent, without any
selection, are checked by men of rank always wishing to
increase their wealth and power ; and this they effect
by marrying heiresses. But the daughters of parents
who have produced single children, are themselves, as
Mr. Galton has shewn,12 apt to be sterile ; and thus
noble families are continually cut off in the direct line,
and their wealth flows into some side channel; but
unfortunately this channel is not determined by supe-
riority of any kind.

Although civilisation thus checks in many ways the
action of natural selection, it apparently favours, by
means of improved food and the freedom from occa-
sional hardships, the better development of the body.
This may be inferred from civilised men having been

12

'Hereditary Genius,' 1S70, p. 132-140.

Chap. V. CIVILISED NATIONS. 171

found, wherever compared, to be physically stronger than
savages. They appear also to have equal powers of
endurance, as has been proved in many adventurous
expeditions. Even the great luxury of the rich can be
but little detrimental ; for the expectation of life of our
aristocracy, at all ages and of both sexes, is very little
inferior to that of healthy English lives in the lower
classes.13

We will now look to the intellectual faculties alone.
If in each grade of society the members were divided
into two equal bodies, the one including the intel-
lectually superior and the other the inferior, there can
be little doubt that the former would succeed best in
all occupations and rear a greater number of children.
Even in the lowest walks of life, skill and ability must
be of some advantage, though in many occupations,
owing to the great division of labour, a very small
one. Hence in civilised nations there will be some
tendency to an increase both in the number and in
the standard of the intellectually able. But I do not
wish to assert that this tendency may not be more than
counterbalanced in other ways, as by the multiplication
of the reckless and improvident ; but even to such as
these, ability must be some advantage.

It has often been objected to views like the fore-
going, that the most eminent men who have ever lived
have left no offspring to inherit their great intellect.
Mr. Gal ton says,14 " I regret I am unable to solve the
" simple question whether, and how far, men and women
" who are prodigies of genius are infertile. I have, how-
" ever, shewn that men of eminence are by no means so."

13 See the fifth and sixth columns, compiled from good authorities,
in the table given in Mr. E. K. Lankester's ' Comparative Longevity,'
1870r p. 115.

14 'Hereditary Genius,' 1870, p. 330.

172 THE DESCENT OF MAN. Part I.

Great lawgivers, the founders of beneficent religions,
great philosophers and discoverers in science, aid the
progress of mankind in a far higher degree by their
works than by leaving a numerous progeny. In the
case of corporeal structures, it is the selection of
the slightly better-endowed and the elimination of the
slightly less well-endowed individuals, and not the pre-
servation of strongly-marked and rare anomalies, that
leads to the advancement of a species.15 So it will be
with the intellectual faculties, namely from the some-
what more able men in each grade of society succeeding
rather better than the less able, and consequently in-
creasing in number, if not otherwise prevented. When
in any nation the standard of intellect and the number
of intellectual men have increased, we may expect from
the law of the deviation from an average, as shewn by
Mr. Galton, that prodigies of genius will appear some-
what more frequently than before.

In regard to the moral qualities, some elimination of
the worst dispositions is always in progress even in the
most civilised nations. Malefactors are executed, or
imprisoned for long periods, so that they cannot freely
transmit their bad qualities. Melancholic and insane
persons are confined, or commit suicide. Violent and
quarrelsome men often come to a bloody end. Kestless
men who will not follow any steady occupation — and
this relic of barbarism is a great check to civilisation16 —
emigrate to newly- sett led countries, where they prove
useful pioneers. Intemperance is so highly destructive,
that the expectation of life of the intemperate, at the
age, for instance, of thirty, is only 13 -8 years ; whilst for
the rural labourers of England at the same a^e it is

15 ' Origin of Species' (fifth edition, 1869), p. 104.

16 'Hereditary Genius,' 1870, p. 347.

Chap. V. CIVILISED NATIONS. 173

4059 years.17 Profligate women bear few children, and
profligate men rarely marry; both suffer from disease.
In the breeding of domestic animals, the elimination of
those individuals, though few in number, which are in
any marked manner inferior, is by no means an unim-
portant element towards success. This especially holds
good with injurious characters which tend to reappear
through reversion, such as blackness in sheep ; and
with mankind some of the worst dispositions, which
occasionally without any assignable cause make their
appearance in families, may perhaps be reversions to
a savage state, from which we are not removed by very
many generations. This view seems indeed recognised
in the common expression that such men are the black
sheep of the family.

With civilised nations, as far as an advanced stand-
ard of morality, and an increased number of fairly
well-endowed men are concerned, natural selection ap-
parently effects but little ; though the fundamental
social instincts were originally thus gained. But I
have already said enough, whilst treating of the lower
races, on the causes which lead to the advance of
morality, namely, the approbation of our fellow-men —
the strengthening of our sympathies by habit — example
and imitation — reason — experience and even self-inte-
rest— instruction during youth, and religious feelings.

A most important obstacle in civilised countries to
an increase in the number of men of a superior class has
been strongly urged by Mr. Greg and Mr. Galton,18

17 E. Eay Lankester, 'Comparative Longevity,' 1870, p. 115. The
table of the intemperate is from Neison's ' Vital Statistics.' In regard
to profligacy, see Dr. Farr, " Influence of Marriage on Mortality," ' Nat,
Assoc, for the Promotion of Social Science,' 1858.

18 ' Fraser'.i Magazine,' Sept. 1868, p. 358. 'Macmillan's Magazine,'
Aug. 1805, p. 318. The Eev. F. W. Farrar (' Fraser's Mag.,' Aug. 1870,
p. 261) tahes a different view.

174 THE DESCENT OF MAN. Paet I.

namely, the fact that the very poor and reckless, who
are often degraded by vice, almost id variably marry
early, whilst the careful and frugal, who are generally
otherwise virtuous, marry late in life, so that they may
be able to support themselves and their children in
comfort. Those who marry early produce within a
given period not only a greater number of generations,
but, as shewn by Dr. Duncan,19 they produce many more
children. The children, moreover, that are born by
mothers during the prime of life are heavier and larger,
and therefore probably more vigorous, than those born
at other periods. Thus the reckless, degraded, and
often vicious members of society, tend to increase at a
quicker rate than the provident and generally virtuous
members. Or as Mr. Greg puts the case : " The care-
" less, squalid, unaspiring Irishman multiplies like
" rabbits : the frugal, foreseeing, self-respecting, am-
" bitious Scot, stern in his morality, spiritual in his
" faith, sagacious and disciplined in his intelligence,
" passes his best years in struggle and in celibacy,
" marries late, and leaves few behind him. Given a
" land originally peopled by a thousand Saxons and a
" thousand Celts — and in a dozen generations five-sixths
" of the population would be Celts, but five-sixths of
"the property, of the power, of the intellect, would
" belong to the one-sixth of Saxons that remained.
" In the eternal ' struggle for existence,' it would be
" the inferior and less favoured race that had prevailed
" — and prevailed by virtue not of its good qualities
" but of its faults."

There are, however, some checks to this downward
tendency. We have seen that the intemperate suffer

19 "On the Laws of the Fertility of Women," in * Transact. Eoyal
Soc.' Edinburgh, vol. xxiv. p. 287. See, also, Mr. Galton, ' Hereditary
Genius,' p. 352-357, for observations to the above effect.

Chap. V. CIVILISED NATIONS. 175

from a high rate of mortality, and the extremely pro-
fligate leave few offspring. The poorest classes crowd
into towns, and it has been proved by Dr. Stark from
the statistics of ten years in Scotland,20 that at all ages
the death-rate is higher in towns than in rural districts,
" and during the first five years of life the town cleath-
" rate is almost exactly double that of the rural districts."
As these returns include both the rich and the poor, no
doubt more than double the number of births would be
requisite to keep up the number of the very poor inha-
bitants in the towns, relatively to those in the country.
With women, marriage at too early an age is highly
injurious ; for it has been found in France that, " twice
" as many wives under twenty die in the year, as died out
" of the same number of the unmarried." The mortality,
also, of husbands under twenty is " excessively high,"21
but what the cause of this may be seems doubtful.
Lastly, if the men who prudently delay marrying until
they can bring up their families in comfort, were to
select, as they often do, women in the prime of life, the
rate of increase in the better class would be only slightly
lessened.

It was established from an enormous body of statistics,
taken during 1853, that the unmarried men throughout
France, between the ages of twenty and eighty, die in a
much larger proportion than the married : for instance,
out of every 1000 unmarried men, between the ages of
twenty and thirty, 11*3 annually died, whilst of the
married only 6*5 died.22 A similar law was proved to

20 ' Tenth Annual Report of Births, Deaths, &c., in Scotland,' 1867,
p. xxix.

21 These quotations are taken from our highest authority on such
questions, namely, Dr. Farr, in his paper " On the Influence of Marriage
on the Mortality of the French People," read before the Nat. Assoc.
for the Promotion of Social Science, 1858.

22 Dr. Farr, ibid. The quotations given below are extracted from
the same striking paper.

176 THE DESCENT OF MAN. Part I.

hold good, during the years 1863 and 1864, with the
entire population above the age of twenty in Scotland :
for instance, out of every 1000 unmarried men, between
the ages of twenty and thirty, 14*97 annually died,
whilst of the married only 7*24 died, that is less than
half.23 Dr. Stark remarks on this, " Bachelorhood is
" more destructive to life than the most unwholesome
u trades, or than residence in an unwholesome house or
" district where there has never been the most distant
" attempt at sanitary improvement." He considers that
the lessened mortality is the direct result of " marriage,
" and the more regular domestic habits which attend that
" state." He admits, however, that the intemperate,
profligate, and criminal classes, whose duration of life
is low, do not commonly marry ; and it must like-
wise be admitted that men with a weak constitution,
ill health, or any great infirmity in body or mind, will
often not wish to marry, or will be rejected. Dr. Stark
seems to have come to the conclusion that marriage in
itself is a main cause of prolonged life, from finding
that ao'ed married men still have a considerable advan-
tage in this respect over the unmarried of the same
advanced age ; but every one must have known instances
of men, who with weak health during youth did not
marry, and yet have survived to old age, though
remaining weak and therefore always with a lessened
chance of life. There is another remarkable circum-
stance which seems to support Dr. Stark's conclusion,
namely, that widows and widowers in France suffer in
comparison with the married a very heavy rate of mor-
tality; but Dr. Fan* attributes this to the poverty and

-z I have taken the mean of the quinquennial means, given in 'The
Tenth Annual Keport of Births, Deaths, &c, in Scotland,' 1867. The
quotation from Dr. Stark is copied from an article in the * Daily News,'
Oct. 17th, 1S68, which Dr. Farr considers very carefully written.

Chap. V. CIVILISED NATIONS. 177

evil habits consequent on the disruption of the family,
and to grief. On the whole we may conclude with Dr.
Farr that the lesser mortality of married than of unmar-
ried men, which seems to be a general law, " is mainly
" due to the constant elimination of imperfect types, and
" to the skilful selection of the finest individuals out of
" each successive generation;" the selection relating only
to the marriage state, and acting on all corporeal, in-
tellectual, and moral qualities. We may, therefore,
infer that sound and good men who out of prudence
remain for a time unmarried do not sutler a high rate
of mortal it v.

If the various checks specified in the two last para-
graphs, and perhaps others as yet unknown, do not
prevent the reckless, the vicious and otherwise inferior
members of society from increasing at a quicker rate
than the better class of men, the nation will retro-
grade, as has occurred too often in the history of the
world. We must remember that progress is no invari-
able rule. It is most difficult to say why one civilised
nation rises, becomes more powerful, and spreads more
widely, than another ; or why the same nation progresses
more at one time than at another. We can only say
that it depends on an increase in the actual number of
the population, on the number of the men endowed
with high intellectual and moral faculties, as well as
on their standard of excellence. Corporeal structure,
except so far as vigour of body leads to vigour of mind,
appears to have little influence.

It has been urged by several writers that as high
intellectual powers are advantageous to a nation, the
old Greeks, who stood some grades higher in intellect
than any race that has ever existed,24 ought to have

24 See the ingenious and original argument on this subject by Mr.
Galton, 'Hereditary Genius,' p. 310-342.

VOL. I. N

178 THE DESCENT OF MAN. Part I.

risen, if the power of natural selection were real, still
higher in the scale, increased in number, and stocked
the whole of Europe. Here we have the tacit assump-
tion, so often made with respect to corporeal structures,
that there is some innate tendency towards continued
development in mind and body. But development of
all kinds depends on many concurrent favourable cir-
cumstances. Natural selection acts only in a tentative
manner. Individuals and races may have acquired
certain indisputable advantages, and yet have perished
from failing in other characters. The Greeks may have
retrograded from a want of coherence between the many
small states, from the small size of their whole country,
from the practice of slavery, or from extreme sensuality ;
for they did not succumb until "they were enervated
" and corrupt to the very core."25 The western nations
of Europe, who now so immeasurably surpass their
former savage progenitors and stand at the summit of
civilisation, owe little or none of their superiority to
direct inheritance from the old Greeks; though they
owe much to the written works of this wonderful people.
Who can positively say why the Spanish nation,
so dominant at one time, has been distanced in the
race. The awakening of the nations of Europe from
the dark ages is a still more perplexing problem. At
this early period, as Mr. Galton 26 has remarked, almost
all the men of a gentle nature, those given to medi-
tation or culture of the mind, had no refuge except in
the bosom of the Church which demanded celibacy ;

25 Mr. Greg, ' Fraser's Magazine,' Sept. 1868, p. 357.

26 'Hereditary Genius,' 1870, p. 357-359. The Eev. F. H. Farrar
(' Fraser's Mag.', Aug. 1870, p. 257) advances arguments on the other
side. Sir C. Lyell had already (* Principles of Geology,' vol. ii. 1868,
p. 489) called attention, in a striking passage, to the evil influence of
the Holy Inquisition in having lowered, through selection, the general
standard of intelligence in Europe.

Chap. V. CIVILISED NATIONS. 179

and this could hardly fail to have had a deteriorating
influence on each successive generation. During this
same period the Holy Inquisition selected with extreme
care the freest and boldest men in order to burn or
imprison them. In Spain alone some of the best men —
those who doubted and questioned, and without doubting
there can be no progress — were eliminated during three
centuries at the rate of a thousand a year. The evil
which the Catholic Church has thus effected, though
no doubt counterbalanced to a certain, perhaps large
extent in other ways, is incalculable ; nevertheless,
Europe has progressed at an unparalleled rate.

The remarkable success of the English as colonists
over other European nations, which is well illustrated by
comparing the progress of the Canadians of English and
French extraction, has been ascribed to their " daring
" and persistent energy ; " but who can say how the
English gained their energy. There is apparently
much truth in the belief that the wonderful progress
of the United States, as well as the character of the
people, are the results of natural selection ; the more
energetic, restless, and courageous men from all parts
of Europe having emigrated during the last ten or
twelve generations to that great country, and having there
succeeded best.27 Looking to the distant future, I do
not think that the Rev. Mr. Zincke takes an exaggerated
view when he says :28 " All other series of events — as
" that which resulted in the culture of mind in Greece,
" and that which resulted in the empire of Rome — only
" appear to have purpose and value when viewed in
" connection with, or rather as subsidiarv to ... . the
" great stream of Anglo-Saxon emigration to the west."

27 Mr. Gal ton, • Macmillan's Magazine,' August, 1865, p. 325. See,
also, ' Nature,' " On Darwinism and National Life," Dec. 1869, p. 184.

28 ' Last Winter in the United States,' 1868, p. 29.

N 2

180 THE DESCENT OF MAN. Part I.

Obscure as is the problem of the advance of civilisation,
we can at least see that a nation which produced during
a lengthened period the greatest number of highly intel-
lectual, energetic, brave, patriotic, and benevolent men,
would generally prevail over less favoured nations.

Natural selection follows from the struggle for exist-
ence ; and this from a rapid rate of increase. It is
impossible not bitterly to regret, but whether wisely
is another question, the rate at which man tends to
increase ; for this leads in barbarous tribes to infan-
ticide and many other evils, and in civilised nations to
abject poverty, celibacy, and to the late marriages of
the prudent. But as man suffers from the same physical
evils with the lower animals, he has no right to expect
an immunity from the evils consequent on the struggle
for existence. Had he not been subjected to natural
selection, assuredlv he would never have attained to
the rank of manhood. When we see in many parts
of the world enormous areas of the most fertile land
peopled by a few wandering savages, but which are
capable of supporting numerous happy homes, it might
be argued that the struggle for existence had not been
sufficiently severe to force man upwards to his highest
standard. Judoino- from all that we know of man and
the lower animals, there has always been sufficient
variability in the intellectual and moral faculties, for
their steady advancement through natural selection.
No doubt such advancement demands many favourable
concurrent circumstances ; but it may well be doubted
whether the most favourable would have sufficed, had
not the rate of increase been rapid, and the consequent
struggle for existence severe to an extreme degree.

On tlie evidence that all civilised nations were once bar-
barous,— As we have had to consider the steps by which

Ciiap. V. CIVILISED NATIONS. 181

some semi-bum an creature has been gradually raised to
the rank of man in his most perfect state, the present
subject cannot be quite passed over. But it has been
treated in so full and admirable a manner by Sir J.
Lubbock,39 Mr. Tylor, Mr. M'Lennan, and others, that
1 need here give only the briefest summary of their
results. The arguments recently advanced by the
Duke of Argyll ^ and formerly by Archbishop Whately,
in favour of the belief that man came into the world
as a civilised being and that all savages have since
undergone degradation, seem to me weak in comparison
with those advanced on the other side. Many nations,
no doubt, have fallen away in civilisation, and some
may have lapsed into utter barbarism, though on this
latter head I have not met with any evidence. The
Fuegians were probably compelled by other conquering
hordes to settle in their inhospitable country, and they
may have become in consequence somewhat more
degraded ; but it would be difficult to prove that they
have fallen much below the Botocudos who inhabit the
finest parts of Brazil.

The evidence that all civilised nations are the de-
scendants of barbarians, consists, on the one side, of
clear traces of their former low condition in still-existing
customs, beliefs, language, &c. ; and on the other side,
of proofs that savages are independently able to raise
themselves a few steps in the scale of civilisation, and
have actually thus risen. The evidence on the first
head is extremely curious, but cannot be here given :
I refer to such cases as that, for instance, of the art of
enumeration, which, as Mr. Tylor clearly shews by the
words still used in some places, originated in counting

29 'On tke Origin of Civilisation,' ' Proc. Ethnological Soc' Nov.
26, 1867.

29 'Primeval Man,' 1869.

182 THE DESCENT OF MAN. Part I.

the fingers, first of one hand and then of the other,
and lastly of the toes. We have traces of this in our
own decimal system, and in the Roman numerals, "which
after reaching to the number V., change into VI., &c,
when the other hand no doubt was used. So again,
" when we speak of three-score and ten, we are count-
" ing by the vigesimal system, each score thus ideally
" made, standing for 20 — for ' one man ' as a Mexican
" or Carib would put it." 31 According to a large and
increasing school of philologists, every language bears
the marks of its slow and gradual evolution. So it
is with the art of writing, as letters are rudiments of
pictorial representations. It is hardly possible to read
Mr. M'Lennan's work32 and not admit that almost all
civilised nations still retain some traces of such rude
habits as the forcible capture of wives. What ancient
nation, as the same author asks, can be named that was
originally monogamous ? The primitive idea of justice,
as shewn by the law of battle and other customs of
which traces still remain, was likewise most rude. Many
existing superstitions are the remnants of former false
religious beliefs. The highest form of religion — the
grand idea of God hating sin and loving righteousness
— was unknown during primeval times.

Turning to the other kind of evidence : Sir J. Lub-
bock has shewn that some savages have recently im-
proved a little in some of their simpler arts. From the

31 'Koyal Institution of Great Britain,' March 15, 1867. Also,
' Researches into the Early History of Mankind,' 1865.

32 'Primitive Marriage,' 1865. See, likewise, an excellent article,
evidently by the same author, in the ' North British Review,' July,
1869. Aho, Mr. L. H. Morgan, " A Conjectural Solution of the Origin
of the Class. System of Relationship," in 'Proc. American Acad, of
Sciences,' vol. vii. Feb. 1868. Prof. Schaaffhausen (' Anthropolog.
Review,' Oct. 1869, p. 373) remarks on "the vestiges of human sacri-
" fices found both in Homer and the Old Testament."

Chap. V. CIVILISED NATIONS. 183

extremely curious account which he gives of the weapons,
tools, and arts, used or practised by savages in various
parts of the world, it cannot be doubted that these have
nearly all been independent discoveries, excepting per
haps the art of making fire.33 The Australian boomerang
is a good instance of one such independent discovery.
The Tahitians when first visited had advanced in many
respects beyond the inhabitants of most of the other
Polynesian islands. There are no just grounds for the
belief that the high culture of the native Peruvians
and Mexicans was derived from any foreign source ; 34
many native plants were there cultivated, and a few
native animals domesticated. We should bear in mind
that a wandering crew from some semi-civilised land,
if washed to the shores of America, would not, judging
from the small influence of most missionaries, have pro-
duced any marked effect on the natives, unless they had
already become somewhat advanced. Looking to a very
remote period in the history of the world, we find, to
use Sir J. Lubbock's well-known terms, a paleolithic
and neolithic period ; and no one will pretend that the
art of grinding rough flint tools was a borrowed one.
In all parts of Europe, as far east as Greece, in Palestine,
India, Japan, New Zealand, and Africa, including Egypt,
flint tools have been discovered in abundance; and of
their use the existing inhabitants retain no tradition.
There is also indirect evidence of their former use by the
Chinese and ancient Jews. Hence there can hardly be a
doubt that the inhabitants of these many countries, which
include nearly the wdiole civilised world, were once in a
barbarous condition. To believe that man was abori-

33 Sir J. Lubbock, 'Prehistoric Times,' 2nd edit. 1869, chap. xv. and
xvi. et passim.

34 Dr. F. Midler has made some good remarks to this effect in the
'Keise der Novara : Anthropolog. Theil,' Abtheil. iii. 1868, s. 127.

184: THE DESCENT OF MAN. Part 1.

ginally civilised and then suffered utter degradation in
so many regions, is to take a pitiably low view of
hum an nature. It is apparently a truer and more
cheerful view that progress has been much more general
than retrogression ; that man has risen, though by slow
and interrupted steps, from a lowly condition to the
highest standard as yet attained by him in knowledge,
morals, and religion.

Chap. VI. AFFINITIES AND GENEALOGY. 185

CHAPTEE VI.

On the Affinities axd Genealogy of Man.

Position of man in the animal series — The natural system genea-
logical— 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 — Conclusion.

Even if it be granted that the difference between man
and his nearest allies is as great in corporeal structure as
some naturalists maintain, and although we must grant
that the difference between them is immense in mental
power, yet the facts given in the previous chapters
declare, as it appears to me, in the plainest manner,
that man is descended from some lower form, notwith-
standing that connecting-links have not hitherto been
discovered.

Man is liable to numerous, slight, and diversified
variations, which are induced by the same general
causes, are governed and transmitted in accordance
with the same general laws, as in the lower animals.
Man tends to multiply at so rapid a rate that his off-
spring are necessarily exposed to a struggle for existence,
and consequently to natural selection. He has given
rise to many races, some of which are so different that
they have often been ranked by naturalists as distinct
species. His body is constructed on the same homo-
logical plan as that of other mammals, independently
of the uses to which the several parts may be put. He

186 THE DESCENT OF MAN. Part I.

passes through the same phases of ernbryological de-
velopment. He retains many rudimentary and useless
structures, which no doubt were once serviceable. Cha-
racters occasionally make their re-appearance in him,
which we have every reason to believe were possessed by
his early progenitors. If the origin of man had been
wholly different from that of all other animals, these
various appearances would be mere empty deceptions ;
but such an admission is incredible. These appearances,
on the other hand, are intelligible, at least to a large
extent, if man is the co-descendant with other mammals
of some unknown and lower form.

Some naturalists, from being deeply impressed with
the mental and spiritual powers of man, have divided the
whole organic world into three kingdoms, the Human,
the Animal, and the Vegetable, thus giving to man a
separate kingdom.1 Spiritual powers cannot be com-
pared or classed by the naturalist ; but he may endea-
vour to shew, as I have done, that the mental faculties
of man and the lower animals do not differ in kind,
although immensely in degree. A difference in degree,
however great, does not justify us in placing man in a
distinct kingdom, as will perhaps be best illustrated
by comparing the mental powers of two insects, namely,
a coccus or scale-insect and an ant, which undoubt-
edly belong to the same class. The difference is here
greater, though of a somewhat different kind, than
that between man and the highest mammal. The
female coccus, whilst young, attaches itself by its pro-
boscis to a plant ; sucks the sap but never moves again ;
is fertilised and lays eggs ; and this is its whole history.
On the other hand, to describe the habits and mental

1 Isidore Geoffroy St.-Hilaire gives a detailed account of the position
assigned to man by various naturalists in their classifications : ' Hist.
Nat. Gen.' torn. ii. 1859, p. 170-189.

Chap. VI. AFFINITIES AND GENEALOGY. 187

powers of a female ant, would require, as Pierre Huber
has shewn, a large volume; I may, however, briefly
specify a few points. Ants communicate information
to each other, and several unite for the same work,
or games of play. They recognise their fellow-ants
after months of absence. They build great edifices,
keep them clean, close the doors in the evening, and
post sentries. They make roads, and even tunnels
under rivers. They collect food for the community,
and when an object, too large for entrance, is brought
to the nest, they enlarge the door, and afterwards
build it up again.2 They go out to battle in regular
bands, and freely sacrifice their lives for the common
weal. They emigrate in accordance with a precon-
certed plan. They capture slaves. They keep Aphides
as milch-cows. They move the eggs of their aphides,
as well as their own eggs and cocoons, into warm parts
of the nest, in order that they may be quickly hatched ;
and endless similar facts could be given. On the
whole, the difference in mental power between an ant
and a coccus is immense ; yet no one has ever dreamed
of placing them in distinct classes, much less in distinct
kingdoms. No doubt this interval is bridged over by
the intermediate mental powers of many other insects ;
and this is not the case with man and the higher apes.
But we have every reason to believe that breaks in the
series are simply the result of many forms having be-
come extinct.

Professor Owen, relying chiefly on the structure of
the brain, has divided the mammalian series into four
sub-classes. One of these he devotes to man ; in another
he places both the marsupials and the monotremata;
so that he makes man as distinct from all other mam-

2 See the very interesting article, " L'lnstinct cliez les Insectes," by
M. George Pouchet, ' Revue des Deux Mondes,' Feb. 1870, p. 682.

188 THE DESCENT OF MAN. Part I.

. inals as are these two latter groups conjoined. This
view has not been accepted, as far as I am aware, by
any naturalist capable of forming an independent judg-
ment, and therefore need not here be further con-
sidered.

We can understand why a classification founded on
any single character or organ — even an organ so won-
derfully complex and important as the brain — -or on the
high development of the mental faculties, is almost sure
to prove unsatisfactory. This principle has indeed been
tried with hymenopterous insects; but when thus classed
by their habits or instincts, the arrangement proved
thoroughly artificial.3 Classifications may, of course, be
based on any character whatever, as on size, colour, or
the element inhabited ; but naturalists have long felt a
profound conviction that there is a natural system. This
system, it is now generally admitted, must be, as far
as possible, genealcgical in arrangement, — that is, the
co-descendants of the same form must be kept together
in one group, separate from the co-descendants of any
other form ; but if the parent-forms are related, so will
be their descendants, and the two groups together will
form a larger group. The amount of difference between
the several groups — that is the amount of modification
which each has undergone — will be expressed by such
terms as genera, families, orders, and classes. As we
have no record of the lines of descent, these lines can
be discovered only by observing the degrees of re-
semblance between the beings which are to be classed.
For this object numerous points of resemblance are of
much more importance than the amount of similarity
or dissimilarity in a few points. If two languages
were found to resemble each other in a multitude of

3 Westwood, ' Modern Class, of Insects,' vol. ii. 1840, p. S7.

Chap. VI. AFFINITIES AND GENEALOGY. 189

words and points of construction, they would be uni-
versally recognised as having sprung from a common
source, notwithstanding that they differed greatly in
some few words or points of construction. But with
organic beings the points of resemblance must not con-
sist of adaptations to similar habits of life : two animals
may, for instance, have had their whole frames modified
for living in the water, and yet they will not be brought
any nearer to each other in the natural system. Hence
we can see how it is that resemblances in unimportant
structures, in useless and rudimentary organs, and in
parts not as yet fully developed or functionally active,
are by far the most serviceable for classification ; for
they can hardly be due to adaptations within a late
period ; and thus they reveal the old lines of descent
or of true affinity.

We can further see why a great amount of modifi-
cation in some one character ought not to lead us to
separate widely any two organisms. A part which
already differs much from the same part in other allied
forms has already, according to the theory of evolution,
varied much ; consequently it would (as long as the
organism remained exposed to the same exciting con-
ditions) be liable to further variations of the same kind ;
and these, if beneficial, would be preserved, and thus
continuallv augmented. In manv cases the continued
development of a part, for instance, of the beak of a
bird, or of the teeth of a mammal, would not be advan-
tageous to the species for gaining its food, or for any
other object; but with man we can see no definite limit,
as far as advantage is concerned, to the continued de-
velopment of the brain and mental faculties. Therefore
in determining the position of man. in the natural or
genealogical system, the extreme development of his
brain ought not to outweigh a multitude of resem-

190 THE DESCENT OF MAX. Part I.

blances in other less important or quite unimportant
points.

The greater number of naturalists who have taken
into consideration the whole structure of man, including
his mental faculties, have followed Blumenbach and
Cuvier, and have placed man in a separate Order, under
the title of the Bimana, and therefore on an equality
with the Orders of the Quadrumana, Carnivora, &c.
liecently many of our best naturalists have recurred to
the view first propounded by Linnaeus, so remarkable for
his sagacity, and have placed man in the same Order
with the Quadrumana, under the title of the Primates.
The justice of this conclusion will be admitted if, in
the first place, we bear in mind the remarks just
made on the comparatively small importance for classi-
fication of the great development of the brain in man ;
bearing, also, in mind that the strongly-marked differ-
ences between the skulls of man and the Quadrumana
(lately insisted upon by Bischoff, Aeby, and others)
apparently follow from their differently developed brains.
In the second place, we must remember that nearly all
the other and more important differences between man
and the Quadrumana are manifestly adaptive in their
nature, and relate chiefly to the erect position of man ;
such as the structure of his hand, foot, and pelvis, the
curvature of his spine, and the position of his head.
The family of seals offers a good illustration of the
small importance of adaptive characters for classification.
These animals differ from all other Carnivora in the
form of their bodies and in the structure of their limbs,
far more than does man from the higher apes : yet in
every system, from that of Cuvier to the most recent
one by Mr. Flower,4 seals are ranked as a mere familv

4 ' Proc. Zoolog. Soc' 1869, p. 4.

Chap. VI. AFFINITIES AND GENEALOGY. 191

in the Order of the Carnivora. If man had not been
his own classifier, he would never have thought of
founding a separate order for his own reception.

It would be beyond my limits, and quite beyond my
knowledge, even to name the innumerable points of
structure in which man agrees with the other Primates.
Our great anatomist and philosopher, Prof. Huxley, has
fully discussed this subject,5 and has come to the con-
clusion that man in all parts of his organisation differs
less from the higher apes, than these do from the lower
members of the same group. Consequently there " is
" no justification for placing man in a distinct order."

In an early part of this volume I brought forward
various facts, shewing how closely man agrees in con-
stitution with the higher mammals ; and this agreement,
no doubt, depends on our close similarity in minute
structure and chemical composition. I gave, as
instances, our liability to the same diseases, and to the
attacks of allied parasites ; our tastes in common for the
same stimulants, and the similar effects thus produced,
as wrell as by various drugs ; and other such facts.

As small unimportant points of resemblance between
man and the higher apes are not commonly noticed in
systematic works, and as, when numerous, they clearly
reveal our relationship, I will specify a few such points.
The relative position of the features are manifestly the
same in man and the Quadrumana; and the various
emotions are displayed by nearly similar movements of
the muscles and skin, chiefly above the eyebrows and
round the mouth. Some few expressions are, indeed,
almost the same, as in the weeping of certain kinds of
monkeys, and in the laughing noise made by others,
durino: which the corners of the mouth are drawn back-

5 «

Evidence as to Man's Place in Nature,' 1SG3, p. 70, et passim.

192 THE DESCENT OF MAX. Part I.

wards, and the lower eyelids wrinkled. The external
ears are curiously alike. In man the nose is much
more prominent than in most monkeys ; but we may
trace the commencement of an aquiline curvature in
the nose of the Hoolock Gibbon ; and this in the Sem-
nopitheeus nasica is carried to a ridiculous extreme.

The faces of many monkeys are ornamented with
beards, whiskers, or moustaches. The hair on the head
grows to a great length in some species of Semno-
pithecus ; 6 and in the Bonnet monkey (Macacus
radiatiis) it radiates from a point on the crown, with a
parting down the middle, as in man. It is commonly
said that the forehead gives to man his noble and intel-
lectual appearance ; but the thick hair on the head of
the Bonnet monkey terminates abruptly downwards,
and is succeeded by such short and fine hair, or down,
that at a little distance the forehead, with the exception
of the eyebrows, appears quite naked. It has been
erroneously asserted that eyebrows are not present in
any monkey. In the species just named the degree of
nakedness of the forehead differs in different individuals;
and Eschricht states7 that in our children the limit
between the hairy scalp and the naked forehead is
sometimes not well defined ; so that here we seem to
have a trifling case of reversion to a progenitor, in whom
the forehead had not as yet become quite naked.

It is well known that the hair on our arms tends to
converge from above and below to a point at the elbow.
This curious arrangement, so unlike that in most of the
lower mammals, is common to the gorilla, chimpanzee,
orang, some species of Hylobates, and even to some few
American monkeys. But in Hylobates agUis the hair

6 Isid. Geoffroy, ' Hist. Nat. Gen.' torn. ii. 1859, p. 217.

7 " Ueher die Richtung der Haare," &c, Miillei-'s ' Arcliiv fur Anat.
und Phys.' 1837, s. 51.

Chap. VI. AFFINITIES AND GENEALOGY. 193

on the fore-arm is directed downwards or towards the
wrist in the ordinary manner ; and in H. lar it is nearly
erect, with only a very slight forward inclination ; so
that in this latter species it is in a transitional state.
It can hardly be doubted that with most mammals the
thickness of the hair and its direction on the back is
adapted to throw off the rain ; even the transverse hairs
on the fore-legs of a dog may serve for this end when he
is coiled up asleep. Mr. Wallace remarks that the con-
vergence of the hair towards the elbow on the arms
of the orang (whose habits he has so carefully studied)
serves to throw off the rain, when, as is the custom
of this animal, the arms are bent, with the hands
clasped round a branch or over its own head. We
should, however, bear in mind that the attitude of an
animal may perhaps be in part determined by the
direction of the hair ; and not the direction of the hair
by the attitude. If the above explanation is correct in
the case of the orang, the hair on our fore-arms offers a
curious record of our former state ; for no one supposes
that it is now of any use in throwing off the rain, nor in
our present erect condition is it properly directed for this
purpose.

It would, however, be rash to trust too much to the
principle of adaptation in regard to the direction of the
hair in man or his early progenitors ; for it is impossible
to study the figures given by Eschricht of the arrange-
ment of the hair on the human foetus (this being the
same as in the adult) and not agree with this excellent
observer that other and more complex causes have
intervened. The points of convergence seem to stand
in some relation to those points in the embryo which
are last closed in during development. There appears,
also, to exist some relation between the arrangement

VOL. I. O

1 94 THE DESCENT OF MAN. Part I.

of the hair on the limbs, and the course of the medullary-
arteries.8

It must not be supposed that the resemblances be-
tween man and certain apes in the above and many
other points — such as in having a naked forehead,
long tresses on the head, &c. — are all necessarily the
result of unbroken inheritance from a common pro-
genitor thus characterised, or of subsequent reversion.
Many of these resemblances are more probably due
to analogous variation, which follows, as I have else-
where attempted to shew,9 from co-descended organisms
having a similar constitution and having been acted
on by similar causes inducing variability. With re-
spect to the similar direction of the hair on the fore-
arms of man and certain monkeys, as this character is
common to almost all the anthropomorphous apes, it
may probably be attributed to inheritance ; but not
certainly so, as some very distinct American monkeys
are thus characterised. The same remark is applicable
to the tailless condition of man ; for the tail is absent
in all the anthropomorphous apes. Nevertheless this
character cannot with certainty be attributed to inheri-
tance, as the tail, though not absent, is rudimentary
in several other Old World and in some New World
species, and is quite absent in several species belonging
to the allied group of Lemurs.

Although, as we have now seen, man has no just right
to form a separate Order for his own reception, he may

8 On the hair in Hylobates, see ' Nat. Hist, of Mammals,' by C. L.
Martin, 1841, p. 415. Also, Isid. Geoffroy on the American monkeys
and other kinds, ' Hist. Nat. Gen.' vol. ii. 1859, p. 216, 243. Esch-
richt, ibid, s. 46, 55, 61. Owen, ' Anat. of Vertebrates,' vol. iii. p. 619.
Wallace, ' Contributions to the Theory of Natural Selection,' 1870, p.
344.

9 'Origin of Species,' 5th edit. 1869, p. 194. 'The Variation of
Animals and Plants under Domestication,' vol. ii. 1868, p. 348.

Chap. VI. AFFINITIES AND GENEALOGY. 195

perhaps claim a distinct Sub-order or Family. Prof.
Huxley, in liis last work,10 divides the Primates into
three Sub-orders ; namely, the Anthropidae with man
alone, the Simiadse including monkeys of all kinds, and
the Lemuridaa with the diversified genera of lemurs. As
far as differences in certain important points of structure
are concerned, man may no doubt rightly claim the
rank of a Sub-order ; and this rank is too low, if we look
chiefly to his mental faculties. Nevertheless, under a
genealogical point of view it appears that this rank is
too high, and that man ought to form merely a Family,
or possibly even only a Sub-family. If we imagine
three lines of descent proceeding from a common source,
it is quite conceivable that two of them might after
the lapse of ages be so slightly changed as still to
remain as species of the same genus ; whilst the third
line might become so greatly modified as to deserve
to rank as a distinct Sab-family, Family, or even
Order. But in this case it is almost certain that
the third line would still retain through inheritance
numerous small points of resemblance with the other
two lines. Here then would occur the difficulty, at
present insoluble, how much weight we ought to assign
in our classifications to strongly-marked differences in
some few points, — that is to the amount of modification
undergone ; and how much to close resemblance in
numerous unimportant points, as indicating the lines of
descent or genealogy. The former alternative is the
most obvious, and perhaps the safest, though the latter
appears the most correct as giving a truly natural
classification.

To form a judgment on this head, with reference
to man we must glance at the classification of the

10 ' An Introduction to the Classification of Animals,' 1869, p. 99.

o 2

19(3 THE DESCENT OF MAN. Part I.

Siniiadse. This family is divided by almost all natura-
lists into the Catarhine group, or Old World monkeys,
all of which are characterised (as their name expresses)
by the peculiar structure of their nostrils and by having
four premolars in each jaw ; and into the Platyrhine
group or New World monkeys (including two very
distinct sub-groups), all of which are characterised by
differently-constructed nostrils and by having six pre-
molars in each jaw. Some other small differences might
be mentioned. Now man unquestionably belongs in
his dentition, in the structure of his nostrils, and some
other respects, to the Catarhine or Old World division ;
nor does he resemble the Platyrhines more closely than
the Catarhines in any characters, excepting in a few
of not much importance and apparently of an adaptive
nature. Therefore it would be against all probability
to suppose that some ancient New World species had
varied, and had thus produced a man-like creature with
all the distinctive characters proper to the Old World
division ; losing at the same time all its own distinctive
characters. There can consequently hardly be a doubt
that man is an offshoot from the Old World Simian stem ;
and that under a genealogical point of view, he must
be classed with the Catarhine division.11

The anthropomorphous apes, namely the gorilla,
chimpanzee, orang, and hylobates, are separated as a
distinct sub-group from the other Old World monkeys
by most naturalists. I am aware that Gratiolet, relying
on the structure of the brain, does not admit the exist-

11 This is nearly the same classification as that provisionally adopted
by Mr. St. George Mivart (' Transact. Philosoph. Soc.' 1867, p. 300),
who, after separating the Lemuridoe, divides the remainder of the
Primates into the Hominidaj, the Simiadae answering to the Catarhines,
the Cebidae, and the Hapalidse, — these two latter groups answering to
the Plat vrl tines.

Chap. VI. AFFINITIES AND GENEALOGY. 197

ence of this sub-group, and no doubt it is a broken
one ; tbus the orang, as Mr. St. G, Mivart remarks,12
" is one of the most peculiar and aberrant forms to be
" found in the Order." The remaining, non-anthropo-
morphous, Old World monkeys, are again divided by
some naturalists into two or three smaller sub-groups ;
the genus Semnopithecus, with its peculiar sacculated
stomach, being the type of one such sub-group. But
it appears from M. Gaudry's wonderful discoveries in
Attica, that during the Miocene period a form existed
there, which connected Semnopithecus and Macacus;
and this probably illustrates the manner in which the
other and higher groups were once blended together.

If the anthropomorphous apes be admitted to form
•a natural sub-group, then as man agrees with them,
not only in all those characters which he possesses in
common with the whole Catarhine group, but in other
peculiar characters, such as the absence of a tail and
of callosities and in general appearance, we may infer
that some ancient member of the anthropomorphous
sub-group gave birth to man. It is not probable
that a member of one of the other lower sub-groups
should, through the law of analogous variation, have
given rise to a man-like creature, resembling the higher
anthropomorphous apes in so many respects. No
doubt man, in comparison with most of his allies, has
undergone an extraordinary amount of modification,
chiefly in consequence of his greatly developed brain
and erect position ; nevertheless we should bear in
mind that he " is but one of several- exceptional forms
" of Primates."13

Every naturalist, who believes in the principle of

12 < Transact. Zoolog. Soc.' vol. vi. 1867, p. 214.

13 Mr. St. G. Mivart, ' Transact. Phil. Soc' 18G7, p. 410.

198 THE DESCENT OF MAN. Part I.

evolution, will grant that the two main divisions of the
Simiadse, namely the Catarhine and Platyrhine mon-
keys, with their sub-groups, have all proceeded from
some one extremely ancient progenitor. The early
descendants of this progenitor, before they had diverged
to any considerable extent from each other, would still
have formed a single natural group ; but some of the
species or incipient genera would have already begun
to indicate by their diverging characters the future
distinctive marks of the Catarhine and Platyrhine divi-
sions. Hence the members of this supposed ancient
group would not have been so uniform in their dentition
or in the structure of their nostrils, as are the existing
Catarhine monkeys in one way and the Platyrhines in
another way, but would have resembled in this respect
the allied Lemuridse which differ ^reatlv from each
other in the form of their muzzles,14 and to an extra-
ordinary degree in their dentition.

The Catarhine and Platyrhine monkeys agree in
a multitude of characters, as is shewn by their unques-
tionably belonging to one and the same Order. The
many characters which they possess in common can
hardly have been independently acquired by so many
distinct species; so that these characters must have
been inherited. But an ancient form which possessed
many characters common to the Catarhine and Platy-
rhine monkeys, and others in an intermediate condition,
and some few perhaps distinct from those now present
in either group, wrould undoubtedly have been ranked,
if seen by a naturalist, as an aue or monkey. And as
man under a genealogical puint of view belongs to the
Catarhine or Old World stock, we must conclude, how-

14 Messrs. Murie and Mivart on the Lenmroidea, 'Transact. Zoolug.
Soc.' vol. vii. 1869, p. 5.

Chap. VI. AFFINITIES AND GENEALOGY. 199

ever much the conclusion may revolt our pride, that
our early progenitors would have been properly thus
designated.15 But we must not fall into the error of
supposing that the early progenitor of the whole Simian
stock, including man, was identical with, or even closely
resembled, any existing ape or monkey.

On the Birthplace and Antiquity of Man. — We are
naturally led to enquire where was the birthplace of
man at that stage of descent when our progenitors
diverged from the Catarhine stock. The fact that
they belonged to this stock clearly shews that they
inhabited the Old World ; but not Australia nor any
oceanic island, as we may infer from the laws of geogra-
phical distribution. In each great region of the world
the living mammals are closely related to the extinct
species of the same region. It is therefore probable
that Africa wras formerly inhabited by extinct apes
closely allied to the gorilla and chimpanzee ; and as
these two species are now man's nearest allies, it is
somewhat more probable that our early progenitors
lived on the African continent than elsewhere. But
it is useless to speculate on this subject, for an ape
nearly as large as a man, namely the Dryopithecus
of Lartet, which was closely allied to the anthropo-
morphous Hylobates, existed in Europe during the
Upper Miocene period ; and since so remote a period
the earth has certainly undergone many great revo-
lutions, and there has been ample time for migration
on the largest scale.

15 H'ackel has come to this same conclusion. See ' Ueher die Ent-
stehung des Menschengeschlechts,' in Virchow's ' Sammlung. gemein.
wissen. Vortrage,' 18G8, s. 61. Also his 'Naturliche Schopfungs-
geschichte,' 1868, in which he gives in detail his views on the geuea-
logy of man.

200 THE DESCENT OF MAN. Part I.

At the period and place, whenever and wherever it
may have been, when man first lost his hairy covering,
he probably inhabited a hot country; and this would
have been favourable for a frugiferous diet, on which,
judging from analogy, he subsisted. We are far from
knowing how long ago it was when man first diverged
from the Catarhine stock ; but this may have occurred
at an epoch as remote as the Eocene period ; for the
higher apes had diverged from the lower apes as
early as the Upper Miocene period, as shewn by the
existence of the Dryopithecus. We are also quite
ignorant at how rapid a rate organisms, whether high
or low in the scale, may under favourable circumstances
be modified : we know, however, that some have retained
the same form during an enormous lapse of time. From
what we see going on under domestication, we learn that
within the same period some of the co-descendants of
the same species may be not at all changed, some a
little, and some greatly changed. Thus it may have
been with man, who has undergone a great amount of
modification in certain characters in comparison with
the higher apes.

The great break in the organic chain between man
and his nearest allies, which cannot be bridged over
by any extinct or living species, has often been ad-
vanced as a grave objection to the belief that man
is descended from some lower form ; but this objection
will not appear of much weight to those who, convinced
by general reasons, believe in the general principle
of evolution. Breaks incessantly occur in all parts
of the series, some being wide, sharp and defined, others
less so in various degrees; as between the orang and
its nearest allies — between the Tarsius and the other
Lemuridas — between the elephant and in a more
striking manner between the Ornithorhynchus or

Chap. VI. AFFINITIES AND GENEALOGY. 201

Echidna, and other mammals. But all these breaks
depend merely on the number of related forms which
have become extinct. At some future period, not
very distant as measured by centuries, the civilised races
of man will almost certainly exterminate and replace
throughout the world the savage races. At the
same time the anthropomorphous apes, as Professor
Schaaffhausen has remarked,16 will no doubt be exter-
minated. The break will then be rendered wider, for
it will intervene between man in a more civilised state,
as we may hope, than the Caucasian, and some ape as
low as a baboon, instead of as at present between the
negro or Australian and the gorilla.

With respect to the absence of fossil remains, serving to
connect man with his ape-like progenitors, no one will
lay much stress on this fact, who will read Sir C. Lyell's
discussion,17 in which he shews that in all the vertebrate
classes the discovery of fossil remains has been an
extremely slow and fortuitous process. Nor should it
be forgotten that those regions which are the most
likely to afford remains connecting man with some
extinct ape-like creature, have not as yet been searched
by geologists.

Lower Stages in the Genealogy of Man. — "We have seen
that man appears to have diverged from the Catarhine
or Old World division of the Simiadse, after these had
diverged from the New World division. We will now
endeavour to follow the more remote traces of his
genealogy, trusting in the first place to the mutual
affinities between the various classes and orders, with
some slight aid from the periods, as far as ascertained,

16 4

17 «

Anthropological Keview,' April, 1867, p. 236. y^C\ C A

Elements of Geology/ 1865, p. 583-585. 'Antiquity of/MH^y
1863, p. 115. A^000° *

L LIBRA

202 THE DESCENT OF MAN. Part I.

of their successive appearance on the earth. The
LemuridsB stand below and close to the Simiadae, con-
stituting a very distinct family of the Primates, or,
according to Hackel, a distinct Order. This group is
diversified and broken to an extraordinary degree, and
includes many aberrant forms. It has, therefore, pro-
bablv suffered much extinction. Most of the remnants

at

survive on islands, namely in Madagascar and in the
islands of the Malayan archipelago, where they have
not been exposed to such severe competition as they
would have been on well-stocked continents. This
group likewise presents many gradations, leading, as
Huxley remarks,18 " insensibly from the crown and
" summit of the animal creation down to creatures
" from which there is but a step, as it seems, to the
" lowest, smallest, and least intelligent of the placental
" mammalia." From these various considerations it is
probable that the Simiadae were originally developed
from the progenitors of the existing Lemuriclae ; and
these in their turn from forms standing very low in the
mammalian series.

The Marsupials stand in many important characters
below the placental mammals. They appeared at an
earlier geological period, and their range was formerly
much more extensive than what it now is. Hence the
Placentata are generally supposed to have been derived
from the Implacentata or Marsupials ; not, however, from
forms closely like the existing Marsupials, but from
their early progenitors. The Monotremata are plainly
allied to the Marsupials ; forming a third and still
lower division in the great mammalian series. They
are represented at the present day solely by the Orni-
thorhynchus anrl Echidna ; and these two forms may

18 ' Man's Place in Nature,' p. 105.

Chap. VI. AFFINITIES AND GENEALOGY. 203

be safely considered as relics of a much larger group
which have been preserved in Australia through some
favourable concurrence of circumstances. The 3Iono-
tremata are eminently interesting, as in several
important points of structure they lead towards the
class of reptiles.

In attempting to trace the genealogy of the Mam-
malia, and therefore of man, lower down in the series,
we become involved in greater and greater obscurity.
He who wishes to see what ingenuity and knowledge
can effect, may consult Prof. Hackel's works.19 I will
content myself with a few general remarks. Every
evolutionist will admit that the five great vertebrate
classes, namely, mammals, birds, reptiles, amphibians,
and fishes, are all descended from some one prototype ;
for they have much in common, especially during their
embryonic state. As the class of fishes is the most
lowly organised and appeared before the others, we may
conclude that all the members of the vertebrate king-
dom are derived from some fish-like animal, less highly
organised than any as yet found in the lowest known
formations. The belief that animals so distinct as a
monkey or elephant and a humming-bird, a snake, frog,
and fish, &c, could all have sprung from the same
parents, will appear monstrous to those who have not
attended to the recent progress of natural history. For
this belief implies the former existence of links closely
binding together all these forms, now so utterly unlike.

19 Elaborate tables are given in his ' Generelle Morphologie ' (B. ii.
s. cliii. and s. 425) ; and with more especial reference to man in his
' Natiirliche Schopfungsgeschichte,' 1868. Prof. Huxley, in reviewing
this latter work (' The Academy/ 1869, p. 42) says, that he considers
the phylum or lines of descent of the Vertebrata to be admirably dis-
cussed by H'aekel, although he differs on some points. He expresses,
also, his high estimate of the value of the general tenor and spirit of
the whole work.

204 THE DESCENT OF MAN. Part I.

Nevertheless it is certain that groups of animals have
existed, or do now exist, which serve to connect more or
less closely the several great vertebrate classes. We
have seen that the Ornithorhynchus graduates towards
reptiles; and Prof. Huxley has made the remarkable
discovery, confirmed by Mr. Cope and others, that the
old Dinosaurians are intermediate in many important
respects between certain reptiles and certain birds — the
latter consisting of the ostrich-tribe (itself evidently a
widely-diffused remnant of a larger group) and of the
Archeopteryx, that strange Secondary bird having a
long tail like that of the lizard. Again, according to
Prof. Owen,20 the Ichthyosaurians — great sea-lizards fur-
nished with paddles — present many affinities with fishes,
or rather, according to Huxley, with amphibians. This
latter class (including in its highest division frogs and
toads) is plainly allied to the Ganoid fishes. These
latter fishes swarmed during the earlier geological
periods, and were constructed on what is called a highly
generalised type, that is they presented diversified affi-
nities with other groups of organisms. The amphibians
and fishes are also so closely united by the Lepidosiren,
that naturalists long disputed in which of these two
classes it ought to be placed. The Lepidosiren and
some few Ganoid fishes have been preserved from utter
extinction by inhabiting our rivers, which are harbours
of refuge, bearing the same relation to the great waters
of the ocean that islands bear to continents.

Lastly, one single member of the immense and diver-
sified class of fishes, namely the lancelet or amphioxus,
is so different from all other fishes, that Hackel main-
tains that it ought to form a distinct class in the
vertebrate kingdom. This fish is remarkable for its

20 ' ralceontoiu-v,' 1860, p. 199.

Chap. VI. AFFINITIES AND GENEALOGY. 205

negative characters ; it can hardly be said to possess a
brain, vertebral column, or heart, &c. ; so that it was
classed by the older naturalists amongst the worms.
Many years ago Prof. Goodsir perceived that the
lancelet presented some affinities with the Ascidians,
which are invertebrate, hermaphrodite, marine crea-
tures permanently attached to a support. They hardly
appear like animals, and consist of a simple, tough,
leathery sack, with two small projecting orifices. They
belong to the Molluscoida of Huxley — a lower division
of the great kingdom of the Mollusca ; but they have
recently been placed by some naturalists amongst the
Vermes or worms. Their larvae somewhat resemble
tadpoles in shape,21 and have the power of swimming
freely about. Some observations lately made by M.
Kowalevsky,22 since confirmed by Prof. Kuppfer, will
form a discovery of extraordinary interest, if still further
extended, as I hear from M. Kowalevsky in Naples he
has now effected. The discovery is that the larvae of
Ascidians are related to the Vertebrata, in their manner
of development, in the relative position of the nervous
system, and in possessing a structure closely like the
chorda dor salts of vertebrate animals. It thus appears,
if we may rely on embryology, which has always proved
the safest guide in classification, that we have at last
gained a clue to the source whence the Vertebrata have

21 I had the satisfaction of seeing, at the Falkland Islands, in April,
1833, and therefore some years before any other naturalist, the loco-
motive larvae of a compound Ascidian, closely allied to, but apparently
generically distinct from, Synoicum. The tail was about five times as
long as the oblong head, and terminated in a very fine filament. It
was plainly divided, as sketched by me under a simple microscope, by
transverse opaque, partitions, which I presume represent the great cells
figured by Kowalevsky. At an early stage of development the tail was
closely coiled round the head of the larva.

22 ' Memoires de l'Acad. des Sciences de St. Petersbourg,' torn. x.
No. 15, 18G6.

206 THE DESCENT OF MAN. Part I.

been derived. We should thus be justified iu believing
that at an extremely remote period a group of animals
existed, resembling in many respects the larvae of our
present Ascidians, which diverged into two great
branches — the one retrograding in development and
producing the present class of Ascidians, the other rising
to the crown and summit of the animal kingdom by
giving birth to the Vertebrata.

We have thus far endeavoured rudely to trace the
genealogy of the Vertebrata by the aid of their mutual
affinities. We will now look to man as he exists ; and
we shall, I think, be able partially to restore during
successive periods, but not in due order of time, the
structure of our early progenitors. This can be effected
by means of the rudiments which man still retains, by
the characters which occasionally make their appear-
ance in him through reversion, and by the aid of the
principles of morphology and embryology. The various
facts, to which I shall here allude, have been given in
the previous chapters. The early progenitors of man
were no doubt once covered with hair, both sexes
having beards ; their ears were pointed and capable of
movement ; and their bodies were provided with a tail,
having the proper muscles. Their limbs and bodies
were also acted on by many muscles which now
only occasionally reappear, but are normally present
in the Quadrumana. The great artery and nerve of
the humerus ran through a supra-condyloid foramen.
At this or some earlier period, the intestine gave forth
a much larger diverticulum or caecum than that now
existing. The foot, judging from the condition of the
great toe in the foetus, was then prehensile; and our
progenitors, no doubt, were arboreal m their habits,
frequenting some warm, forest-clad land. The males

Chap. VI. AFFINITIES AND GENEALOGY. 207

were provided with great canine teeth, which served
them as formidable weapons.

At a much earlier period the uterus was double ; the
excreta were voided through a cloaca ; and the eye
was protected by a third eyelid or nictitating mem-
brane. At a still earlier period the progenitors of
man must have been aquatic in, their habits ; for
morphology plainly tells us that our lungs consist of a
modified swim-bladder, which once served as a float.
The clefts on the neck in the embryo of man show
where the branchiae once existed. At about this period
the true kidneys were replaced by the corpora wolffiana.
The heart existed as a simple pulsating vessel; and
the chorda dorsalis took the place of a vertebral column.
These early predecessors of man, thus seen in the dim
recesses of time, must have been as lowly organised
as the lancelet or amphioxus, or even still more lowly
organised.

There is one other point deserving a fuller notice.
It has long been known that in the vertebrate king-
dom one sex bears rudiments of various accessory
parts, appertaining to the reproductive system, which
properly belong to the opposite sex; and it has now
been ascertained that at a very early embryonic period
both sexes possess true male and female glands. Hence
some extremely remote progenitor of the whole verte-
brate kingdom appears to have been hermaphrodite or
androgynous.23 But here we encounter a singular

23 This is the conclusion of one of the highest authorities in com-
parative anatomy, namely, Prof. Gegenbaur : ' Grundziige der vergleich.
Anat." 1870, s. 876. The result has been arrived at chiefly from the
study of the Amphibia ; but it appears from the researches of Waldeyer
(as quoted in Humphry's 'Journal of Anat. and Phys.' 1869, p. 161),
that the sexual organs of even " the higher vertebrata are, in their early
" condition, hermaphrodite." Similar views have long been held by
some authors, though until recently not well based.

208 THE DESCENT OF MAN. Part I.

difficulty. In the mammalian class the males possess
in their vesiculaa prostraticae rudiments of a uterus with
the adjacent passage ; they bear also rudiments of
mammae, and some male marsupials have rudiments
of a marsupial sack.24 Other analogous facts could be
added. Are we, then, to suppose that some extremely
ancient mammal possessed organs proper to both sexes,
that is, continued androgynous after it had acquired
the chief distinctions of its proper class, and therefore
after it had diverged from the lower classes of the
vertebrate kingdom ? This seems improbable in the
highest degree; for had this been the case, we might
have expected that some few members of the two
lower classes, namely fishes25 and amphibians, would
still have remained androgynous. We must, on the
contrary, believe that when the five vertebrate classes
diverged from their common progenitor the sexes
had already become separated. To account, however,
for male mammals possessing rudiments of the acces-
sory female organs, and for female mammals possessing
rudiments of the masculine organs, we need not suppose
that their early progenitors were still androgynous after
they had assumed their chief mammalian characters.
It is quite possible that as the one sex gradually
acquired the accessory organs proper to it, some of the
successive steps or modifications were transmitted to
the opposite sex. When we treat of sexual selection,
we shall meet writh innumerable instances of this form
of transmission, — as in the case of the spurs, plumes,

24 The male Thylacinus offers the best instance. Owen, ' Anatomy
of Vertebrates,' vol. iii. p. 771.

25 Serranus is well known often to be in an hermaphrodite condition ;
but Dr. Giinther informs me that he is convinced that this is not its
normal state. Descent from an ancient androgynous prototype would,
however, naturally favour end explain, to a certain extent, the recur-
rence of this condition in these fishes.

Chap. VI. AFFINITIES AND GENEALOGY. 209

and brilliant colours, acquired by male birds for battle
or ornament, and transferred to the females in an im-
perfect or rudimentary condition.

The possession by male mammals of functionally
imperfect mammary organs is, in some respects, espe-
cially curious. The Monotremata have the proper milk-
secreting glands with orifices, but no nipples; and as
these animals stand at the very base of the mam-
malian series, it is probable that the progenitors of the
class possessed, in like manner, the milk-secreting
glands, but no nipples. This conclusion is supported
by what is known of their manner of development ;
for Professor Turner informs me, on the authority of
Kolliker and Lauger, that in the embryo the mammary
glands can be distinctly traced before the nipples are
in the least visible ; and it should be borne in mind that
the development of successive parts in the individual
generally seems to represent and accord with the deve-
lopment of successive beings in the same line of descent.
The Marsupials differ from the Monotremata by possess-
ing nipples ; so that these organs were probably first
acquired by the Marsupials after they had diverged
from, and risen above, the Monotremata, and were
then transmitted to the placental mammals. No one
will suppose that after the Marsupials had approxi-
mately acquired their present structure, and therefore
at a rather late period in the development of the
mammalian series, any of its members still remained
androgynous. We seem, therefore, compelled to recur
to the foregoing view, and to conclude, that the nipples
were first developed in the females of some very early
marsupial form, and were then, in accordance with a
common law of inheritance, transferred in a functionally
imperfect condition to the males.

Nevertheless a suspicion has sometimes crossed my
vol. i. p

210 THE DESCENT OF MAN. Part I.

mind that long after the progenitors of the whole
mammalian class had ceased to be androgynous, both
sexes might have yielded milk and thus nourished
their young; and in the case of the Marsupials, that
both sexes might have carried their young in mar-
supial sacks. This will not appear utterly incredible, if
we reflect that the males of svno-nathous fishes receive
the eggs of the females in their abdominal pouches,
hatch them, and afterwards, as some believe, nourish
the young;26 — that certain other male fishes hatch the
eggs within their mouths or branchial cavities ; — that
certain male toads take the chaplets of eggs from the
females and wind them round their own thighs, keep-
ing them there until the tadpoles are born ; — that cer-
tain male birds undertake the whole duty of incubation,
and that male pigeons, as well as the females, feed their
nestlings with a secretion from their crops. But the
above suspicion first occurred to me from the mammary
glands in male mammals being developed so much more
perfectly than the rudiments of those other accessory
reproductive parts, which are found in the one sex
though proper to the other. The mammary glands
and nipples, as they exist in male mammals, can indeed
hardly be called rudimentary ; they are simply not
fully developed and not functionally active. They are
sympathetically affected under the influence of certain
diseases, like the same organs in the female. At birth
they often secrete a few drops of milk ; and they have

26 Mr. Lockwood believes (as quoted in ' Quart. ,Journal of Science,'
April, 1868, p. 269), from what he has observed of the development of
Hippocampus, that the walls of the abdominal pouch of the male
in some way afford nourishment. On male fishes hatching the ova in
their mouths, see a very interesting paper by Prof. Wyman, in ' Proc.
Boston Soc. of Nat. Hist.' Sept. 15, 1857 ; also Prof. Turner, in 'Journal
of Anat. and Phys.' Nov. 1, 1866, p. 78. Dr. Giinther has likewise
described similar cases.

Chap. VI. AFFINITIES AND GENEALOGY. 211

been known occasionally in man and other mammals to
become well developed, and to yield a fair supply of
milk. Now if we suppose that during a former pro-
longed period male mammals aided the females in
nursing their offspring, and that afterwards from some
cause, as from a smaller number of young being pro-
duced, the males ceased giving this aid, disuse of the
organs during maturity would lead to their becoming
inactive ; and from two well-known principles of in-
heritance this state of inactivity would probably be
transmitted to the males at the corresponding age of
maturity. But at all earlier ages these organs would
be left unaffected, so that they would be equally well
developed in the young of both sexes.

Conclusion. — The best definition of advancement or
progress in the organic scale ever given, is that by
Von Baer; and this rests on the amount of differ-
entiation and specialisation of the several parts of
the same being, when arrived, as I should be inclined
to add, at maturity. Now as organisms have become
slowly adapted by means of natural selection for
diversified lines of life, their parts wrill have become,
from the advantage gained by the division of physio-
logical labour, more and more differentiated and spe-
cialised for various functions. The same part appears
often to have been modified first for one purpose, and
then long afterwards for some other and quite distinct
purpose ; and thus all the parts are rendered more and
more complex. But each organism will still retain the
general type of structure of the progenitor from which
it was aboriginally derived. In accordance with this
view it seems, if we turn to geological evidence, that
organisation on the whole has advanced throughout the
world by slow and interrupted steps. In the great

p 2

212 THE DESCENT OF MAN. Part I.

kingdom of the Vertebrata it has culminated in man.
It must not, however, be supposed that groups of organic
beings are always supplanted and disappear as soon as
they have given birth to other and more perfect groups.
The latter, though victorious over their predecessors,
may not have become better adapted for all places in
the economy of nature. Some old forms appear to have
survived from inhabiting protected sites, where they
have not been exposed to very severe competition ; and
these often aid us in constructing our genealogies,
by giving us a fair idea of former and lost populations.
But we must not fall into the error of looking at the
existing members of any lowly-organised group as per-
fect representatives of their ancient predecessors.

The most ancient progenitors in the kingdom of the
Vertebrata, at which we are able to obtain an obscure
glance, apparently consisted of a group of marine animals,27
resembling the larvae of existing Ascidians. These
animals probably gave rise to a group of fishes, as lowly
organised as the lancelet ; and from these the Ganoids,
and other fishes like the Lepidosiren, must have been
developed. From such fish a very small advance would

27 All vital functions tend to run their course in fixed and recurrent
periods, and with tidal animals the periods would probably be lunar ;
for such animals must have been left dry or covered deep with water, —
supplied with copious food or stinted, — during endless generations, at
regular lunar intervals. If then the Vertebrata are descended from an
animal allied to the existing tidal Ascidians, the mysterious fact, that
with the higher and now terrestrial Vertebrata, not to mention other
classes, many normal and abnormal vital processes run their course
according to lunar periods, is rendered intelligible. A recurrent period,
if approximately of the right duration, when once gained, would not, as
far as we can judge, be liable to be changed ; consequently it might
be thus transmitted during almost any number of generations. This
conclusion, if it could be proved sound, would be curious ; for we should
then see that the period of gestation in each mammal, and the hatching
of each bird's eggs, and many other vital processes, still betrayed the
primordial birthplace of these animals.

Chap. VI. AFFINITIES AND GENEALOGY. 213

carry us on to the amphibians. We have seen that
birds and reptiles were once intimately connected
together ; and the Monotremata now, in a slight degree,
connect mammals with reptiles. But no one can at
present say by what line of descent the three higher
and related classes, namely, mammals, birds, and rep-
tiles, were derived from either of the two lower verte-
brate classes, namely amphibians and fishes. In the
class of mammals the steps are not difficult to con-
ceive which led from the ancient Monotremata to
tne ancient Marsupials ; and from these to the early
progenitors of the placental mammals. We may thus
ascend to the Lemuridse ; and the interval is not wide
from these to the Simiadse. The Simiadae then branched
off into two great stems, the New World and Old World
monkeys ; and from the latter, at a remote period, Man,
the wonder and glory of the Universe, proceeded.

Thus we have given to man a pedigree of prodigious
length, but not, it may be said, of noble quality. The
world, it has often been remarked, appears as if it had
long been preparing for the advent of man ; and this, in
one sense is strictly true, for he owes his birth to a long
line of progenitors. If any single link in this chain
had never existed, man would not have been exactly
what he now is. Unless we wilfully close our eyes, we
may, with our present knowledge, approximately re-
cognise our parentage ; nor need we feel ashamed of it.
The most humble organism is something much higher
than the inorganic dust under -our feet ; and no one
with an unbiassed mind can study any living creature,
however humble, without being struck with enthusiasm
at its marvellous structure and properties.

214 THE DESCENT OF MAN, Part I.

CHAPTEK VII.

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 — Mono-
genists 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 ex-
tinction of races — The formation of races — The effects of cross-
ing — Slight influence of the direct action of the conditions of life
Slight or no influence of natural selection— Sexual selection.

*n

*&

It is not my intention here to describe the several
so-called races of men; but to inquire what is the
value of the differences between them under a classi-
ficatory point of view, and how they have originated.
In determining whether two or more allied forms
ought to be ranked as species or varieties, natu-
ralists are practically guided by the following con-
siderations ; namely, the amount of difference between
them, and whether such differences relate to few or
many points of structure, and whether they are of
physiological importance ; but more especially whether
they are constant. Constancy of character is what is
chiefly valued and sought for by naturalists. Whenever
it can be shewn, or rendered probable, that the forms
in question have remained distinct for a long period,
this becomes an argument of much weight in favour
of treating them as species. Even a slight degree of
sterility between any two forms when first crossed, or
in their offspring, is generally considered as a decisive

Chap. VII. THE RACES OF MAN. 215

test of tlieir specific distinctness ; and their continued
persistence without blending within the same area, is
usually accepted as sufficient evidence, either of some
degree of mutual sterility, or in the case of animals of
some repugnance to mutual pairing.

Independently of blending from intercrossing, the
complete absence, in a well-investigated region, of
varieties linking together any two closely-allied forms,
is probably the most important of all the criterions
of their specific distinctness; and this is a somewhat
different consideration from mere constancy of character,
for two forms may be highly variable and yet not
yield intermediate varieties. Geographical distribution
is often unconsciously and sometimes consciously brought
into play ; so that forms living in two widely separated
areas, in which most of the other inhabitants are speci-
fically distinct, are themselves usually looked at as dis-
tinct ; but in truth this affords no aid in distinguishing
geographical races from so-called good or true species.

Now let us apply these generally-admitted principles
to the races of man, viewing him in the same spirit as
a naturalist would any other animal. In regard to the
amount of difference between the races, we must make
some allowance for our nice powers of discrimination
gained by the long habit of observing ourselves. In
India, as Elphinstone remarks,1 although a newly-arrived
European cannot at first distinguish the various native
races, yet they soon appear to him extremely dissimilar ;
and the Hindoo cannot at first perceive any difference
between the several European nations. Even the most
distinct races of man, with the exception of certain
negro tribes, are much more like each other in form

1 ' History of India,' 1841, vol. i. p. 323. Father Ripa makes exactly
tiie tame remark with respect to the Chinese.

216 THE DESCENT OF MAN. Part I.

than would at first be supposed. This is well shewn by
the French photographs in the Collection Anthropolo-
gique du Museum of the men belonging to various
races, the greater number of which, as many persons
to whom I have shown them have remarked, mioht
pass for Europeans. Nevertheless, these men if seen
alive would undoubtedly appear very distinct, so that
we are clearly much influenced in our judgment by
the mere colour of the skin and hair, by slight differ-
ences in the features, and by expression.

There is, however, no doubt that the various races,
when carefully compared and measured, differ much
from each other, — as in the texture of the hair, the
relative proportions of all parts of the body,2 the capa-
city of the lungs, the form and capacity of the skull,
and even in the convolutions of the brain.3 But it
would be an endless task to specify the numerous points
of structural difference. The races differ also in con-
stitution, in acclimatisation, and in liability to certain
diseases. Their mental characteristics are likewise very
distinct ; chiefly as it would appear in their emotional,
but partly in their intellectual, faculties. Every one
who has had the opportunity of comparison, must have
been struck with the contrast between the taciturn,
even morose, aborigines of S. America and the light-
hearted, talkative negroes. There is a nearly similar
contrast between the Malays and the Papuans,4 who live

2 A vast number of measurements of Whites, Blacks, and Indians, are
given in the ' Investigations in the Military and Anthropolog. Statistics
of American Soldiers,' by B. A. Gould, 1869, p. 298-358 ; on the
capacity of the lungs, p. 471. See also the numerous and valuable
tables, by Dr. Weisbach, from the observations of Dr. Scherzer and
Dr. Schwarz, in the 'Beise der Novara: Anthropolog. Theil,' 1867.

* See, for instance, Mr. Marshall's account of the brain of a Bush-
woman, in ' Bhil. Transact.' 1861, p. 519.

4 Wallace, ' The Malay Archipelago,' vol. ii. 1869, p. 178.

Chap. VII. THE EACES OF MAN. 217

under the same physical conditions, and are separated
from each other only by a narrow space of sea.

We -will first consider the arguments wdiich may be
advanced in favour of classing the races of man as
distinct species, and then those on the other side. If a
naturalist, who had never before seen such beings, were
to compare a Negro, Hottentot, Australian, or Mongolian,
he would at once perceive that they differed in a multi-
tude of characters, some of slight and some of consider-
able importance. On inquiry he would find that they
were adapted to live under widely different climates, and
that they differed somewhat in bodily constitution and
mental disposition. If he were then told that hundreds
of similar specimens could be brought from the same
countries, he would assuredly declare that they were as
good species as many to which he had been in the
habit of affixing specific names. This conclusion would
be greatly strengthened as soon as he had ascertained
that these forms had all retained the same character for
many centuries ; and that negroes, apparently identical
with existing negroes, had lived at least 4000 years
ago.5 He would also hear from an excellent observer,

5 With respect to the figures in the famous Egyptian caves of Abou-
Simbel, M. Pouchet says (' The Plurality of the Human Kaces,' Eng.
translat. 1861, p. 50), that he was far from finding recognisable repre-
sentations of the dozen or more nations which some authors believe
that they can recognise. Even some of the most strongly-marked
races cannot be identified witli that degree of unanimity which might
have been expected from what has been written on the subject.
Thus Messrs. Nott and Gliddon (' Types of Mankind,' p. 148) state
that Rameses II., or the Great, has features superbly European; where-
as Knox, another firm believer in the specific distinction of the races
of man ('Races of Man,' 1850, p. 201), speaking of young Memnon
(the same person with Rameses II., as I am infurmedby Mr. Birch) in-
sists in the strongest manner that he is identical in character with the
Jews of Antwerp. Again, whilst looking in the British Museum with
two competent judges, officers of the establishment, at the statue of
Amunoph III., we agreed that he had a strongly negro cast of features ;

218 THE DESCENT OF MAN. Pabt I.

Dr. Lund,6 that the human skulls found in the caves of
Brazil, entombed with many extinct mammals, belonged
to the same type as that now prevailing throughout the
American Continent.

Our naturalist would then perhaps turn to geogra-
phical distribution, and he would probably declare that
forms differing not only in appearance, but fitted for
the hottest and dampest or driest countries, as well as
for the arctic regions, must be distinct species. He
might appeal to the fact that no one species in the
group next to man, namely the Quaclrumana, can resist
a low temperature or any considerable change of climate ;
and that those species which come nearest to man have
never been reared to maturity, even under the temperate
climate of Europe. He would be deeply impressed with
the fact, first noticed by Agassiz,7 that the different
races of man are distributed over the world in the same
zoological provinces, as those inhabited by undoubtedly
distinct species and genera of mammals. This is mani-
festly the case with the Australian, Mongolian, and
Negro races of man ; in a less well-marked manner with
the Hottentots ; but plainly with the Papuans and
Malays, who are separated, as Mr. Wallace has shewn,
by nearly the same line which divides the great Malayan
and Australian zoological provinces. The aborigines
of America range throughout the Continent ; and this
at first appears opposed to the above rule, for most of
the productions of the Southern and Northern halves
differ widely ; yet some few living forms, as the

but Messrs. Nott and Gliddon (ibid. p. 146, fig. 53) describe him as
" a hybrid, but not of negro intermixture."

6 As quoted by Nott and Gliddon, « Types of Mankind,' 1854, p. 439.
They give also corroborative evidence ; but C. Vogt thinks that the
subject requires further investigation.

7 " Diversity of Origin of the Human Kaces," in the ' Christian
Examiner,' July, 1850.

Chap. VII. THE EACES OF MAN. 219

opossum, range from the one into the other, as did
formerly some of the gigantic Edentata. The Esqui-
maux, like other Arctic animals, extend round the
whole polar regions. It should be observed that the
mammalian forms which inhabit the several zoological
provinces, do not differ from each other in the same
degree; so that it can hardly be considered as an
anomaly that the Negro differs more, and the American
much less, from the other races of man than do the
mammals of the same continents from those of the other
provinces. Man, it may be added, does not appear to
have aboriginally inhabited any oceanic island ; and in
this respect he resembles the other members of his class.
In determining whether the varieties of the same
kind of domestic animal should be ranked as specifically
distinct, that is, whether any of them are descended from
distinct wild species, every naturalist would lay much
stress on the fact, if established, of their external parasites
being specifically distinct. All the more stress would be
laid on this fact, as it would be an exceptional one, for
I am informed by Mr. Denny that the most different
kinds of dogs, fowls, and pigeons, in England, are infested
by the same species of Pediculi or lice. Now Mr. A.
Murray has carefully examined the Pediculi collected in
different countries from the different races of man ; 8
and he finds that they differ, not only in colour, but
in the structure of their claws and limbs. In every
case in which numerous specimens were obtained the
differences were constant. The surgeon of a whaling
ship in the Pacific assured me that^ when the Pediculi,
with which some Sandwich Islanders on board swarmed,
strayed on to the bodies of the English sailors, they
died in the course of three or four clays. These Pediculi

8 'Transact. E. Soc. of Edinburgh,' vol. xxii. 1861. p. 567.

220 THE DESCENT OF MAN. Part I.

were darker coloured and appeared different from those
proper to the natives of Chiloe in South America, of
which he gave me specimens. These, again, appeared
larger and much softer than European lice. Mr. Murray
procured four kinds from Africa, namely from the
Negroes of the Eastern and Western coasts, from the
Hottentots and Caffres ; two kinds from the natives
of Australia ; two from North, and two from South
America. In these latter cases it may be presumed
that the Pediculi came from natives inhabiting different
districts. With insects slight structural differences, if
constant, are generally esteemed of specific value : and
the fact of the races of man being infested by parasites,
which appear to be specifically distinct, might fairly
be urged as an argument that the races themselves
ought to be classed as distinct species.

Our supposed naturalist having proceeded thus far
in his investigation, would next inquire whether the
races of men, when crossed, were in any degree sterile.
He might consult the work 9 of a cautious and philo-
sophical observer, Professor Broca ; and in this he would
find good evidence that some races were quite fertile
together ; but evidence of an opposite nature in regard to
other races. Thus it has been asserted that the native
women of Australia and Tasmania rarely produce
children to European men ; the evidence, however, on
this head has now been shewn to be almost valueless.
The half-castes are killed by the pure blacks ; and an
account has lately been published of eleven half-caste
youths murdered and burnt at the same time, whose
remains were found by the police.10 Again, it has often

9 'On the Phenomena of Hybridity in the Genus Homo,' Eng.
translat. 1864.

10 See the interesting letter by Mr. T. A. Murray, in the ' Anthro-
polog. Keview,' April, 1868, p. liii. In this letter Count Strzeleeki's

Chap. VII. THE KACES OF MAN. 221

been said that when mulattoes intermarry they produce
few children ; on the other hand, Dr. Bachman of
Charlestown u positively asserts that he has known
mulatto families which have intermarried for several
generations, and have continued on an average as fertile
as either pure whites or pure blacks. Inquiries formerly
made by Sir C. Lyell on this subject led him, as he
informs me, to the same conclusion. In the United
States the census for the year 1854 included, according
to Dr. Bachman, 405,751 mulattoes ; and this number,
considering all the circumstances of the case, seems
small ; but it may partly be accounted for by the de-
graded and anomalous position of the class, and by the
profligacy of the women. A certain amount of absorp-
tion of mulattoes into negroes must always be in pro-
gress ; and this would lead to an apparent diminution
of the former. The inferior vitality of mulattoes is
spoken of in a trustworthy work12 as a well-known
phenomenon ; but this is a different consideration from
their lessened fertility ; and can hardly be advanced as
a proof of the specific distinctness of the parent races.
No doubt both animal and vegetable hybrids, when
produced from extremely distinct species, are liable to
premature death ; but the parents of mulattoes cannot
be put under the category of extremely distinct species.
The common Mule, so notorious for long life and vigour,
and yet so sterile, shews how little necessary connection

statement, that Australian women who have borne children to a white
man are afterwards sterile with their own race^ is disproved. M. A. de
Quatrefages has also collected (' Eevue des Cours Scientifiques,' March,
1869, p. 239) much evidence that Australians and Europeans are not
sterile when crossed.

11 ' An Examination of Prof. Agassiz's Sketch of the Nat. Provinces
of the Animal World,' Charleston, 1855, p. 44.

12 'Military and Anthropolog. Statistics of American Soldiers,' by
B. A. Gould, 18G9, p. 319.

222 THE DESCENT OF MAN. Part I.

there is in hybrids between lessened fertility and vitality :
other analogous cases could be added.

Even if it should hereafter be proved that all the
races of men were perfectly fertile together, he who was
inclined from other reasons to rank them as distinct
species, might with justice argue that fertility and
sterility are not safe criterions of specific distinctness.
We know that these qualities are easily affected by
changed conditions of life or by close inter-breeding,
and that they are governed by highly complex laws, for
instance that of the unequal fertility of reciprocal crosses
between the same two species. With forms which must
be ranked as undoubted species, a perfect series exists
from those which are absolutely sterile when crossed,
to those which are almost or quite fertile. The degrees
of sterility do not coincide strictly with the degrees of
difference in external structure or habits of life. Man
in many respects may be compared with those animals
which have long been domesticated, and a large body
of evidence can be advanced in favour of the Pallasian
doctrine13 that domestication tends to eliminate the

13 ' The Variation of Animals and Plants under Domestication,' vol. ii.
p. 109. I may here remind the reader that the sterility of species
when crossed is not a specially-acquired quality ; but, like the inca-
pacity of certain trees to be grafted together, is incidental on other
acquired differences. The nature of these differences is unknown, but
they relate more especially to the reproductive system, and much less
to external structure or to ordinary differences in constitution. One
important element in the sterility of crossed species apparently lies in
one or both having been long habituated to fixed conditions ; for we
know that changed conditions have a special influence on the repro-
ductive system, and we have good reason to believe (as before re-
marked) that the fluctuating conditions of domestication tend to elimi-
nate that sterility which is so general with species in a natural state
when crossed. It has elsewhere been shewn by me (ibid. vol. ii. p. 185,
and' ' Origin of Species,' oth edit. p. 317) that the sterility of crossed
species has not been acquired through natural selection : we can see that
when two forms have already been rendered very sterile, it is scarcely

Chap. VII. THE EACES OF MAN. 223

sterility which is so general a result of the crossing of
species in a state of nature. From these several con-
siderations, it may be justly urged that the perfect ferti-
lity of the intercrossed races of man, if established,
would not absolutely preclude us from ranking them as
distinct species.

Independently of fertility, the character of the off-
spring from a cross has sometimes been thought to
afford evidence whether the parent-forms ought to be
ranked as species or varieties ; but after carefully study-
ing the evidence, I have come to the conclusion that no
general rules of this kind can be trusted. Thus with
mankind the offspring of distinct races resemble in all
respects the offspring of true species and of varieties.
This is shewn, for instance, by the manner in which
the characters of both parents are blended, and by
one form absorbing another through repeated crosses.
In this latter case the progeny both of crossed species
and varieties retain for a long period a tendency to
revert to their ancestors, especially to that one which
is prepotent in transmission. When any character has
suddenly appeared in a race or species as the result of a

possible that their sterility should he augmented by the preservation or
survival of the more and more sterile individuals ; for as the sterility
increases fewer and fewer offspring will be produced i'rom which to
breed, and at last only single individuals will be produced, at the rarest
intervals. But there is even a higher grade of sterility than this. Both
Gartner and Kolreuter have proved that in genera of plants including
numerous species, a series can be formed from species which when
crossed yield fewer and fewer seeds, to species which never produce a
single seed, but yet are affected by the pollen of the other species, for
the germen swells. It is here manifestly impossible to select the more
sterile individuals, which have already ceased to yield seeds ; so tW_ . ^
the acme of sterility, wheu the germen alone is affected, caunpif*ne\ P A >
gained through selection. This acme, and no doubt the olhei^i\&es
of sterility, are the incidental results of certain unknowm d^^iicO*^' ' ^<
in the constitution of the reproductive system of the species/ "wiucKarj^^.,-^
crossed. / ^0

Lr LIBRAE

224 THE DESCENT OF MAN. Part I.

single act of variation, as is general with monstrosities,14
and this race is crossed with another not thus charac-
terised, the characters in question do not commonly
appear in a blended condition in the young, but are
transmitted to them either perfectly developed or not at
all. As with the crossed races of man cases of this kind
rarely or never occur, this may be used as an argument
against the view suggested by some ethnologists, namely
that certain characters, for instance the blackness of the
negro, first appeared as a sudden variation or sport.
Had this occurred, it is probable that mulattoes would
often have been born, either completely black or com-
pletely white.

We have now seen that a naturalist might feel him-
self fully justified in ranking the races of man as distinct
species ; for he has found that they are distinguished by
many differences iu structure and constitution, some
being of importance. These differences have, also, re-
mained nearly constant for very long periods of time.
He will have been in some degree influenced by the
enormous range of man, which is a great anomaly in
the class of mammals, if mankind be viewed as a single
species. He will have been struck with the distribution
of the several so-called races, in accordance with that
of other undoubtedly distinct species of mammals.
Finally he might urge that the mutual fertility of all
the races has not as yet been fully proved ; and even if
proved would not be an absolute proof of their specific
identity.

On the other side of the question, if our supposed
naturalist were to enquire whether the forms of man
kept distinct like ordinary species, when mingled to-

14 ' The Variation of Animals,' &c, vol. ii. p. 92.

Chap. VII. THE EACES OF MAN. 225

gether in large numbers in the same country, lie would
immediately discover that this was by no means the
case. In Brazil he would behold an immense mongrel
population of Negroes and Portuguese ; in Chiloe and
other parts of South America, he would behold the
whole population consisting of Indians and Spaniards
blended in various degrees.15 In many parts of the
same continent he would meet with the most complex
crosses between Negroes, Indians, and Europeans ; and
such triple crosses afford the severest test, judging from
the vegetable kingdom, of the mutual fertility of the
parent-forms. In one island of the Pacific he would
find a small population of mingled Polynesian and
English blood ; and in the Viti Archipelago a popula-
tion of Polynesians and Negritos crossed in all degrees.
Many analogous cases could be added, for instance, in
South Africa. Hence the races of man are not suffi-
ciently distinct to co-exist without fusion ; and this it
is, which in all ordinary cases affords the usual test of
specific distinctness.

Our naturalist would likewise be much disturbed as
soon as he perceived that the distinctive characters of
every race of man were highly variable. This strikes
every one when he first beholds the negro-slaves in
Brazil, who have been imported from all parts of Africa.
The same remark holds good with the Polynesians, and
with many other races. It may be doubted whether
any character can be named which is distinctive of a
race and is constant. Savages, even within the limits of
the same tribe, are not nearly so uniform in character,
as has often been said. Hottentot women offer certain

15 M. de Quatrefages has given (' Anthropolog. Eeview,' Jan. 1860,
p. 22) an interesting account of the success and energy of the Paulistas
in Brazil, who are a much crossed race of Portuguese and Indians, with
a mixture of the blood of other races.

VOL. I. Q

226 THE DESCENT OF MAN. Part I.

peculiarities, more strongly marked than those occur-
ring in any other race, but these are known not to
be of constant occurrence. In the several American
tribes, colour and hairyness differ considerably ; as does
colour to a certain degree, and the shape of the features
greatly, in the Negroes of Africa. The shape of the
skull varies , much in some races ; 16 and so it is with
every other character. Now all naturalists have learnt
by dearly-bought experience, how rash it is to attempt
to define species by the aid of inconstant characters.

But the most weighty of all the arguments against
treating the races of man as distinct species, is that they
graduate into each other, independently in many cases,
as far as we can judge, of their having intercrossed.
Man has been studied more carefully than any other
organic being, and yet there is the greatest possible
diversity amongst capable judges whether he should be
classed as a single species or race, or as two (Virey), as
three (Jacquinot), as four (Kant), five (Blumenbach),
six (Buffon), seven (Hunter), eight (Agassiz), eleven
(Pickering), fifteen (Bory St. Vincent), sixteen (Des-
moulins), twenty- two (Morton), sixty (Crawfurd), or as
sixty-three, according to Burke.17 This diversity of
judgment does not prove that the races ought not to be
ranked as species, but it shews that they graduate into
each other, and that it is hardly possible to discover
clear distinctive characters between them.

Every naturalist who has had the misfortune to under-

16 For instance with the aborigines of America and Australia. Prof.
Huxley says (' Transact. Internat. Congress of Prehist. Arch.' 1868, p.
105) that the skulls of many South Germans and Swiss are " as short
" and as broad as those of the Tartars," &c.

17 See a good discussion on this subject in Waitz, ' Introduct. to
Anthropology,' Eng. translat. 1863, p. 198-208, 227. I have taken
some of the above statements from H. Tuttle's ' Origin and Antiquity
of Physical Man,' Boston, 1866, p. 35.

Chap. VII. THE RACES OF MAN. 227

take the description of a group of highly varying
organisms, has encountered cases (I speak after ex-
perience) precisely like that of man ; and if of a cautious
disposition, he will end by uniting all the forms which
graduate into each other as a single species ; for he will
say to himself that he has no right to give names to
objects which he cannot define. Cases of this kind occur
in the Order which includes man, namely in certain
genera of monkeys ; whilst in other genera, as in Cerco-
pithecus, most of the species can be determined with
certainty. In the American genus Cebus, the various
forms are ranked by some naturalists as species, by
others as mere geographical races. Now if numerous
specimens of Cebus were collected from all parts of
South America, and those forms which at present ap-
pear to be specifically distinct, were found to graduate
into each other by close steps, they would be ranked by
most naturalists as mere varieties or races ; and thus the
greater number of naturalists have acted with respect
to the races of man. Nevertheless it must be confessed
that there are forms, at least in the vegetable king-
dom,18 which we cannot avoid naming as species, but
which are connected together, independently of inter-
crossing, by numberless gradations.

Some naturalists have lately employed the term
" sub-species " to designate forms which possess many of
the characteristics of true species, but which hardly de-
serve so high a rank. Now if we reflect on the weighty
arguments, above given, for raising the races of man to
the dignity of species, and the insuperable difficulties
on the other side in defining them, the term " sub-

18 Prof. N'ageli has carefully described several striking cases in his
' Botanische Mittheilungen,' B. ii. 1866, s. 294-369. Prof. Asa Gray
has made analogous remarks on some intermediate forms in the Com-
posite of N. America.

Q 2

228 THE DESCENT OF MAN. Part I.

species " might here be used with much propriety. But
from long habit the term " race " will perhaps always
be employed. The choice of terms is only so far im-
portant as it is highly desirable to use, as far as that
may be possible, the same terms for the same degrees of
difference. Unfortunately this is rarely possible ; for
within the same family the larger genera generally
include closely-allied forms, which can be distinguished
only with much difficulty, whilst the smaller genera
include forms that are perfectly distinct ; yet all must
equally be ranked as species. So again the species
within the same large genus by no means resemble
each other to the same degree : on the contrary, in
most cases some of them can be arranged in little groups
round other species, like satellites round planets.19

The question whether mankind consists of one or
several species has of late years been much agitated by
anthropologists, who are divided into two schools of
monogenists and polygenists. Those who do not admit
the principle of evolution, must look at species either
as separate creations or as in some manner distinct
entities ; and they must decide what forms to rank as
species by the analogy of other organic beings which
are commonly thus received. But it is a hopeless en-
deavour to decide this point on sound grounds, until
some definition of the term " species " is generally ac-
cepted ; and the definition must not include an element
which cannot possibly be ascertained, such as an act of
creation. We might as well attempt without any defi-
nition to decide whether a certain number of houses
should be called a village, or town, or city. We have a
practical illustration of the difficulty in the never-

19

' Origin of Species,' 5th edit. p. 68.

Chap. VII. THE RACES OF MAN. 229

ending donbts whether many closely-allied mammals,
birds, insects, and plants, which represent each other in
North America and Europe, should be ranked species
or geographical races ; and so it is with the productions
of many islands situated at some little distance from the
nearest continent.

Those naturalists, on the other hand, who admit the
principle of evolution, and this is now admitted by the
greater number of rising men, will feel no doubt that
all the races of man are descended from a single primi-
tive stock ; whether or not they think fit to designate
them as distinct species, for the sake of expressing their
amount of difference.20 With our domestic animals the
question whether the various races have arisen from
one or more species is different. Although all such
races, as well as all the natural species within the same
genus, have undoubtedly sprung from the same primi-
tive stock, yet it is a fit subject for discussion, whether,
for instance, all the domestic races of the dog have
acquired their present differences since some one species
was first domesticated and bred by man ; or whether they
owe some of their characters to inheritance from distinct
species, which had already been modified in a state of
nature. With mankind no such question can arise, for
he cannot be said to have been domesticated at any
particular period.

When the races of man diverged at an extremely
remote epoch from their common progenitor, they will
have differed but little from each other, and been few
in number ; consequently they will then, as far as their
distinguishing characters are concerned, have had less
claim to rank as distinct species, than the existing so-

20 See Prof. Huxley to this effect in the ' Fortnightly Review,' 1865,
p. 275.

230 THE DESCENT OF MAN. Part I.

called races. Nevertheless such early races would per-
haps have been ranked by some naturalists as distinct
species, so arbitrary is the term, if their differences,
although extremely slight, had been more constant than
at present, and had not graduated into each other.

It is, however, possible, though far from probable,
that the early progenitors of man might at first have
diverged much in character, until they became more
unlike each other than are any existing races ; but that
subsequently, as suggested by Vogt,21 they converged
in character. When man selects for the same object
the offspring of two distinct species, he sometimes
induces, as far as general appearance is concerned,
a considerable amount of convergence. This is the
case, as shewn by Von Nathusius,22 with the improved
breeds of pigs, which are descended from two distinct
species ; and in a less well-marked manner with the
improved breeds of cattle. A great anatomist, Gratiolet,
maintains that the anthropomorphous apes do not form
a natural sub-group ; but that the orang is a highly
developed gibbon or semnopithecus ; the chimpanzee
a highly developed macacus ; and the gorilla a highly
developed mandrill. If this conclusion, which rests
almost exclusively on brain-characters, be admitted,
we should have a case of convergence at least in
external characters, for the anthropomorphous apes are
certainly more like each other in many points than
they are to other apes. All analogical resemblances,
as of a whale to a fish, may indeed be said to be
cases of convergence ; but this term has never been
applied to superficial and adaptive resemblances. It

21 ' Lectures on Man,' Eng, translat. 1864, p. 468.

22 'Die Kacen des Schweines,' 1860, s. 46. 'Vorstudien fur Ge-
schichte, &c, Schwemeschadel/ 1864, s. 104. With respect to cattle,
tee M. de Quatrefages, ' Unite' de l'Espece Humaine,' 1861, p. 119.

Chap. VII. THE KACES OF MAN. 231

would be extremely rash in most cases to attribute to
convergence close similarity in many points of struc-
ture in beings which had once been widely different.
The form of a crystal is determined solely by the
molecular forces, and it is not surprising that dissimilar
substances should sometimes assume the same form;
but with organic beings we should bear in mind that
the form of each depends on an infinitude of complex
relations, namely on the variations which have arisen,
these being due to causes far too intricate to be followed
out, — on the nature of the variations which have been
preserved, and this depends on the surrounding physical
conditions, and in a still higher degree on the sur-
rounding organisms with which each has come into
competition, — and lastly, on inheritance (in itself a
fluctuating element) from innumerable progenitors, all
of which have had their forms determined through
equally complex relations. It appears utterly incredible
that two organisms, if differing in a marked manner,
should ever afterwards converge so closely as to lead
to a near approach to identity throughout their whole
organisation. In the case of the convergent races of pigs
above referred to, evidence of their descent from two pri-
mitive stocks is still plainly retained, according to Von
Nathusius, in certain bones of their skulls. If the races
of man were descended, as supposed by some naturalists,
from two or more distinct species, which had differed as
much, or nearly as much, from each other, as the orang
differs from the gorilla, it can hardly be doubted that
marked differences in the structure of certain bones would
still have been discoverable in man as he now exists.

Although the existing races of man differ in many
respects, as in colour, hair, shape of skull, proportions
of the body, &c, yet if their whole organisation be taken

232 THE DESCENT OF MAN. Part I.

into consideration they are found to resemble each
other closely in a multitude of points. Many of these
points are of so unimportant or of so singular a nature,
that it is extremely improbable that they should have
been independently acquired by aboriginally distinct
species or races. The same remark holds good with
equal or greater force with respect to the numerous
points of mental similarity between the most distinct
races of man. The American aborigines, Negroes and
Europeans differ as much from each other in mind as any
three races that can be named ; yet I was incessantly
struck, whilst living with the Fuegians on board the
" Beagle," with the many little traits of character,
shewing how similar their minds were to ours ; and so
it was with a full-blooded negro with whom I happened
once to be intimate.

He who will carefully read Mr. Tylor's and Sir
J. Lubbock's interesting works 23 can hardly fail to be
deeply impressed with the close similarity between
the men of all races in tastes, dispositions and habits.
This is shewn by the pleasure which they all take in
dancing, rude music, acting, painting, tattooing, and
otherwise decorating themselves, — in their mutual
comprehension of gesture-language — and, as I shall be
able to shew in a future essay, by the same expression
in their features, and by the same inarticulate cries,
when they are excited by various emotions. This
similarity, or rather identity, is striking, when contrasted
with the different expressions which may be observed
in distinct species of monkeys. There is good evi-
dence that the art of shooting with bows and arrows has
not been handed down from any common progenitor of

23 Tylor's « Early History of Mankind,' 1865 ; for the evidence with
respect to gesture-language, see p. 54. Lubbock's • Prehistoric Times,'
2nd edit. 1869.

Chap. VII. THE KACES OF MAN. 233

mankind, yet the stone arrow-heads, brought from the
most distant parts of the world and manufactured at the
most remote periods, are, as Mlsson has shewn,24 almost
identical ; and this fact can only be accounted for by
the various races having similar inventive or mental
powers. The same observation has been made by
archaeologists 25 with respect to certain widely-prevalent
ornaments, such as zigzags, &c; and with respect to
various simple beliefs and customs, such as the burying
of the dead under megalithic structures. I remember
observing in South America,26 that there, as in so many
other parts of the world, man has generally chosen the
summits of lofty hills, on which to throw up piles of
stones, either for the sake of recording some remarkable
event, or for burying his dead.

Now when naturalists observe a close agreement in
numerous small details of habits, tastes and dispositions
between two or more domestic races, or between nearly-
allied natural forms, they use this fact as an argument
that all are descended from a common progenitor who
was thus endowed ; and consequently that all should be
classed under the same species. The same argument
may be applied with much force to the races of man.

As it is improbable that the numerous and unim-
portant points of resemblance between the several races
of man in bodily structure and mental faculties (I do
not here refer to similar customs) should all have been
independently acquired, they must have been inherited
from progenitors who were thus characterised. We
thus gain some insight into the early state of man,

34 ' The Primitive Inhabitants of Scandinavia/ Eng. translaj^mAJQ A j
by Sir J. Lubbock, 1868, p. 104. /V ~* ,

25 Hodder M. Westropp, on Cromlechs, &c, ' JournajAf^Et]jliP-S **<v
logical Soc' as given in 'Scientific Opinion,' June 2nd, 18^9£p~. 2r ^^.z*** «

26 ' Journal of Kesearches : Voyage of the " Beagle," ' n.39. n ,

1. 1. LIBRA*

234 THE DESCENT OF MAN. Part I.

before he had spread step by step over the face of the
earth. The spreading of man to regions widely sepa-
rated by the sea, no doubt, preceded any considerable
amount of divergence of character in the several races ;
for otherwise we should sometimes meet with the same
race in distinct continents ; and this is never the case.
Sir J. Lubbock, after comparing the arts now practised
by savages in all parts of the world, specifies those
which man could not have known, when he first wan-
dered from his original birth-place ; for if once learnt
they would never have been forgotten. 27 He thus shews
that "the spear, which is but a development of the
" knife-point, and the club, which is but a long hammer,
" are the only things left." He admits, however, that
the art of making fire probably had already been dis-
covered, for it is common to all the races now existing,
and was known to the ancient cave-inhabitants of
Europe. Perhaps the art of making rude canoes or
rafts was likewise known ; but as man existed at a re-
mote epoch, when the land in many places stood at a
very different level, he would have been able, without
the aid of canoes, to have spread widely. Sir J. Lubbock
further remarks how improbable it is that our earliest
ancestors could have " counted as high as ten, consider-
" ing that so many races now in existence cannot get
" beyond four." Nevertheless, at this early period, the
intellectual and social faculties of man could hardly have
been inferior in any extreme degree to those now pos-
sessed by the lowest savages ; otherwise primeval man
could not have been so eminently successful in the
struggle for life, as proved by his early and wide
diffusion.

From the fundamental differences between certain

' Prehistoric Times,' 1869, p. 574.

Chap. VII. THE EACES OF MAN. 235

languages, some philologists have inferred that when
man first became widely diffused he was not a speaking
animal; but it may be suspected that languages, far
less perfect than any now spoken, aided by gestures,
might have been used, and yet have left no traces
on subsequent and more highly-developed tongues.
Without the use of some language, however imperfect,
it appears doubtful whether man's intellect could have
risen to the standard implied by his dominant position
at an early period.

Whether primeval man, when he possessed very few
arts of the rudest kind, and when his power of language
was extremely imperfect, would have deserved to be
called man, must depend on the definition which we
employ. In a series of forms graduating insensibly
from some ape-like creature to man as he now exists,
it would be impossible to fix on any definite point when
the term " man " ought to be used. But this is a matter
of very little importance. So again it is almost a
matter of indifference whether the so-called races of
man are thus designated, or are ranked as species
or sub-species; but the latter term appears the most
appropriate. Finally, we may conclude that when
the principles of evolution are generally accepted, as
they surely will be before long, the dispute between the
monogenists and the polygenists will die a silent and
unobserved death.

One other question ought not to be passed over
without notice, namely, whether, as is sometimes
assumed, each sub-species or race of man has sprung
from a single pair of progenitors. With our domestic
animals a new race can readily be formed from a single
pair possessing some new character, or even from a
single individual thus characterised, by carefully match-

236 THE DESCENT OF MAN. Part I.

ing the varying offspring ; but most of our races have
been formed, not intentionally from a selected pair,
but unconsciously by the preservation of many indi-
viduals which have varied, however slightly, in some
useful or desired manner. If in one country stronger
and heavier horses, and in another country lighter and
fleeter horses, were habitually preferred, we may feel
sure that two distinct sub-breeds would, in the course
of time, be produced, without any particular pairs
or individuals having been separated and bred from
in either country. Many races have been thus formed,
and their manner of formation is closely analogous with
that of natural species. We know, also, that the
horses which have been brought to the Falkland
Islands have become, during successive generations,
smaller and weaker, whilst those which have run wild
on the Pampas have acquired larger and coarser
heads; and such changes are manifestly due, not to
any one pair, but to all the individuals having been
subjected to the same conditions, aided, perhaps, by
the principle of reversion. The new sub-breeds in
none of these cases are descended from any single
pair, but from many individuals which have varied in
different degrees, but in the same general manner;
and we may conclude that the races of man have been
similarly produced, the modifications being either the
direct result of exposure to different conditions, or the
indirect result of some form of selection. But to this
latter subject we shall presently return.

On the Extinction of the Races of Man. — The partial
and complete extinction of many races and sub-races
of man are historically known events. Humboldt saw
in South America a parrot which was the sole living
creature that could speak the language of a lost tribe.

Chap. VII. THE KACES OF MAN. 237

Ancient monuments and stone implements found in
all parts of the world, of which no tradition is pre-
served by the present inhabitants, indicate much
extinction. Some small and broken tribes, remnants
of former races, still survive in isolated and gene-
rally mountainous districts. In Europe the ancient
races were all, according to Schaaffhausen,28 " lower in
" the scale than the rudest living savages ; " they must
therefore have differed, to a certain extent, from any
existing race. The remains described by Professor
Broca29 from Les Eyzies, though they unfortunately
appear to have belonged to a single family, indicate a
race with a most singular combination of low or simious
and high characteristics, and is "entirely different
" from any other race, ancient or modern, that we have
" ever heard of." It differed, therefore, from the qua-
ternary race of the caverns of Belgium.

Unfavourable physical conditions appear to have had
but little effect in the extinction of races.30 Man has
long lived in the extreme regions of the North, with
no wood wherewith to make his canoes or other imple-
ments, and with blubber alone for burning and giving
him warmth, but more especially for melting the snow.
In the Southern extremity of America the Fuegians
survive without the protection of clothes, or of any
building worthy to be called a hovel. In South Africa
the aborigines wander over the most arid plains, where
dangerous beasts abound. Man can withstand the
deadly influence of the Terai at the foot of the Hima-
laya, and the pestilential shores of tropical Africa.

58 Translation in ' Anthropological Eeview,' Oct. 1S68, p. 431.

29 ' Transact. Internat. Congress of Prehistoric Arch.' 1868, p. 172-
175. See also Broca (translation) in ' Anthropological Kcview,' Oct.
1868, p. 410.

30 Dr. Gerland ' Ueber das Aussterben der Natnrvolkcr/ 1868, s. 82.

238 THE DESCENT OF MAN. Part I.

Extinction follows chiefly from the competition of
tribe with tribe, and race with race. Various checks
are always in action, as specified in a former chapter,
which serve to keep down the numbers of each savage
tribe, — such as periodical famines, the wandering of
the parents and the consequent deaths of infants, pro-
longed suckling, the stealing of women, wars, accidents,
sickness, licentiousness, especially infanticide, and,
perhaps, lessened fertility from less nutritious food, and
many hardships. If from any cause any one of these
checks is lessened, even in a slight degree, the tribe
thus favoured will tend to increase ; and when one
of two adjoining tribes becomes more numerous and
powerful than the other, the contest is soon settled by
war, slaughter, cannibalism, slavery, and absorption.
Even when a weaker tribe is not thus abruptly swept
away, if it once begins to decrease, it generally goes on
decreasing until it is extinct.31

When civilised nations come into contact with bar-
barians the struggle is short, except where a deadly cli-
mate gives its aid to the native race. Of the causes which
lead to the victory of civilised nations, some are plain
and some very obscure. We can see that the cultivation
of the land will be fatal in many ways to savages, for
they cannot, or will not, change their habits. New
diseases and vices are highly destructive ; and it appears
that in every nation a new disease causes much death,
until those who are most susceptible to its destructive
influence are gradually weeded out ;32 and so it may be
with the evil effects from spirituous liquors, as well as
with the unconquerably strong taste for them shewn by
so many savages. It further appears, mysterious as is

31 Gerland (ibid. s. 12) gives facts in support of this statement.

32 See remarks to this effect in Sir H. Holland's ' Medical Notes and
Reflections,' 1839, p. 390.

Chap. VII. THE EACES OF MAN. 239

the fact, that the first meeting of distinct and separated
people generates disease.33 Mr. Sproat, who in Van-
couver Island closely attended to the subject of extinc-
tion, believes that changed habits of life, which always
follow from the advent of Europeans, induces much ill-
health. He lays, also, great stress on so trifling a cause
as that the natives become " bewildered and dull by the
" new life around them ; they lose the motives for exer-
" tion, and get no new ones in their place." M

The grade of civilisation seems a most important
element in the success of nations which come in compe-
tition. A few centuries ago Europe feared the inroads
of Eastern barbarians ; now, any such fear would be ridi-
culous. It is a more curious fact, that savages did not
formerly waste away, as Mr. Bagehot has remarked,
before the classical nations, as they now do before
modern civilised nations ; had they done so, the old
moralists would have mused over the event ; but there
is no lament in any writer of that period over the perish-
ing barbarians.35

Although the gradual decrease and final extinction
of the races of man is an obscure problem, we can see
that it depends on many causes, differing in different
places and at different times. It is the same difficult
problem as that presented by the extinction of one of
the higher animals — of the fossil horse, for instance,
which disappeared from South America, soon afterwards
to be replaced, within the same districts, by countless
troops of the Spanish horse. The New Zealander seems

33 1 have collected (; Journal of Besearches, Voyage of the " Beagle," '
p. 435) a good many oases bearing on this subject : see also Gerland,
ibid. s. 8. Poeppig speaks of the " breath of civilisation as poisonous
" to savages."

34 Sproat, ' Scenes and Studies of Savage Life,' 1868, p. 284.

35 Bagehot, " Physics and Politics," ' Fortnightly Beview,' April 1,
1868, p. 455.

240 THE DESCENT OF MAN. Part I.

conscious of this parallelism, for he compares his future
fate with that of the native rat almost exterminated by
the European rat. The difficulty, though great to our
imagination, and really great if we wish to ascertain
the precise causes, ought not to be so to our reason,
as long as we keep steadily in mind that the increase of
each species and each race is constantly hindered by
various checks ; so that if any new check, or cause of
destruction, even a slight one, be superadded, the race
will surely decrease in number ; and as it has every-
where been observed that savages are much opposed to
any change of habits, by which means injurious checks
could be counterbalanced, decreasing numbers will
sooner or later lead to extinction ; the end, in most
cases, being promptly determined by the inroads of
increasing and conquering tribes.

On the Formation of the Baces of Man. — It may be
premised that when we find the same race, though
broken up into distinct tribes, ranging over a great
area, as over America, we may attribute their general
resemblance to descent from a common stock. In some
cases the crossing of races already distinct has led to
the formation of new races. The singular fact that
Europeans and Hindoos, who belong to the same Aryan
stock and speak a language fundamentally the same,
differ widely in appearance, whilst Europeans differ
but little from Jews, who belong to the Semitic stock
and speak quite another language, has been accounted
for by Broca36 through the Aryan branches having been
largely crossed during their wide diffusion by various
indigenous tribes. When two races in close contact

36 " On Anthropology," translation, ' Anthropolog. Eeview,' Jan.
1868, p. 38.

Chap. VIL THE RACES OF MAN. 241

cross, the first result is a heterogeneous mixture :
thus Mr. Hunter, in describing the Santali or hill-
tribes of India, says that hundreds of imperceptible
gradations may be traced " from the black, squat tribes
" of the mountains to the tall olive-coloured Brahman,
" with his intellectual brow, calm eyes, and high but
" narrow head;" so that it is necessary in courts of
justice to ask the witnesses whether they are Santalis
or Hindoos.37 Whether a heterogeneous people, such
as the inhabitants of some of the Polynesian islands,
formed by the crossing of two distinct races, with few
or no pure members left, would ever become homo-
geneous, is not known from direct evidence. But as
with our domesticated animals, a crossed breed can
certainly, in the course of a few generations, be fixed
and made uniform by careful selection,38 we may infer
that the free and prolonged intercrossing during many
generations of a heterogeneous mixture would supply
the place of selection, and overcome any tendency to
reversion, so that a crossed race would ultimately be-
come homogeneous, though it might not partake in an
equal degree of the characters of the two parent-races.

Of all the differences between the races of man, the
colour of the skin is the most conspicuous and one of
the best marked. Differences of this kind, it was for-
merly thought, could be accounted for by long expo-
sure under different climates ; but Pallas first shewed
that this view is not tenable, and he has been followed
by almost all anthropologists.39 The view has been

37 ' The Annals of Rural Bengal,' 1868, p. 134.

38 ' The Variation of Animals and Plants under Domestication,' vol.
ii. p. 95.

89 Pallas, 'Act. Acad. St. Petersburgh,' 1780, part ii. p. 69. He
was followed by Rudolphi, in his ' Beytr'age zur Anthropologic, ' 1812.
An excellent summary of the evidence is given by Godron, ' De
l'Espece,' 1859, vol. ii. p. 246, &c.

VOL. I. R

242 THE DESCENT OF MAN. Paet I.

rejected chiefly because the distribution of the variously
coloured races, most of whom must have long inhabited
their present homes, does not coincide with correspond-
ing differences of climate. Weight must also be given
to such cases as that of the Dutch families, who, as
we hear on excellent authority,40 have not undergone
the least change of colour, after residing for three cen-
turies in South Africa. The uniform appearance in
various parts of the world of gypsies and Jews, though
the uniformity of the latter has been somewhat exagge-
rated,41 is likewise an argument on the same side. A
very damp or a very dry atmosphere has been supposed
to be more influential in modifying the colour of the
skin than mere heat ; but as D'Orbigny in South
America, and Livingstone in Africa, arrived at diame-
trically opposite conclusions with respect to dampness
and dryness, any conclusion on this head must be con-
sidered as very doubtful.42

Various facts, which I have elsewhere given, prove
that the colour of the skin and hair is sometimes corre-
lated in a surprising manner with a complete immunity
from the action of certain vegetable poisons and from
the attacks of certain parasites. Hence it occurred to
me, that negroes and other dark races might have
acquired their dark tints by the darker individuals
escaping during a long series of generations from
the deadly influence of the miasmas of their native
countries.

I afterwards found that the same idea had long ago

40 Sir Andrew Smith, as quoted by Knox, ' Races of Man,' 1850,
p. 473.

41 See De Quatrefages on this head, • Revue des Cours Scientifiques,'
Oct. 17, 1868, p. 731.

4" Livingstone's ' Travels and Researches in S. Africa,' 1857, p. 338,
329. D'Orbigny, as quoted by Godron, • De l'Espece,' vol. ii. p. 266.

Chap. VII. THE RACES OF MAN. 243

occurred to Dr. Wells.43 That negroes, and even mulat-
toes, are almost completely exempt from the yellow-
fever, which is so destructive in tropical America, has
long been known.44 They likewise escape to a large
extent the fatal intermittent fevers that prevail along,
at least, 2600 miles of the shores of Africa, and which
annually cause one-fifth of the white settlers to die, and
another fifth to return home invalided.45 This immu-
nity in the negro seems to be partly inherent, de-
pending on some unknown peculiarity of constitution,
and partly the result of acclimatisation. Pouchet46
states that the negro regiments, borrowed from the
Viceroy of Egypt for the Mexican war, which had been
recruited near the Soudan, escaped the yellow-fever
almost equally well with the negroes originally brought
from various parts of Africa, and accustomed to the
climate of the West Indies. That acclimatisation plays
a part is shewn by the many cases in which negroes,
after having resided for some time in a colder climate,
have become to a certain extent liable to tropical
fevers.47 The nature of the climate under which the
white races have long resided, likewise has some in-
fluence on them ; for during the fearful epidemic of
yellow-fever in Demerara during 1837, Dr. Blair found
that the death-rate of the immigrants was proportional

43 See a paper read before the Royal Soc. in 1813, and published in
his Essays in 1818. I have given an account of Dr. Wells' views in the
Historical Sketch (p. svi) to my ' Origin of Species/ Various cases of
colour correlated with constitutional peculiarities are given in my
' Variation of Animals under Domestication,' vol. ii. p. 227, 335.

44 See, for instance, Nott and Gliddon, ' Types of Mankind,' p. 68.

45 Major Tulloch, in a paper read before the Statistical Society,
April 20th, 1840, and given in the ' Athenaeum,' 1840, p. 353.

46 * The Plurality of the Human Race ' (translat.), 1864, p. 60.

4' Quatrefages, ' Unite de l'Espece Humaine,' 1861, p. 205. Waitz,
'Introduct. to Anthropology,' translat. vol. i. 1863, p. 124. Living-
stone gives analogous cases in his ' Travels.'

R 2

244 THE DESCENT OF MAN. Part I.

to the latitude of the country whence they had come.
With the negro the immunity, as far as it is the result
of acclimatisation, implies exposure during a prodigious
length of time ; for the aborigines of tropical America,
who have resided there from time immemorial, are not
exempt from yellow-fever; and the Rev. B. Tristram
states, that there are districts in Northern Africa which
the native inhabitants are compelled annually to leave,
though the negroes can remain with safety.

That the immunity of the negro is in any degree
correlated with the colour of his skin is a mere conjec-
ture : it may be correlated with some difference in his
blood, nervous system, or other tissues. Nevertheless,
from the facts above alluded to, and from some connec-
tion apparently existing between complexion and a
tendency to consumption, the conjecture seemed to me
not improbable. Consequently I endeavoured, with but
little success,48 to ascertain how far it held good. The

48 In the spring of 1862 I obtained permission from the Director-
General of the Medical department of the Army, to transmit to the
surgeons of the various regiments on foreign service a blank table,
with the following appended remarks, but I have received no returns.
" As several well-marked cases have been recorded with our domestic
" animals of a relation between the colour of the dermal appendages
" and the constitution ; and it being notorious that there is some limited
" degree of relation between the colour of the races of man and the
" climate inhabited by them ; the following investigation seems worth
" consideration. Namely, whether there is any relation in Europeans
" between the colour of their hair, and their liability to the diseases of
" tropical countries. If the surgeons of the several regiments, when
" stationed in unhealthy tropical districts, would be so good as first to
" count, as a standard of comparison, how many men, in the force
" whence the sick are drawn, have dark and light-coloured hair, and
" hair of intermediate or doubtful tints ; and if a similar account were
" kept by the same medical gentlemen, of all the men who suffered
" from malarious and yellow fevers, or from dysentery, it would soon
" be apparent, after some thousand cases had been tabulated, whether
" there exists any relation between the colour of the hair and consti-
" tutional liability to tropical diseases. Perhaps no such relation would

Chap. VII. THE RACES OF MAN. 245

late Dr. Daniell, who had long lived on the West Coast
of Africa, told me that he did not believe in any such
relation. He was himself unusually fair, and had with-
stood the climate in a wonderful manner. When he
first arrived as a boy on the coast, an old and expe-
rienced negro chief predicted from his appearance that
this would prove the case. Dr. Nicholson, of Antigua,
after having attended to this subject, wrote to me that
he did not think that dark-coloured Europeans escaped
the yellow-fever better than those that were light-
coloured. Mr. J. M. Harris altogether denies49 that
Europeans with dark hair withstand a hot climate
better than other men ; on the contrary, experience has
taught him in making a selection of men for service
on the coast of Africa, to choose those with red hair.
As far, therefore, as these slight indications serve, there
seems no foundation for the hypothesis, which has been
accepted by several writers, that the colour of the black
races may have resulted from darker and darker indi-
viduals having survived in greater numbers, during
their exposure to the fever-generating miasmas of their
native countries.

Although with our present knowledge we cannot
account for the strongly-marked differences in colour
between the races of man, either through correlation
with constitutional peculiarities, or through the direct
action of climate ; yet we must not quite ignore the

" be discovered, but the investigation is well worth making. In case
"any positive result were obtained, it might be of some practical use
" in selecting men for any particular service. Theoretically the result
" would be of high interest, as indicating one means by which a race
" of men inhabiting from a remote period an unhealthy tropical climate,
" might have become dark-coloured by the better preservation of dark-
" haired or dark-complexioned individuals during a long succession of
" generations."

49 ' Anthropological Review,' Jan. 18GG, p. xxi.

246 THE DESCENT OF MAN. Part I.

latter agency, for there is good reason to believe that
some inherited effect is thus produced.50

We have seen in our third chapter that the condi-
tions of life, such as abundant food and general comfort,
affect in a direct manner the development of the bodily
frame, the effects being transmitted. Through the
combined influences of climate and changed habits of
life, European settlers in the United States undergo, as
is generally admitted, a slight but extraordinarily rapid
change of appearance. There is, also, a considerable
body of evidence shewing that in the Southern States
the house-slaves of the third generation present a
markedly different appearance from the field-slaves.51

If, however, we look to the races of man, as distri-
buted over the world, we must infer that their charac-
teristic differences cannot be accounted for by the
direct action of different conditions of life, even after
exposure to them for an enormous period of time. The
Esquimaux live exclusively on animal food; they are
clothed in thick fur, and are exposed to intense cold
and to prolonged darkness ; yet they do not differ in
any extreme degree from the inhabitants of Southern
China, who live entirely on vegetable food and are ex-
posed almost naked to a hot, glaring climate. The un-
clothed Fuegians live on the marine productions of their
inhospitable shores ; the Botocudos of Brazil wander

50 See, for instance, Quatrefages ('Eevue des Cours Scientifiques,'
Oct. 10, 1868, p. 724) on the effects of residence in Abyssinia and
Arabia, and other analogous cases. Dr. Eolle ('Der Mensch, seine
Abstammung,' &c, 1865, s. 99) states, on the authority of Khanikof,
that the greater number of German families settled in Georgia, have
acquired in the course of two generations dark hair and eyes. Mr. D.
Forbes informs me that the Quichuas in the Andes vary greatly in
colour, according to the position of the valleys inhabited by them.

51 Harlan, ' Medical Researches,' p. 532. Quatrefages (' Unite' de
l'Espece Humaine,' 1861, p. 128) has collected much evidence on this
head.

Chap. VII. THE EACES OF MAN. 247

about the hot forests of the interior and live chiefly on
vegetable productions ; yet these tribes resemble each
other so closely that the Fuegians on board the " Beagle"
were mistaken by some Brazilians for Botocudos. The
Botocudos again, as well as the other inhabitants of
tropical America, are wholly different from the Negroes
who inhabit the opposite shores of the Atlantic, are
exposed to a nearly similar climate, and follow nearly
the same habits of life.

Nor can the differences between the races of man be
accounted for, except to a quite insignificant degree, by
the inherited effects of the increased or decreased use of
parts. Men who habitually live in canoes, may have
their legs somewhat stunted ; those who inhabit lofty
regions have their chests enlarged ; and those who con-
stantly use certain sense-organs have the cavities in
which they are lodged somewhat increased in size, and
their features consequently a little modified. With
civilised nations, the reduced size of the jaws from
lessened use, the habitual play of different muscles
serving to express different emotions, and the increased
size of the brain from greater intellectual activity, have
together produced a considerable effect on their general
appearance in comparison with savages.52 It is also
possible that increased bodily stature, with no corre-
sponding increase in the size of the brain, may have
given to some races (judging from the previously ad-
duced cases of the rabbits) an elongated skull of the
dolichocephalic type.

Lastly, the little-understood principle of correlation
will almost certainly have come into action, as in the
case of great muscular development and strongly pro-

52 See Prof. Schaaffhausen, translat. in 'Anthropological Keview,'
Oct. 1868, p. 429.

248 THE DESCENT OF MAN. Part I.

jecting supra-orbital ridges. It is not improbable that
the texture of the hair, which differs much iu the dif-
ferent races, may stand in some kind of correlation with
the structure of the skin ; for the colour of the hair
and skin are certainly correlated, as is its colour and
texture with the Mandans. 53 The colour of the skin
and the odour emitted by it are likewise in some man-
ner connected. With the breeds of sheep the number
of hairs within a given space and the number of the
excretory pores stand in some relation to each other.54
If we may judge from the analogy of our domesticated
animals, many modifications of structure in man pro-
bably come under this principle of correlated growth.

We have now seen that the characteristic differences
between the races of man cannot be accounted for in a
satisfactory manner by the direct action of the condi-
tions of life, nor by the effects of the continued use of
parts, nor through the principle of correlation. We
are therefore led to inquire whether slight individual
differences, to which man is eminently liable, may
not have been preserved and augmented during a long
series of generations through natural selection. But
here we are at once met by the objection that beneficial
variations alone can be thus preserved ; and as far as
we are enabled to judge (although always liable to
error on this head) not one of the external differ-
ences between the races of man are of any direct or

53 Mr. Catlin states (' N. American Indians,' 3rd edit. 1842, to] . i. p.
49) that in the whole tribe of the Mandans, about one in ten or twelve
of the members of all ages and both sexes have bright silvery grey hair,
which is hereditary. Now this hair is as coarse and harsh as that of
a horse's mane, whilst the hair of other colours is fine and soft.

54 On the odour of the skin, Godron, ' Sur 1'Espece,' torn. ii. p. 217.
On the pores in the skin, Dr. Wilckens, ' Die Aufgaben der landwirth.
Zootechnik,' 1869, s. 7.

Chap. VII. THE KACES OF MAN. 249

special service to him. The intellectual and moral or
social faculties must of course be excepted from this re-
mark ; but differences in these faculties can have had
little or no influence on external characters. The vari-
ability of all the characteristic differences between the
races, before referred to, likewise indicates that these
differences cannot be of much importance ; for, had
they been important, they would long ago have been
either fixed and preserved, or eliminated. In this
respect man resembles those forms, called by naturalists
protean or polymorphic, which have remained extremely
variable, owing, as it seems, to their variations being of
an indifferent nature, and consequently to their having
escaped the action of natural selection.

We have thus far been baffled in all our attempts
to account for the differences between the races of man ;
but there remains one important agency, namely Sexual
Selection, which appears to have acted as powerfully
on man, as on many other animals. I do not intend
to assert that sexual selection will account for all the
differences between the races. An unexplained resi-
duum is left, about which we can in our ignorance
only say, that as individuals are continually born with,
for instance, heads a little rounder or narrower, and
with noses a little longer or shorter, such slight dif-
ferences might become fixed and uniform, if the un-
known agencies which induced them were to act in a
more constant manner, aided by long-continued inter-
crossing. Such modifications come under the provi-
sional class, alluded to in our fourth chapter, which for
the want of a better term have been called spontaneous
variations. Nor do I pretend that the effects of sexual
selection can be indicated with scientific precision ; but
it can be shewn that it would be an inexplicable fact if
man had not been modified by this agency, which has

250 THE DESCENT OF MAN. Paet I.

acted so powerfully on innumerable animals, both high
and low in the scale. It can further be shewn that the
differences between the races of man, as in colour, hairy-
ness, form of features, &c, are of the nature which it
might have been expected would have been acted on by
sexual selection. But in order to treat this subject in a
fitting manner, I have found it necessary to pass the
whole animal kingdom in review ; I have therefore de-
voted to it the Second Part of this work. At the close
I shall return to man, and, after attempting to shew
how far he has been modified through sexual selection,
will give a brief summary of the chapters in this First
Part,

Part II.

SEXUAL SELECTION.

Part II.— SEXUAL SELECTION.

CHAPTER VIII.

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 — On the limita-
tion of the numbers of the two sexes through natural selection.

With animals which have their sexes separated, the
males necessarily differ from the females in their organs
of reproduction ; and these afford the primary sexual
characters. But the sexes often differ in what Hunter
has called secondary sexual characters, which are not
directly connected with the act of reproduction; for
instance, in the male possessing certain organs of sense
or locomotion, of which the female is quite destitute, or
in having them more highly-developed, in order that
he may readily find or reach her ; or again, in the male
having special organs of prehension so as to hold her
securely. These latter organs of infinitely diversified
kinds graduate into, and in some cases can hardly be
distinguished from, those which are commonly ranked
as primary, such as the complex appendages at the
apex of the abdomen in male insects. Unless indeed

254 THE PRINCIPLES OF Part II.

we confine the term " primary ': to the reproductive
glands, it is scarcely possible to decide, as far as the
organs of prehension are concerned, which ought to
be called primary and which secondary.

The female often differs from the male in having
organs for the nourishment or protection of her young,
as the mammary glands of mammals, and the ab-
dominal sacks of the marsupials. The male, also, in
some few cases differs from the female in possessing
analogous organs, as the receptacles for the ova pos-
sessed by the males of certain fishes, and those tem-
porarily developed in certain male frogs. Female bees
have a special apparatus for collecting and carrying
pollen, and their ovipositor is modified into a sting for
the defence of their larvae and the community. In the
females of many insects the ovipositor is modified in
the most complex manner for the safe placing of the
eggs. Numerous similar cases could be given, but they
do not here concern us. There are, however, other
sexual differences quite disconnected with the primary
organs with which we are more especially concerned —
such as the greater size, strength, and pugnacity of the
male, his weapons of offence or means of defence
against rivals, his gaudy colouring and various orna-
ments, his power of song, and other such characters.

Besides the foregoing primary and secondary sexual
differences, the male and female sometimes differ in
structures connected with different habits of life, and
not at all, or only indirectly, related to the reproductive
functions. Thus the females of certain flies (Culicidae
and Tabanidse) are blood-suckers, whilst the males live
on flowers and have their mouths destitute of man-
dibles.1 The males alone of certain moths and of some

* Westwood, 'Modern Class, of Insects,' vol. ii. 1840, p. 541. In

Chap. VIII. SEXUAL SELECTION. 255

crustaceans (e.g. Tanais) have imperfect, closed mouths,
and cannot feed. The Complemental males of certain
cirripedes live like epiphytic plants either on the female
or hermaphrodite form, and are destitute of a mouth
and prehensile limbs. In these cases it is the male
which has been modified and has lost certain import-
ant organs, which the females and other members of the
same group possess. In other cases it is the female
which has lost such parts ; for instance, the female glow-
worm is destitute of wings, as are many female moths,
some of which never leave their cocoons. Many female
parasitic crustaceans have lost their natatory legs. In
some weevil-beetles (Curculionidse) there is a great
difference between the male and female in the length
of the rostrum or snout ; 2 but the meaning of this and
of many analogous differences, is not at all understood.
Differences of structure between the two sexes in rela-
tion to different habits of life are generally confined to
the lower animals ; but with some few birds the beak
of the male differs from that of the female. No doubt
in most, but apparently not in all these cases, the dif-
ferences are indirectly connected with the propagation
of the species : thus a female which has to nourish a
multitude of ova will require more food than the male,
and consequently will require special means for procur-
ing it. A male animal which lived for a very short
time might without detriment lose through disuse its
organs for procuring food ; but he would retain his
locomotive organs in a perfect state, so that he might
reach the female. The female, on the other hand,
might safely lose her organs for -flying, swimming,

regard to the statement about Tanais, mentioned below, I am indebted
to Fritz Muller.

2 Kirby and Spence, 'Introduction to Entomology,' vol. iii. 1826,
p. 309.

256 THE PKINCIPLES OF Part II.

or walking, if she gradually acquired habits which ren-
dered such powers useless.

We are, however, here concerned only with that kind
of selection, which I have called sexual selection. This
depends on the advantage which certain individuals have
over other individuals of the same sex and species, in
exclusive relation to reproduction. When the two sexes
differ in structure in relation to different habits of life,
as in the cases above mentioned, they have no doubt
been modified through natural selection, accompanied
by inheritance limited to one and the same sex. So
again the primary sexual organs, and those for nourish-
ing or protecting the young, come under this same head ;
for those individuals which generated or nourished their
offspring best, would leave, ceteris paribus, the greatest
number to inherit their superiority ; whilst those which
generated or nourished their offspring badly, would leave
but few to inherit their weaker powers. As the male
has to search for the female, he requires for this purpose
organs of sense and locomotion, but if these organs are
necessary for the other purposes of life, as is generally
the case, they will have been developed through natural
selection. When the male has found the female he
sometimes absolutely requires prehensile organs to hold
her ; thus Dr. WTallace informs me that the males of cer-
tain moths cannot unite with the females if their tarsi
or feet are broken. The males of many oceanic crusta-
ceans have their legs and antennae modified in an extra-
ordinary manner for the prehension of the female ;
h^nce we may suspect that owing to these animals
being washed about by the waves of the open sea, they
absolutely require these organs in order to propagate
their kind, and if so their development will have been
the result of ordinary or natural selection.

W7hen the two sexes follow exactly the same habits

Chap. VIII. SEXUAL SELECTION. 257

of life, and the male has more highly developed sense
or locomotive organs than the female, it may be that
these in their perfected state are indispensable to the
male for finding the female ; but in the vast majority
of cases, they serve only to give one male an advan-
tage over another, for the less well-endowed males,
if time were allowed them, would succeed in pair-
ing with the females ; and thev would in all other
respects, judging from the structure of the female, be
equally well adapted for their ordinary habits of life.
In such cases sexual selection must have come into
action, for the males have acquired their present struc-
ture, not from being better fitted to survive in the
struggle for existence, but from having gained an ad-
vantage over other males, and from having transmitted
this advantage to their male offspring alone. It was the
importance of this distinction which led me to designate
this form of selection as sexual selection. So again,
if the chief service rendered to the male by his pre-
hensile organs is to prevent the escape of the female
before the arrival of other males, or when assaulted by
them, these organs will have been perfected through
sexual selection, that is by the advantage acquired by
certain males over their rivals. But in most cases it
is scarcely possible to distinguish between the effects
of natural and sexual selection. Whole chapters could
easily be filled with details on the differences between
the sexes in their sensorv, locomotive, and prehensile
organs. As, however, these structures are not more
interesting than others adapted for the ordinary pur-
poses of life, I shall almost pass them over, giving only
a few instances under each class.

There are many other structures and instincts which
must have been developed through sexual selection —
such as the weapons of offence and the means of defence

VOL. L s

258 THE PRINCIPLES OF Part II.

possessed by the males for fighting with and driving
away their rivals — their courage and pugnacity — their
ornaments of many kinds — their organs for producing
vocal or instrumental music — and their glands for
emitting odours; most of these latter structures serv-
ing only to allure or excite the female. That these
characters are the result of sexual and not of ordinary
selection is clear, as unarmed, unornamented, or un-
attractive males would succeed equally well in the
battle for life and in leaving a numerous progeny, if
better endowed males were not present. We may infer
that this would be the case, for the females, which are
unarmed and unornamented, are able to survive and
procreate their kind. Secondary sexual characters of
the kind just referred to, will be fully discussed in the
following chapters, as they are in many respects in-
teresting, but more especially as they depend on the
will, choice, and rivalry of the individuals of either sex.
When we behold two males fighting for the possession
of the female, or several male birds displaying their
gorgeous plumage, and performing the strangest antics
before an assembled bodv of females, we cannot doubt
that, though led by instinct, they know what they are
about, and consciously exert their mental and bodily
powers.

In the same manner as man can improve the breed
of his game-cocks by the selection of those birds which
are victorious in the cockpit, so it appeal's that the
strongest and most vigorous males, or those provided
with the best weapons, have prevailed under nature,
and have led to the improvement of the natural breed
or species. Through repeated deadly contests, a slight-
degree of variability, if it led to some advantage, how-
ever slight, would suffice for the work of sexual selec-
tion ; and it is certain that secondarv sexual characters

Chap; VIII. SEXUAL SELECTION. 259

are eminently variable. In the same manner as man
can give beauly, according to his standard of taste, to his
male poultry — can give to the Sebright bantam a new
and elegant plumage, an erect and peculiar carriage —
so it appears that in a state of nature female birds, by
having long selected the more attractive males, have
added to their beauty. No doubt this implies powers
of discrimination and taste on the part of the female
which will at first appear extremely improbable; but
I hope hereafter to shew that this is not the case.

From our ignorance on several points, the precise
manner in which sexual selection acts is to a certain
extent uncertain. Nevertheless if those naturalists who
already believe in the mutability of species, will read
the following chapters, they will, I think, agree with
me that sexual selection has played an important part
in the history of the organic world. It is certain that
with almost all animals there is a struggle between the
males for the possession of the female. This fact is so
notorious that it would be superfluous to give instances.
Hence the females, supposing that their mental capacity
sufficed for the exertion of a choice, could select. one out
of several males. But in numerous cases it appears as
if it had been specially arranged that there should be
a struggle between many males. Thus with migratory
birds, the males generally arrive before the females at
their place of breeding, so that many males are ready
to contend for each female. The bird-catchers assert
that this is invariably the case with the nightingale and
blackcap, as I am informed by Mr. ^Tenner Weir, who
confirms the statement with respect to the latter species.

Mr. Swayslancl of Brighton, who has been in the habit,
during the last forty years, of catching our migratory
birds on their first arrival, writes to me that he has
never known the females of any species to arrive before

s 2

260 THE PRINCIPLES OF Part II.

their males. During one spring he shot thirty-nine
males of Ray's wagtail (Budytes Baii) before he saw a
single female. Mr. Gould has ascertained by dissection,
as he informs me, that male snipes arrive in this
country before the females ; but this hardly concerns us,
as snipes do not breed here. In the case of fish, at the
period when the salmon ascend our rivers, the males in
large numbers are ready to breed before the females.
So it apparently is with frogs and toads. Throughout
the great class of insects the males almost always
emerge from the pupal state before the other sex, so
that they generally swarm for a time before any females
can be seen.3 The cause of this difference between
the males and females in their periods of arrival and
maturity is sufficiently obvious. Those males which
annually first migrated into any country, or which in
the spring were first ready to breed, or were the most
eager, would leave the largest number of offspring ;
and these would tend to inherit similar instincts and
constitutions. On the whole there can be no doubt that
with almost all animals, in which the sexes are separate,
there is a constantly recurrent struggle between the
males for the possession of the females.

Our difficulty in regard to sexual selection lies in
understanding how it is that the males which conquer
other males, or those which prove the most attractive
to the females, leave a greater number of offspring
to inherit their superiority than the beaten and less

3 Even with those of plants in which the sexes are separate, the male
flowers are generally mature before the female. Many hermaphrodite
plants are, as first shewn by C. K. Sprengel, dichogamous ; that is,
their male and female organs are not ready at the same time, so thai
they cannot be self-fertilised. Now with such plants the pollen is
generally mature in the same flower before the stigma, though there
are some exceptional species in which the female organs are mature
before the male.

Chap. VIII. SEXUAL SELECTION. 261

attractive males. Unless this result should follow, the
characters which give to certain males an advantage
over others, could not be perfected and augmented
through sexual selection. When the sexes exist in
exactly equal numbers, the worst-endowed males will
ultimately find females (except where polygamy pre-
vails), and leave as many offspring, equally well fitted for
their general habits of life, as the best-endowed males.
From various facts and considerations, I formerly in-
ferred that with most animals, in which secondary sexual
characters are well developed, the males considerably
exceeded the females in number; and this does hold
good in some few cases. If the males were to the
females as two to one, or as three to two, or even in
a somewhat lower ratio, the whole affair would be
simple ; for the better-armed or more attractive males
would leave the largest number of offspring. But after
investigating, as far as possible, the numerical propor-
tions of the sexes, I do not believe that any great
inequality in number commonly exists. In most cases
sexual selection appears to have been effective in the
following manner.

Let us take any species, a bird for instance, and
divide the females inhabiting a district into two equal
bodies: the one consisting of the more vigorous and
better-nourished individuals, and the other of the less
vigorous and healthy. The former, there can be little
doubt, would be ready to breed in the spring before the
others ; and this is the opinion of Mr. Jenner Weir, who
has during many years carefully attended to the habits
of birds. There can also be no doubt that the most
vigorous, healthy, and best-nourished females would on
an average succeed in rearing the largest number of
offspring. The males, as we have seen, are generally
ready to breed before the females ; of the males the

262 THE PRINCIPLES OF Part II.

strongest, and with some species the best armed, drive
away the weaker males ; and the former would then
unite with the more vigorous and best-nourished fe-
males, as these are the first to breed. Such vigorous
pairs would surely rear a larger number of offspring
than the retarded females, which would be compelled,
supposing the sexes to be numerically equal, to unite
with the conquered and less powerful males ; and tliis
is all that is wanted to add, in the course of successive
generations, to the size, strength and courage of the
males, or to improve their weapons.

But in a multitude of cases the males which conquer
other males, do not obtain possession of the females,
independently of choice on the part of the latter. The
courtship of animals is by no means so simple and short
an affair as might be thought. The females are most
excited by, or prefer pairing with, the more ornamented
males, or those which are the best songsters, or play the
best antics ; but it is obviously probable, as has been
actually observed in some cases, that they would at the
same time prefer the more vigorous and lively males.4
Thus the more vigorous females, which are the first to
breed, will have the choice of many males ; and though
they may not always select the strongest or best armed,
they will select those which are vigorous and well armed,
and in other respects the most attractive. Such early
pairs would have the same advantage in rearing off-
spring on the female side as above explained, and nearly
the same advantage on the male side. And this ap-
parently has sufficed during a long course of generations
to acid not only to the strength and fighting-powers of

4 I have received information, hereafter to be given, to this effect
with respect to poultry. Even with birds, such as pigeons, which
pair for life, the female, as I hear from Mr. Jenner Weir, will desert
her mate if he is injured or grows weak.

Chap. VIII. SEXUAL SELECTION. 263

the males, but likewise to their various ornaments or
other attractions.

In the converse and much rarer case of the males
selecting particular females, it is plain that those which
were the most vigorous and had conquered others, would
have the freest choice ; and it is almost certain that they
would select vigorous as well as attractive females. Such
pairs would have an advantage in rearing offspring, more
especially if the male had the power to defend the
female during the pairing-season, as occurs with some
of the higher animals, or aided in providing for the
young. The same principles would apply if both sexes
mutually preferred and selected certain individuals of
the opposite sex ; supposing that they selected not only
the more attractive, but likewise the more vigorous
individuals.

Numerical Proportion of the Two Sexes. — I have
remarked that sexual selection would be a simple
affair if the males considerably exceeded in number
the females. Hence I w7as led to investigate, as far
as I could, the proportions between the two sexes of
as many animals as possible ; but the materials are
scanty. I will here give only a brief abstract of the
results, retaining the details for a supplementary dis-
cussion, so as not to interfere with the course of my
argument. Domesticated animals alone afford the
opportunity of ascertaining the proportional numbers
at birth ; but no records have been specially kept for
this purpose. By indirect means, however, I have
collected a considerable body of statistical data, from
which it appears that with most^ of our domestic
animals the sexes are nearly equal at birth. Thus with
race-horses, 25,560 births have been recorded during
twenty-one years, and the male births have been to the
female births as 99*7 to 100. With greyhounds the

264 THE PRINCIPLES OF Part II.

inequality is greater than with any other animal, for
during twelve years, out of 6878 births, the male births
have been as HO'l to 100 female births. It is, however,
in some degree doubtful whether it is safe to infer that
the same proportional numbers would hold good under
natural conditions as under domestication ; for slight
and unknown differences in the conditions affect to a
certain extent the proportion of the sexes. Thus with
mankind, the male births in England are as 104*5,
in Kussia as 108'9, and with the Jews of Livonia as
120 to 100 females. The proportion is also mysteriously
affected by the circumstance of the births being legiti-
mate or illegitimate.

For our present purpose we are concerned with the
proportion of the sexes, not at birth, but at maturity,
and this adds another element of doubt ; for it is a well
ascertained fact that with man a considerably larger
proportion of males than of females die before or during
birth, and during the first few years of infancy. So it
almost certainly is with male lambs, and so it may be
with the males of other animals. The males of some
animals kill each other by fighting ; or they drive
each other about until they become greatly emaciated.
They must, also, whilst wandering about in eager search
for the females, be often exposed to various dangers.
With many kinds of fish the males are much smaller
than the females, and they are believed often to be
devoured by the latter or by other fishes. With some
birds the females appear to die in larger proportion
than the males : they are also liable to be destroyed on
their nests, or whilst in charge of their young. With
insects the female larvae are often larger than those of
the males, and would consequently be more likely to be
devoured : in some cases the mature females are less
active and less rapid in their movements than the males,

Chap. VIII. SEXUAL SELECTIOX. 265

and would not be so well able to escape from danger.
Hence, with animals in a state of nature, in order to
judge of the proportions of the sexes at maturity, we
must rely on mere estimation ; and this, except perhaps
when the inequality is strongly marked, is but little
trustworthy. Nevertheless, as far as a judgment can
be formed, we may conclude from the facts given in the
supplement, that the males of some few mammals, of
many birds, of some fish and insects, considerably
exceed in number the females.

The proportion between the sexes fluctuates slightly
during successive years : thus with race-horses, for every
100 females born, the males varied from 107*1 in one
year to 92*0 in another year, and with greyhounds from
116*3 to 95*3. But had larger numbers been tabulated
throughout a more extensive area than England, these
fluctuations would probably have disappeared ; and such
as they are, they would hardly suffice to lead under
a state of nature to the effective action of sexual selec-
tion. Nevertheless with some few wild animals, the
proportions seem, as shewn in the supplement, to fluc-
tuate either during different seasons or in different
localities in a sufficient degree to lead to such action.
For it should be observed that any advantage gained
during certain years or in certain localities by those
males which were able to conquer other males, or were
the most attractive to the females, would probably be
transmitted to the offspring and would not subsequently
be eliminated. During the succeeding seasons, when
from the equality of the sexes every male was every-
where able to procure a female, the stronger or more
attractive males previously produced would still have
at least as good a chance of leaving offspring as the
less strong or less attractive.

Polygamy. — The practice of polygamy leads to the

2G6 THE PKINCIPLES OF Part II.

same results as would follow from au actual inequality
in the number of the sexes ; for if each male secures
two or more females, many males will not be able to
pair ; and the latter assuredly will be the weaker or
less attractive individuals. Many mammals and some
few birds are polygamous, but with animals belonging to
the lower classes I have found no evidence of this habit.
The intellectual powers of such animals are, perhaps,
not sufficient to lead them to collect and guard a harem
of females. That some relation exists between poly-
gamy and the development of secondary sexual cha-
racters, appears nearly certain; and this supports the
view that a numerical preponderance of males would
be eminently favourable to the action of sexual selection.
Nevertheless many animals, especially birds, which are
strictly monogamous, display stroDgly- marked secondary
sexual characters ; whilst some few animals, which are
polygamous, are not thus characterised.

We will first briefly run through the class of mam-
mals, and then turn to birds. The gorilla seems to be
a polygamist, and the male differs considerably from
the female ; so it is with some baboons which live in
herds containing twice as many adult females as males.
In South America the Mycetes car ay a presents well-
marked sexual differences in colour, beard, and vocal
organs, and the male generally lives with two or three
wives : the male of the Cebus ca/pucinus differs some-
what from the female, and appears to be polygamous.5
Little is known on this head with respect to most other
monkeys, but some species are strictly monogamous.
The ruminants are eminently polygamous, and they

5 On the Gorilla, Savage and Wyinan, ' Boston Journal of Nat. Hist.'
vol. v. 1845-47, p. 423. On Cynocephalus, Brehm, 'Illust. Thierleberi,'
B. i. 1864, s. 77. On Mycetes, Kengger, 'Naturgeseh. : Saugethiere
von Paraguay,' 1830, s. 14, 20. On Cebus, Brehin, ibid. s. 108.

Chap. VIII. SEXUAL SELECTION. 207

more frequently present sexual differences than almost
any other group of mammals, especially in their weapons,
but likewise in other characters. Most deer, cattle, and
sheep are polygamous; as are most antelopes, though
some of the latter are monogamous. Sir Andrew
Smith, in speaking of the antelopes of South Africa,
says that in herds of about a dozen there was rarely
more than one mature male. The Asiatic Antilope
saiga appears to be the most inordinate polygamist
in the world; for Pallas6 states that the male drives
away all rivals, and collects a herd of about a hundred,
consisting of females and kids : the female is hornless
and has softer hair, but does not otherwise differ much
from the male. The horse is polygamous, but, except
in his greater size and in the proportions of his body,
differs but little from the mare. The wild boar, in his
great tusks and some other characters, presents well-
marked sexual characters ; in Europe and in India he
leads a solitary life, except during the breeding-season ;
but at this season he consorts in India with several
females, as Sir W. Elliot, who has had large experience
in observing this animal, believes : whether this holds
good in Europe is doubtful, but is supported by some
statements. The adult male Indian elephant, like the
boar, passes much of his time in solitude; but when
associating with others, "it is rare to find," as Dr.
Campbell states, " more than one male with a whole
<; herd of females." The larger males expel or kill the
smaller and weaker ones. The male differs from the
female by his immense tusks and greater size, strength,
and endurance ; so great is the difference in these latter

6 Pallas, ' Spicilegia Zoolog/ Fasc. xii. 1777, p. 29. Sir Andrew
Smith, ' Illustrations of the Zoology of S. Africa/ 1849, pi. 29, on the
Kobus. Owen, in his 'Anatomy of Vertebrates ' (vol. iii. 18l8, p. 633)
gives a table incidentally showing which species of Antelopes pair and
which are gregarious.

268 THE PRINCIPLES OF Part II.

respects, that the males when caught are valued at
twenty per cent, above the females.7 With other pachy-
dermatous animals the sexes differ very little or not at
all, and they are not, as far as known, polygamists.
Hardly a single species amongst the Cheiroptera and
Edentata, or in the great Orders of the Rodents and
Insectivora, presents well-developed secondary sexual
differences ; and I can find no account of any species
being polygamous, excepting, perhaps, the common rat,
the males of which, as some rat-catchers affirm, live
with several females.

The lion in South Africa, as I hear from Sir Andrew
Smith, sometimes lives with a single female, but gene-
rally with more than one, and, in one case, was found
with as many as five females, so that he is polygamous.
He is, as far as I can discover, the sole polygamist in
the whole group of the terrestrial Carnivora, and he
alone presents well-marked sexual characters. If, how-
ever, we turn to the marine Carnivora, the case is widely
different ; for many species of seals offer, as we shall
hereafter see, extraordinary sexual differences, and they
are eminently polygamous. Thus the male sea-ele-
phant of the Southern Ocean, always possesses, accord-
ing to Peron, several females, and the sea-lion of Forster
is said to be surrounded by from twenty to thirty females.
In the North, the male sea-bear of Steller is accom-
panied by even a greater number of females.

With respect to birds, many species, the sexes of
which differ greatly from each other, are certainly
monogamous. In Great Britain we see well-marked
sexual differences in, for instance, the wild-duck which
pairs with a single female, with the common blackbird,

' Dr. Campbell, in ' Proc. Zoolog. Soc.' 1869, p. 138. See also an
interesting paper, by Lieut. Johnstone, in ' Proc. Asiatic Soc. ofBeDgal,'
May, 1868.

Chap. VIII. SEXUAL SELECTION. 269

and with the bullfinch which is said to pair for life. So
it is, as I am informed by Mr. Wallace, with the Chat-
terers or Cotingidoe of South America, and numerous
other birds. In several groups I have not been able to
discover whether the species are polygamous or mono-
gamous. Lesson says that birds of paradise, so re-
markable for their sexual differences, are polygamous,
but Mr. Wallace doubts whether he had sufficient evi-
dence. Mr. Salvin informs me that he has been led
to believe that humming-birds are polygamous. The
male widow-bird, remarkable for his caudal plumes,
certainly seems to be a polygamist.8 I have been
assured by Mr. Jenner Weir and by others, that three
starlings not rarely frequent the same nest ; but whether
this is a case of polygamy or polyandry has not been
ascertained.

The Gallinaceae present almost as strongly marked
sexual differences as birds of paradise or humming-
birds, and many of the species are, as is well known,
polygamous ; others being strictly monogamous. WThat
a contrast is presented between the sexes of the poly-
gamous peacock or pheasant, and the monogamous
guinea-fowl or partridge ! Many similar cases could
be given, as in the grouse tribe, in which the males
of the polygamous capercailzie and black-cock differ
greatly from the females ; whilst the sexes of the mono-
gamous red grouse and ptarmigan differ very little.
Amongst the Cursores, no great number of species
offer strongly - marked sexual differences, except the
bustards, and the great bustard (Otis tarda), is said to

8 ' The Ibis,' vol. iii. 1861, p. 133, on the Progne Widow-bird. Sot-
also on the Vidua axillaris, ibid. vol. ii. 1860, p. 211. On the poly-
gamy of the Capercailzie and Great Bustard, see L. Lloyd, ' Game Birds
of Sweden,' 1867, p. 19, and 182. Montagu and Selby speak of the
Black Grouse as polygamous and of the Bed Grouse as monogamous.

270 THE PRINCIPLES OF Part II.

be polygamous. With the Grallatores, extremely few
species differ sexually, but the ruff" (Machetes pugnax)
affords a strong exception, and this species is believed
by Montagu to be a polygamist. Hence it appears
that with birds there often exists a close relation
between polygamy and the development of strongly-
marked sexual differences. On asking Mr. Bartlett, at
the Zoological Gardens, who has had such large ex-
perience with birds, whether the male tragopan (one of
the Gallinacere) was polygamous, I was struck by his
answering, " I do not know, but should think so from
" his splendid colours."

It deserves notice that the instinct of pairing with a
single female is easily lost under domestication. The
wild-duck is strictly monogamous, the domestic-duck
hiprhlv polv^amous. The Rev. W. D. Fox informs me
that with some half-tamed wild-ducks, kept on a large
pond in his neighbourhood, so many mallards were shot
bv the gamekeeper that only one was left for every
seven or eight females ; yet unusually large broods
were reared. The guinea-fowl is strictly monogamous ;
but Mr. Fox finds that his birds succeed best when he
keeps one cock to two or three hens.9 Canary-birds
pair in a state of nature, but the breeders in England
successfully put one male to four or five females ; never-
theless the first female, as Mr. Fox has been assured,
is alone treated as the wife, she and her young ones
being fed by him ; the others are treated as concubines.
I have noticed these cases, as it renders it in some
degree probable that monogamous species, in a state of
nature, might readily become either temporarily or per-
manently polygamous.

9 The Rev. E. S. Dixon, however, speaks positively (' Ornamental
Poultry,' 1S48, p. 76) about the eggs of the guinea-fowl being infertile
when more than one female is kept with the same male.

Chap. VIII. SEXUAL SELECTION. 271

With respect to reptiles and fishes, too little is known
of their habits to enable ns to sj)eak of their marriage
arrangements. The stickle-back G-asterosteus), however,
is said to be a polygamist ;10 and the male during the
breeding-season differs conspicuously from the female.

To sum up on the means through which, as far as
we can judge, sexual selection has led to the develop-
ment of secondary sexual characters. It has been shewn
that the largest number of vigorous offspring will be
reared from the pairing of the strongest and best-armed
males, which have conquered other males, with the
most vigorous and best-nourished females, which are
the first to breed in the spring. Such females, if
they select the more attractive, and at the same time
vigorous, males, will rear a larger number of offspring
than the retarded females, which must pair writh the
less vigorous and less attractive males. So it will be
if the more vigorous males select the more attractive
and at the same time healthy and vigorous females ;
and this will especially hold good if the male defends
the female, and aids in providing food for the young.
The advantage thus gained by the more vigorous pairs
in rearing a larger number of offspring has apparently
sufficed to render sexual selection efficient. But a lame
preponderance in number of the males over the females
would be still more efficient ; whether the preponder-
ance was only occasional and local, or permanent;
whether it occurred at birth, or subsequently from the
greater destruction of the females ; or whether it in-
directly followed from the practice of polygamy.

TJie Male generally more modified than the Female. —
Throughout the animal kingdom, when the sexes differ

10

Noel Humphreys, 'River Gardens,' 1857,

272 THE PRINCIPLES OF Part II.

from each other in external appearance, it is the male
which, with rare exceptions, has been chiefly modified ;
for the female still remains more like the young of her
own species, and more like the other members of the
same group. The cause of this seems to lie in the
males of almost all animals having stronger passions
than the females. Hence it is the males that fight
together and sedulously display their charms before
the females ; and those which are victorious transmit
their superiority to their male offspring. Why the
males do not transmit their characters to both sexes
will hereafter be considered. That the males of all
mammals eagerly pursue the females is notorious to
every one. So it is with birds ; but many male birds
do not so much pursue the female, as display their
plumage, perform strange antics, and pour forth their
song, in her presence. With the few fish which have
been observed, the male seems much more eager than
the female ; and so it is with alligators, and apparently
with Batrachians. Throughout the enormous class of
insects, as Kirby remarks,11 " the law is, that the male
" shall seek the female." With spiders and crustaceans,
as I hear from two great authorities, Mr. Blackwall and
Mr. C. Spence Bate, the males are more active and more
erratic in their habits than the females. With insects and
crustaceans, when the organs of sense or locomotion are
present in the one sex and absent in the other, or when,
as is frequently the case, they are more highly developed
in the one than the other, it is almost invariably the male,
as far as I can discover, which retains such organs, or has
them most developed ; and this shews that the male is
the more active member in the courtship of the sexes.12

11 Kirby and Spence, 'Introduction to Entomology,' vol. iii. 182G,
p. 342.

12 One parasitic Hymenopterous insect ("Westwood, ' Modern Class,
of Insects,' vol. ii. p. 160) forms an exception to the rule, as the male

Chap. VIII. SEXUAL SELECTION. 273

The female, on the other hand, with the rarest excep-
tion, is less eager than the male. As the illustrious
Hunter 13 long ago observed, she generally " requires to
" be courted ; " she is coy, and may often be seen en-
deavouring for a long time to escape from the male.
Everv one who has attended to the habits of animals

af

will be able to call to mind instances of this kind.
Judging from various facts, hereafter to be given, and
from the results which may fairly be attributed to
sexual selection, the female, though comparatively
passive, generally exerts some choice and accepts one
male in preference to others. Or she may accept, as
appearances would sometimes lead us to believe, not
the male which is the most attractive to her, but the
one which is the least distasteful. The exertion of
some choice on the part of the female seems almost as
general a law as the eagerness of the male.

We are naturally led to enquire why the male in so
many and such widely distinct classes has been ren-
dered more easrer than the female, so that he searches
for her and plays the more active part in courtship.
It would be no advantage and some loss of power if
both sexes were mutually to search for each other ; but
why should the male almost always be the seeker ?
With plants, the ovules after fertilisation have to be
nourished for a time ; hence the pollen is necessarily
brought to the female organs — being placed on the
stigma, through the agency of insects or of the wind,

has rudimentary wings, and never quits the cell in which it is born,
whilst the female has well-developed wings. "- Audouin believes that
the females are impregnated by the males which are born in the same
cells with them ; but it is much more probable that the females visit
other cells, and thus avoid close interbreeding. We shall hereafter
meet with a few exceptional cases, in various classes, in which the
female, instead of the male, is the seeker and wooer.

13 ' Essays and Observations,' edited by Owen, vol. i. 1861, p. 194.

VOL. I. T

274 THE PRINCIPLES OF Part II.

or by the spontaneous movements of the stamens ; and
with the Algae, &c, by the locomotive power of the
antherozooids. With lowly-organised animals perma-
nently affixed to the same spot and having their sexes
separate, the male element is invariably brought to
the female ; and we can see the reason why ; for the
ova, even if detached before being fertilised and not
requiring subsequent nourishment or protection, would
be, from their larger relative size, less easily transported
than the male element. Hence plants 14 and many of
the lower animals are, in this respect, analogous. The
males of affixed animals having been thus led to emit
their fertilising element, it is natural that any of their
descendants, which rose in the scale and became loco-
motive, should retain the same habit, and should closely
approach the female, so that the fertilising element
might not run the risk of a long transit through the
waters of the sea. With some few of the lower ani-
mals, the females alone are iixed, and with these the
males must be the seekers. With respect to forms,
of which the progenitors were primordially free, it is
difficult to understand why the males should inva-
riably have acquired the habit of approaching the
females, instead of being approached by them. But
in all cases, in order that the males should be efficient
seekers, it would be necessary that they should be en-
dowed with strong passions ; and the acquirement of
such passions would naturally follow from the more
eager males leaving a larger number of offspring than
the less eager.

The great eagerness of the male has thus indirectly

14 Prof. Sachs ('Lehrbuch der Botanik,' 1870, s. 633) in speaking of
the male and ft male reproductive cells, remarks, " verlialt sich die eine
" bei der Vereinigung activ, . . . die andere erscheint bei dur Verein-
" igung passiv."

Chap. VIII. SEXUAL SELECTION. 275

led to the much more frequent development of secon-
dary sexual characters in the male than in the female.
But the development of such characters will have been
much aided, if the conclusion at which I arrived after
studying domesticated animals, can be trusted, namely,
that the male is more liable to vary than the female.
I am aware how difficult it is to verify a conclusion of
this kind. Some slight evidence, however, can be gained
by comparing the two sexes in mankind, as man has
been more carefully observed than any other animal.
During the Novara Expedition 15 a vast number of mea-
surements of various parts of the body in different races
were made, and the men were found in almost every
case to present a greater range of variation than the
women ; but I shall have to recur to this subject in
a future chapter. Mr. J. Wood,16 who has carefully
attended to the variation of the muscles in man, puts
in italics the conclusion that " the greatest number of
" abnormalities in each subject is found in the males."
He had previously remarked that "altogether in 102
" subjects the varieties of redundancy were found to
" be half as many again as in females, contrasting
" widely with thft greater frequency of deficiency in
" females before described." Professor Macalister like-
wise remarks17 that variations in the muscles "are
" probably more common in males than females."
Certain muscles which are not normally present in man-
kind are also more frequently developed in the male
than in the female sex, although exceptions to this rule

15 'Reise der Novara: Anthropolog. Tlieil,"* 1867, s. 21G-2G9. TJie
results were calculated by Dr. Weisbach from measurements made by
Drs. K. Scherzer and Schwarz. On the greater variability of the males
of domesticated animals, see my ' Variation of Animals and Plants
under Domestication,' vol. ii. 1868, p-. 75.

16 'Proceedings Royal Soc.' vol. xvi. July, 18GS, p. :">19 and 524.

17 < Proc. Royal Irish Academy,' vol. x. 1868, p. 123.

rti •)

276 THE PRINCIPLES OF Part II.

are said to occur. Dr. Burt Wilder18 has tabulated
the cases of 152 individuals with supernumerary digits,
of which 86 were males, and 39, or less than half,
females ; the remaining 27 being of unknown sex. It
should not, however, be overlooked that women would
more frequently endeavour to conceal a deformity of
this kind than men. Whether the large proportional
number of deaths of the male offspring of man and
apparently of sheep, compared with the female offspring,
before, during, and shortly after birth (see supplement),
has any relation to a stronger tendency in the organs
of the male to vary and thus to become abnormal in
structure or function, I will not pretend to conjecture.

In various classes of animals a few exceptional cases
occur, in which the female instead of the male has
acquired well pronounced secondary sexual characters,
such as brighter colours, greater size, strength, or pug-
nacity. With birds, as we shall hereafter see, there
has sometimes been a complete transposition of the
ordinary characters proper to each sex ; the females
having become the more eager in courtship, the males
remaining comparatively passive, but apparently select-
ing, as we may infer from the results, the more attractive
females. Certain female birds have thus been rendered
more highly coloured or otherwise ornamented, as well
as more powerful and pugnacious than the males, these
characters being transmitted to the female offspring
alone.

It may be suggested that in some cases a double

J DO

process of selection has been carried on ; the males
having selected the more attractive females, and the
latter the more attractive males. This process however,
though it might lead to the modification of both sexes,

13 'Massachusetts Medical Soc' vol. ii. No. 8, 1868, p. 9.

Chap. VIII. SEXUAL SELECTION. 277

would not make the one sex different from the other,
unless indeed their taste for the beautiful differed; but
this is a supposition too improbable in the case of any
animal, excepting man, to be worth considering. There
are, however, many animals, in which the sexes resemble
each other, both being furnished with the same orna-
ments, which analogy would lead us to attribute to the
agency of sexual selection. In such cases it may be
suggested with more plausibility, that there has been a
double or mutual process of sexual selection ; the more
vigorous and precocious females having selected the
more attractive and vigorous males, the latter having
rejected all except the more attractive females. But
from what we know of the habits of animals, this view
is hardly probable, the male being generally eager to
pair with any female. It is more probable that the
ornaments common to both sexes were acquired by one
sex, generally the male, and then transmitted to the off-
spring of both sexes. If, indeed, during a lengthened
period the males of any species were greatly to exceed
the females in number, and then during another
lengthened period under different conditions the reverse
were to occur, a double, but not simultaneous, process
of sexual selection might easily be carried on, by which
the two sexes might be rendered widely different.

We shall hereafter see that many animals exist, of
which neither sex is brilliantly coloured or provided
with special ornaments, and yet the members of both
sexes or of one alone have probably been modified
through sexual selection. The absence of bright tints
or other ornaments may be the result of variations of
the right kind never having occurred, or of the animals
themselves preferring simple colours, such as plain black
or white. Obscure colours have often been acquired
through natural selection for the sake of protection, and

278 THE PRINCIPLES OF Part II.

the acquirement through sexual selection of conspicuous
colours, may have been checked from the danger thus
incurred. But in other cases the males have probably
struggled together during long ages, through brute
force, or by the display of their charms, or by both
means combined, and yet no effect will have been pro-
duced unless a larger number of offspring were left by
the more successful males to inherit their superiority,
than by the less successful males ; and this, as previously
ihewn, depends on various complex contingencies.

Sexual selection acts in a less rigorous manner than
natural selection. The latter produces its effects by the
life or death at all ages of the more or less successful
individuals. Death, indeed, not rarely ensues from the
conflicts of rival males. But generally the less success-
ful male merelv fails to obtain a female, or obtains a
retarded and less vigorous female later in the season,
or, if polygamous, obtains fewer females ; so that they
leave fewer, or less vigorous, or no offspring. In re-
gard to structures acquired through ordinary or natural
selection, there is in most cases, as long as the condi-
tions of life remain the same, a limit to the amount of
advantageous modification in relation to certain special
ends ; but in regard to structures adapted to make one
male victorious over another, either in fighting or in
charming the female, there is no definite limit to the
amount of advantageous modification ; so that as long as
the proper variations arise the work of sexual selection
will go on. This circumstance may partly account for
the frequent and extraordinary amount of variability
presented by secondary sexual characters. Nevertheless,
natural selection will determine that characters of this
kind shall not be acquired by the victorious males,
which would be injurious to them in any high degree,
either by expending too much of their vital powers, or

Chap. VIII. SEXUAL SELECTION. 279

by exposing them to any great danger. The develop-
ment, however, of certain structures — of the horns, for
instance, in certain stags — has been carried to a
wonderful extreme ; and in some instances to an
extreme which, as far as the general conditions of life
are concerned, must be slightly injurious to the male.
From this fact we learn that the advantages which
favoured males have derived from conquering other
males in battle or courtship, and thus leaving a
numerous progeny, have been in the long run greater
than those derived from rather more perfect adaptation
to the external conditions of life. We shall further see,
and this could never have been anticipated, that the
power to charm the female has been in some few in-
stances more important than the power to conquer other
males in battle.

LAWS OF INHERITANCE.

In order to understand how sexual selection has
acted, and in the course of ages has produced conspicuous
results with many animals of many classes, it is neces-
sary to bear in mind the laws of inheritance, as far as
they are known. Two distinct elements are included
under the term " inheritance," namely the transmission
and the development of characters ; but as these
generally go together, the distinction is often over-
looked. We see this distinction in those characters
which are transmitted through the early years of life,
but are developed only at maturity or during old
age. We see the same distinction more clearly with
secondary sexual characters, for these are transmitted
through both sexes, though developed in one alone.
That they are present in both sexes, is manifest when
two species, having strongly-marked sexual characters,
are crossed, for each transmits the characters proper to

280 THE PRINCIPLES OF Part 11.

its own male and female sex to the hybrid offspring of
either sex. The same fact is likewise manifest, when
characters proper to the male are occasionally deve-
loped in the female when she grows old or becomes
diseased ; and so conversely with the male. Again,
characters occasionally appear, as if transferred from
the male to the female, as when, in certain breeds of the
fowl, spurs regularly appear in the young and healthy
females ; but in truth they are simply developed in the
female ; for in every breed each detail in the structure
of the spur is transmitted through the female to her
male offspring. In all cases of reversion, characters
are transmitted through two, three, or many generations,
and are then under certain unknown favourable con-
ditions developed. This important distinction between
transmission and development will be easiest kept in
mind by the aid of the hypothesis of pangenesis, whether
or not it be accepted as true. According to this hypo-
thesis, every unit or cell of the body throws off gemmules
or undeveloped atoms, which are transmitted to the
offspring of both sexes, and are multiplied by self-
division. They may remain undeveloped during the
early years of life or during successive generations ;
their development into units or cells, like those from
wdiich they were derived, depending on their affinity
for, and union with, other units or cells previously
developed in the due order of growth.

Inheritance at Corresponding Periods of Life. — This
tendency is well established. If a new character appears
in an animal whilst young, whether it endures through-
out life or lasts only for a time, it will reappear, as a
general rule, at the same age and in the same manner
in the offspring. If, on the other hand, a new character
appears at maturity, or even during old age, it tends

Chap. VIII. SEXUAL SELECTION. 281

to reappear in the offspring at the same advanced age.
When deviations from this rule occur, the transmitted
characters much oftener appear before than after the
corresponding age. As I have discussed this subject
at sufficient length in another work,19 I will here merely
give two or three instances, for the sake of recalling the
subject to the reader's mind. In several breeds of the
Fowl, the chickens whilst covered with down, the young
birds in their first true plumage and in their adult plum-
age, differ greatly from each other, as well as from their
common parent-form, the Gallus bankiva ; and these
characters are faithfully transmitted by each breed to
their offspring at the corresponding period of life. For
instance, the chickens of spangled Hamburghs, whilst
covered with down, have a few dark spots on the head
and rump, but are not longitudinally striped, as in
many other breeds ; in their first true plumage, " they
"are beautifully pencilled," that is each feather is
transversely marked by numerous dark bars; but in
their second plumage the feathers all become spangled
or tipped with a dark round spot.20 Hence in this
breed variations have occurred and have been trans-
mitted at three distinct periods of life. The Pigeon offers
a more remarkable case, because the aboriginal parent-
species does not undergo with advancing age any change
of plumage, excepting that at maturity the breast
becomes more iridescent ; yet there are breeds which
do not acquire their characteristic colours until they

19 'The Variation of Animals and Plants under Domestication,' vol.
ii. 1868, p. 75. In the latt chapter but one, the provisional hypothesis
of pangenesis, above alluded to, is fully explained.

20 These facts are given on the high authority of a great breeder,
Mr. Teebay, in Tegetmeier's ' Poultry Book,' 1868, p. 158. On the
characters of chickens of different breeds, and on the breeds of the
pigeon, alluded to in the above paragraph, see ' Variation of Animals,'
&c, vol. i. p. 160, 249 ; vol. ii. p. 77.

282 THE PRINCIPLES OF Part II.

have moulted two, three, or four times ; and these
modifications of* plumage are regularly transmitted.

Inheritance at Corresponding Seasons of the Year.
— With animals in a state of nature innumerable
instances occur of characters periodically appearing at
different seasons. We see this with the horns of the
staar, and with the fur of arctic animals which becomes
thick and white during the winter. Numerous birds
acquire bright colours and other decorations during the
breeding-season alone. I can throw but little light on
this form of inheritance from fartts observed under
domestication. Pallas states,21 that in Siberia domestic
cattle and horses periodically become lighter-coloured
during the winter ; and I have observed a similar
marked change of colour in certain ponies in England.
Although I do not know that this tendency to assume a
differently coloured coat durinff different seasons of the
year is transmitted, yet it probably is so, as all shades of
colour are strongly inherited by the horse. Nor is this
form of inheritance, as limited by season, more remark-
able than inheritance as limited by age or sex.

Inheritance as Limited hy Sex. — The equal trans-
mission of characters to both sexes is the commonest
form of inheritance, at least with those animals which
do not present strongly -marked sexual differences, and
indeed with many of these. But characters are not
rarely transferred exclusively to that sex, in which they
first appeared. Ample evidence on this head has been
advanced in my work on Variation under Domestica-

21 'Novae species Quadrupedum e Glirium ordine,' 1778, p. 7. On
the transmission of colour by the horse, see ' Variation of Animals, &c.
under Domestication/ vol. i. p. 21. Also vol. ii. p. 71, for a general
discussion on Inheritance as limited by Sex.

Chap. VIII. SEXUAL SELECTION. 283

tion ; but a few instances may here be given. There
are breeds of the sheep and goat, in which the horns
of the male differ greatly in shape from those of the
female ; and these differences, acquired under domes-
tication, are regularly transmitted to the same sex.
With tortoise-shell cats the females alone, as a general
rule, are thus coloured, the males being rusty-red.
With most breeds of the fowl, the characters proper
to each sex are transmitted to the same sex alone. So
general is this form of transmission that it is an ano-
maly when we see in certain breeds variations trans-
mitted equally to both sexes. There are also certain
sub-breeds of the fowl in which the males can hardly
be distinguished from each other, whilst the females
differ considerably in colour. With the pigeon the
sexes of the parent-species do not differ in any external
character; nevertheless in certain domesticated breeds
the male is differently coloured from the female.22
The wattle in the English Carrier pigeon and the crop
in the Pouter are more highly developed in the male
than in the female ; and although these characters have
been gained through long-continued selection by man,
the difference between the two sexes is wholly due to
the form of inheritance which has prevailed ; for it
has arisen, not from, but rather in opposition to, the
wishes of the breeder.

Most of our domestic races have been formed by the
accumulation of many slight variations ; and as some
of the successive steps have been transmitted to one
sex alone, and some to both sexes, we find in the diffe-
rent breeds of the same species all gradations between
great sexual dissimilarity and complete similarity. In-

32 Dr. Cliapuis, ' Le Pigeon Voyageur Beige,' 1865, p. 87. Boitard
et Corbie, « Les Pigeons de Voliere,' &c, 1824, p. 173.

284 THE PEINCIPLES OF Part II.

stances have already been given with the breeds of the
fowl and pigeon ; and under nature analogous cases are
of frequent occurrence. With animals under domesti-
cation, but whether under nature I will not venture to
say, one sex may lose characters proper to it, and may
thus come to resemble to a certain extent the opposite
sex ; for instance, the males of some breeds of the fowl
have lost their masculine plumes and hackles. On the
other hand the differences between the sexes may be
increased under domestication, as with merino sheep, in
which the ewes have lost their horns. Again, characters
proper to one sex may suddenly appear in the other
sex ; as with those sub-breeds of the fowl in which the
hens whilst young acquire spurs ; or, as in certain
Polish sub-breeds, in which the females, as there is
reason to believe, originally acquired a crest, and sub-
sequently transferred it to the males. All these cases
are intelligible on the hypothesis of pangenesis ; for
they depend on the gemmules of certain units of the
body, although present in both sexes, becoming through
the influence of domestication dormant in the one sex ;
or if naturally dormant, becoming developed.

There is one difficult question which it will be con-
venient to defer to a future chapter ; namely, whether
a character at first developed in both sexes, can be ren-
dered through selection limited in its development to
one sex alone. If, for instance, a breeder observed that
some of his pigeons (in which species characters are
usually transferred in an equal degree to both sexes)
varied into pale blue ; could he by long-continued
selection make a breed, in which the males alone should
be of this tint, whilst the females remained unchanged ?
I will here only say, that this, though perhaps not
impossible, would be extremely difficult ; for the natural
result of breeding from the pale -blue males would be

Chap. VIII. SEXUAL SELECTION. 285

to change his whole stock, including both sexes, into
this tint, If, however, variations of the desired tint
appeared, which were from the first limited in their
development to the male sex, there would not be the
least difficulty in making a breed characterised by the
two sexes being of a different colour, as indeed has been
effected with a Belgian breed, in which the males alone
are streaked with black. In a similar manner, if any
variation appeared in a female pigeon, which was from
the first sexually limited in its development, it would
be easy to make a breed with the females alone thus
characterised ; but if the variation was not thus originally
limited, the process would be extremely difficult, per-
haps impossible.

On the Relation bettveen the period of Development of a
Character and its transmission to one sex or to both sexes.
— Why certain characters should be inherited by both
sexes, and other characters by one sex alone, namely by
that sex in which the character first appeared, is in most
cases quite unknown. We cannot even conjecture why
with certain sub-breeds of the pigeon, black striae, though
transmitted through the female, should be developed in
the male alone, whilst every other character is equally
transferred to both sexes. Why, again, with cats, the
tortoise-shell colour should, with rare exceptions, be
developed in the female alone. The very same cha-
racters, such as deficient or supernumerary digits, colour-
blindness, &c, may with mankind be inherited by the
males alone of one family, and in another family by the
females alone, though in both cases'transmitted through
the opposite as well as the same sex.23 Although we
are thus ignorant, two rules often hold good, namely

Zi Eefercnces are given in my ' Variation of Animals under Domes-
tication,' vol. ii. p. 72.

286 THE PRINCIPLES OF Part II.

that variations which first appear in either sex at a late
period of life, tend to be developed in the same sex
alone ; whilst variations which first appear early in life
in either sex tend to be developed in both sexes. I am,
however, far from supposing that this is the sole de-
termining cause. As I have not elsewhere discussed
this subject, and as it has an important bearing on
sexual selection, I must here enter into lengthy and
somewhat intricate details.

It is in itself probable that any character appearing
at an early age would tend to be inherited equally by
both sexes, for the sexes do not differ much in constitu-
tion, before the power of reproduction is gained. On
the other hand, after this power has been gained and the
sexes have come to differ in constitution, the gemmules
(if I may again use the language of pangenesis) which
are cast off from each varying part in the one sex would
be much more likely to possess the proper affinities
for uniting with the tissues of the same sex, and thus
becoming developed, than with those of the opposite
sex.

I was first led to infer that a relation of this kind
exists, from the fact that whenever and in whatever
manner the adult male has come to differ from the
adult female, he differs in the same manner from the
young of both sexes. The generality of this fact is quite
remarkable : it holds good with almost all mammals,
birds, amphibians, and fishes ; also with many crus-
taceans, spiders and some few insects, namely certain
orthoptera and libellulae. In all these cases the varia-
tions, through the accumulation of which the male ac-
quired his proper masculine characters, must have oc-
curred at a somewhat late period of life ; otherwise the
young males would have been similarly characterised ;
and conformably with our rule, they are transmitted to

Chap. VIII. SEXUAL SELECTION. 287

and developed in the adult males alone. When, on the
other hand, the adult male closely resembles the young
of both sexes (these, with rare exceptions, being- alike),
he generally resembles the adult female ; and in most of
these cases the variations through which the young and
old acquired their present characters, probably occurred
in conformity with our rule during youth. But there is
here room for doubt, as characters are sometimes trans-
ferred to the offspring at an earlier age than that at
which they first appeared in the parents, so that the
parents may have varied wrhen adult, and have trans-
ferred their characters to their offspring whilst young.
There are, moreover, many animals, in which the two
sexes closely resemble each other, and yet both differ
from their young ; and here the characters of the adults
must have been acquired late in life ; nevertheless,
these characters in apparent contradiction to our rule,
are transferred to both sexes. We must not, however,
overlook the possibility or even probability of succes-
sive variations of the same nature sometimes occurring,
under exposure to similar conditions, simultaneously in
both sexes at a rather late period of life ; and in this case
the variations would be transferred to the offspring of
both sexes at a corresponding late age ; and there would
be no real contradiction to our rule of the variations
which occur late in life being transferred exclusively to
the sex in which they first appeared. This latter rule
seems to hold true more generally than the second rule,
namely, that variations which occur in either sex early
in life tend to be transferred to both sexes. As it was
obviously impossible even to estimate in how large a
number of cases throughout the animal kingdom these
two propositions hold good, it occurred to me to inves-
tigate some striking or crucial instances, and to rely
on the result.

288 THE PRINCIPLES OF Part II.

An excellent case for investigation is afforded by the
Deer Family. In all the species, excepting one, the
horns are developed in the male alone, though certainly
transmitted through the female, and capable of occasional
abnormal development in her. In the reindeer, on the
other hand, the female is provided with horns ; so that
in this species, the horns ought, according to our rule,
to appear early in life, long before the two sexes had
arrived at maturity and had come to differ much in
constitution. In all the other species of deer the horns
ought to appear later in life, leading to their develop-
ment in that sex alone, in which they first appeared
in the progenitor of the whole Family. Now in seven
species, belonging to distinct sections of the family and
inhabiting different regions, in which the stags alone
bear horns, I find that the horns first appear at periods
varying from nine months after birth in the roebuck to
ten or twelve or even more months in the stags of the
six other larger species.24 But with the reindeer the
case is widely different, for as I hear from Prof. Nilsson,
who kindly made special enquiries for me in Lapland,
the horns appear in the young animals within four or
five weeks after birth, and at the same time in both
sexes. So that here we have a structure, developed at
a most unusually early age in one species of the family,
and common to both sexes in this one species.

In several kinds of antelopes the males alone are

24 I am much obliged to Mr. Cupples for having made enquiries for
me in regard to the Roebuck and Red Deer of Scotland from Mr.
Robertson, the experienced head-forester to the Marquis of Breadalbane.
In regard to Fallow-deer, I am obliged to Mr. Eyton and others for
information. For the Cervus dices of N. America, see ' Land and Water,'
1868, p. 221 and 254; and for the C. Yirginianus and strongyloceros of
the same continent, see J. D. Caton, in ' Ottawa Acad, of Nat. So.'
1868, p. 13. For Cervus Eldi of Pegu, see Lieut. Beavan, ' Proc.
Zoolog. Soc' 1867, p. 762.

Chap. VIII. SEXUAL SELECTION. 289

provided with horns, whilst in the greater number both
sexes have horns. With respect to the period of de-
velopment, Mr. Blvth informs me that there lived
at one time in the Zoological Gardens a young koodoo
(Ant. strepsiceros), in which species the males alone
are horned, and the young of a closely-allied species,
viz. the eland (Ant. oreas), in which both sexes are
horned. Now in strict conformity with our rule, in the
young male koodoo, although arrived at the age of ten
months, the horns were remarkably small considering
the size ultimately attained by them : whilst in the
young male eland, although only three months old, the
horns were already very much larger than in the koodoo.
It is also worth notice that in the prong-horned antelope,25
in which species the horns, though present in both
sexes, are almost rudimentary in the female, they do not
appear until about five or six months after birth. With
sheep, goats, and cattle, in which the horns are well
developed in both sexes, though not quite equal in size,
they can be felt, or even seen, at birth or soon after-
wards.26 Our rule, however, fails in regard to some
breeds of sheep, for instance merinos, in which the rams
alone are horned ; for I cannot rind on enquiry,27 that

25 Antilocaiyra Americana. Owen, 'Anatomy of Vertebrates,' vol.
iii. p. 627.

2fi I have been assured that the horns of the sheep in North Wales
can always be felt, and are sometimes even an inch in length, at birth.
With cattle Youatt says (' Cattle/ 1834, p. 277) that the prominence of
the frontal bone penetrates the cutis at birth, and that the homy
matter is soon formed over it.

27 I am greatly indebted to Prof. Victor Cams for having made
inquiries for me, from the highest authorities, with respect to the
merino sheep of Saxony. On the Guinea coast of Africa there is a
breed of sheep in which, as with merinos, the rams alone bear horns ;
and Mr. Winwood Eeade informs me that in the one case observed, a
young ram born on Feb. 10th first showed horns on March Gth, so
that in this instance the development of the horns occurred at a later
VOL. I. U

290 THE PRINCIPLES OF Part II.

the horns are developed later in life in this breed than
in ordinary sheep in which both sexes are horned. But
with domesticated sheep the presence or absence of
horns is not a firmly fixed character ; a certain proportion
of the merino ewes bearing small horns, and some of the
rams being hornless ; whilst with ordinary sheep hornless
ewes are occasionally produced.

In most of the species of the splendid family of the
Pheasants, the males differ conspicuously from the
females, and they acquire their ornaments at a rather
late period of life. The eared pheasant (Crossoptilon
auritum), however, offers a remarkable exception, for
both sexes possess the fine caudal plumes, the large ear-
tufts and the crimson velvet about the head ; and I find
on enquiry in the Zoological Gardens that all these
characters, in accordance with our rule, appear very
early in life. The adult male can, however, be distin-
guished from the adult female by one character, namely
by the presence of spurs ; and conformably with our
rule, these do not begin to be developed, as I am assured
by Mr. Bartlett, before the age of six months, and even
at this age, can hardly be distinguished in the two
sexes.28 The male and female Peacock differ con-

period of life, conformably with our rule, than in the Welch sheep, in
which both sexes are horned.

28 In the common peacock (Paro cristatus) the male alone possesses
spurs, whilst botli sexes of the Java peacock (P. muticus) offer the
unusual case of being furnished with spurs. Hence I fully expected
that in the latter species they would have been developed earlier in life
than in the common peacock ; but M. Hegt of Amsterdam informs me,
that with young birds of the previous year, belonging to both species,
compared on April 23rd, 1869, there was no difference in the develop-
ment of the spurs. The spurs, however, were as yet represented merely
by slight knobs or elevations. I presume that I should have been in-
formed if any difference in the rate of development had subsequently
been observed.

Chap. VIII. SEXUAL SELECTION. 291

spicuously from each other in almost every part of their
plumage, except in the elegant head-crest, which is
common to both sexes ; and this is developed very early
in life, long before the other ornaments which are con-
fined to the male. The wild-duck offers an analogous
case, for the beautiful green speculum on the wings
is common to both sexes, though duller and somewhat
smaller in the female, and it is developed early in life,
whilst the curled tail-feathers and other ornaments
peculiar to the male are developed later.29 Between
such extreme cases of close sexual resemblance and
wide dissimilarity, as those of the Crossoptilon and
peacock, many intermediate ones could be given, in
which the characters follow in their order of develop-
ment our two rules.

As most insects emerge from their pupal state in a
mature condition, it is doubtful whether the period of
development determines the transference of their cha-
racters to one or both sexes. But we do not know that
the coloured scales, for instance, in two species of but-
terflies, in one of which the sexes differ in colour, whilst
in the other they are alike, are developed at the same
relative age in the cocoon. Nor do we know whether
all the scales are simultaneously developed on the wings

29 In some other species of the Duck Family the speculum in the
two sexes diners in a greater degree ; but I have not been able to dis-
cover whether its full development occurs later in life in the males of
such species, than in the male of the common duck, as ought to be the
case according to our rule. With the allied Mergus cucullatus we have,
however, a case of this kind : the two sexes differ conspicuously in
general plumage, and to a considerable degree in the speculum, which
is pure white in the male and greyish-white in the female. Now the
young males at first resemble, in all respects, the female, and have a
greyish-white speculum, but this becomes pure white at an earlier age
than that at which the adult male acquires his other more strongly-
marked sexual differences in plumage : see Audubon, ' Ornithological
Biography,' vol. hi. 1835, p. 249-250.

u 2

292 THE PRINCIPLES OF Part II.

of the same species of butterfly, in which certain coloured
marks are confined to one sex, whilst other marks are
common to both sexes. A difference of this kind in the
period of development is not so improbable as it may
at first appear; for with the Orthoptera, which assume
their adult state, not by a single metamorphosis, but by
a succession of moults, the young males of some species
at first resemble the females, and acquire their distinc-
tive masculine characters only during a later moult.
Strictly analogous cases occur during the successive
moults of certain male crustaceans.

We have as yet only considered the transference of
characters, relatively to their period of development, with
species in a natural state ; we will now turn to domes-
ticated animals ; first touching on monstrosities and
diseases. The presence of supernumerary digits, and
the absence of certain phalanges, must be determined
at an early embryonic period- — the tendency to profuse
bleeding is at least congenital, as is probably colour-
blindness— yet these peculiarities, and other similar
ones, are often limited in their transmission to one sex ;
so that the rule that characters which are developed
at an early period tend to be transmitted to both sexes,
here wholly fails. But this rule, as before remarked,
does not appear to be nearly so generally true as the
converse proposition, namely, that characters which
appear late in life in one sex are transmitted exclu-
sively to the same sex. From the fact of the above
abnormal peculiarities becoming attached to one sex,
long before the sexual functions are active, we may
infer that there must be a difference of some kind
between the sexes at an extremely early age. With
respect to sexually-limited diseases, we know too little
of the period at which they originate, to draw any
fair conclusion. Gout, however, seems to fall under

GilAP. VIII. SEXUAL SELECTION. 293

our rule ; for it is generally caused by intemperance
after early youth, and is transmitted from the father
to his sons in a much more marked manner than to his
daughters.

In the various domestic breeds of sheep, goats, and
cattle, the males differ from their respective females
in the shape or development of their horns, forehead,
mane, dewlap, tail, and hump on the shoulders; and
these peculiarities, in accordance with our rule, are not
fully developed until rather late in life. With dogs,
the sexes do not differ, except that in certain breeds,
especially in the Scotch deer-hound, the male is much
larger and heavier than the female ; and as we shall see
in a future chapter, the male goes on increasing in size
to an unusually late period of life, which will account,
according to our rule, for his increased size being trans-
mitted to his male offspring alone. On the other hand,
the tortoise-shell colour of the hair, which is confined
to female cats, is quite distinct at birth, and this case
violates our rule. There is a breed of pigeons in which
the males alone are streaked with black, and the streaks
can be detected even in the nestlings ; but they become
more conspicuous at each successive moult, so that this
case partly opposes and partly supports the rule. With
the English Carrier and Pouter pigeon the full develop-
ment of the wattle and the crop occurs rather late in
life, and these characters, conformably with our rule,
are transmitted in full perfection to the males alone.
The following cases perhaps come within the class pre-
viouslv alluded to, in which the two sexes have varied
in the same manner at a rather late period of life, and
have consequently transferred their new characters to
both sexes at a corresponding late period ; and if so,
such cases are not opposed to our rule. Thus there
are sub-breeds of the pigeon, described by Neumeis-

294 THE PEINCIPLES OF Part II.

ter,30 both sexes of which change colour after moulting
twice or thrice, as does likewise the Almond Tumbler ;
nevertheless these changes, though occurring rather
late in life, are common to both sexes. One variety
of the Canary-bird, namely the London Prize, offers a
nearly analogous case.

With the breeds of the Fowl the inheritance of various
characters by one sex or by both sexes, seems generally
determined by the period at which such characters are
developed. Thus in all the many breeds in which the
adult male differs greatly in colour from the female and
from the adult male parent-species, he differs from the
young male, so that the newly acquired characters must
have appeared at a rather late period of life. On the
other hand with most of the breeds in which the two sexes
resemble each other, the young are coloured in nearly
the same manner as their parents, and this renders it
probable that their colours first appeared early in life.
We have instances of this fact in all black and white
breeds, in which the young and old of both sexes are
alike ; nor can it be maintained that there is something
peculiar in a black or white plumage, leading to its
transference to both sexes ; for the males alone of many
natural species are either black or white, the females
being very differently coloured. With the so-called
Cuckoo sub-breeds of the fowl, in which the feathers are
transversely pencilled with dark stripes, both sexes and
the chickens are coloured in nearly the same manner.
The laced plumage of the Sebright bantam is the same
in both sexes, and in the chickens the feathers are tipped
with black, which makes a near approach to lacing.
Spangled Ham burghs, however, offer a partial exception,

30 'Das Ganze der Taubenzucht,' 1837, s. 21, 24. For the case of
the streaked pigeons, see Dr. Chapuis, 'Le Pigeon Voyageur Beige,'
1865, p. 87.

Chap. VIII. SEXUAL SELECTION. 295

for the two sexes, though not quite alike, resemble each
other more closely than do the sexes of the aboriginal
parent-species, yet they acquire their characteristic
plumage late in life, for the chickens are distinctly
pencilled. Turning to other characters besides colour :
the males alone of the wild parent-species and of most
domestic breeds possess a fairly well developed comb, but
in the young of the Spanish fowl it is largely developed
at a very early age, and apparently in consequence of
this it is of unusual size in the adult females. In the
Game breeds pugnacity is developed at a wonderfully
early age, of which curious proofs could be given ; and
this character is transmitted to both sexes, so that the
hens, from their extreme pugnacity, are now generally
exhibited in separate pens. With the Polish breeds the
bony protuberance of the skull which supports the crest
is partially developed even before the chickens are
hatched, and the crest itself soon begins to grow, though
at first feebly ; 31 and in this breed a great bony protu-
berance and an immense crest characterise the adults of
both sexes.

Finally, from what we have now seen of the relation
which exists in many natural species and domesticated
races, between the period of the development of their
characters and the manner of their transmission — for
example the striking fact of the early growth of the
horns in the reindeer, in which both sexes have horns,
in comparison with their much later growth in the
other species in which the male alone bears horns

31 For full particulars and references on all these points respecting
the several breeds of the Fowl, see ' Variation of Animals and Plants
under Domestication,' vol. i. p. 250, 256. In regard to the higher
animals, the sexual differences which have arisen under domestication
are described in the same work under the head of each species.

296 THE PEINCIPLES OF Part II.

— we may conclude that one cause, though not the sole
cause, of characters being exclusively inherited by one
sex, is their development at a late age. And secondly,
that one, though apparently a less efficient, cause of
characters being inherited by both sexes is their deve-
lopment at an early age, whilst the sexes differ but
little in constitution. It appears, however, that some
difference must exist between the sexes even during an
early embryonic period, for characters developed at this
age not rarely become attached to one sex.

Summary and concluding remarks. — From the fore-
going discussion on the various laws of inheritance, we
learn that characters often or even generally tend to
become developed in the same sex, at the same age,
and periodically at the same season of the year, in
which they first appeared in the parents. But these
laws, from unknown causes, are very liable to change.
Hence the successive steps in the modification of a
species might readily be transmitted in different ways ;
some of the steps being transmitted to one sex,
and some to both; some to the offspring at one age,
and some at all ages. Not only are the laws of inherit-
ance extremely complex, but so are the causes which
induce and govern variability. The variations thus
caused are preserved and accumulated by sexual selec-
tion, which is in itself an extremely complex affair,
depending, as it does, on ardour in love, courage, and \
the rivalry of the males, and on the powers of percep-
tion, taste, and will of the female. Sexual selection will
also be dominated by natural selection for the general
welfare of the species. Hence the manner in which the
individuals of either sex or of both sexes are affected
through sexual selection cannot fail to be complex in
the highest degree.

Chap. VIII. SEXUAL SELECTION. 297

When variations occur late in life in one sex, and are
transmitted to the same sex at the same age, the other
sex and the young are necessarily left unmodified.
When they occur late in life, but are transmitted to
both sexes at the same age, the young alone are left un-
modified. Variations, however, may occur at any period
of life in one sex or in both, and be transmitted to both
sexes at all ages, and then all the individuals of the
species will be similarly modified. In the following
chapters it will be seen that all these cases frequently
occur under nature.

Sexual selection can never act on any animal be-
fore the age for reproduction has arrived. From
the great eagerness of the male it has generally
acted on this sex and not on the females. The males
have thus become provided with weapons for fight-
ing with their rivals, or with organs for discovering
and securely holding the female, or for exciting and
charming her. When the sexes differ in these respects,
it is also, as we have seen, an extremely general law
that the adult male differs more or less from the young
male ; and we may conclude from this fact that the
successive variations, by which the adult male became
modified, have not generally occurred much before the
age for reproduction. Whenever some or many of the
variations have occurred early in life, the young males
will partake in a less or greater degree of the cha-
racters of the adult males. Differences of this kind
between the old and young males may be observed
with many animals, for instance with birds.

It is probable that young male animals have often
tended to vary in a manner which would not only have
been of no use to them at an early age, but would have
been actually injurious, — as in the acquisition of bright
colours, which would have rendered them conspicuous

298 THE PEINCIPLES OF Part II.

to their enemies, or of structures, such as great horns,
which would have expended much vital force in
their development. Variations of this kind occurring
in the young males will almost certainly have been
eliminated through natural selection. With the adult
and experienced males, on the other hand, the advan-
tage derived from the acquisition of such characters,
in their rivalry with other males, will often have
more than counterbalanced exposure to some degree
of clanger.

As variations analogous to those wdrich give to the
male a superiority over other males in fighting, or in
finding, securing, or charming the opposite sex, would,
if they happened to arise in the female, be of no service
to her, they will not have been preserved through
sexual selection in this sex. We have good evidence
that with domesticated animals variations of all kinds
are soon lost through intercrossing and accidental
deaths, if not carefully selected. Consequently, varia-
tions of the above kind, if they chanced to arise in
the female, would be extremely liable to be lost ;
and the females would be left unmodified, as far as
these characters are concerned, excepting in so far
as they were received through transference from the
males. No doubt, if the females varied and trans-
mitted their newly acquired characters to their off-
spring of both sexes, the characters which were ad-
vantageous to the males would be preserved through
sexual selection, although they were of no use to
the females themselves. In this case, both sexes would
be modified in the same manner. But I shall here-
after have to recur to these more intricate contin-
gencies.

Variations occurring late in life, and transmitted to
one sex alone, have incessantly been taken advantage

Chap. VIII. SEXUAL SELECTION. 299

of and accumulated through sexual selection in rela-
tion to the reproduction of the species ; therefore it
appears, at first sight, an unaccountable fact that simi-
lar variations have not frequently been accumulated
through natural selection, in relation to ordinary habits
of life. If this had occurred, the two sexes would fre-
quently have been differently modified, for the sake,
for instance, of capturing prey or of escaping from
clanger. We have already seen and shall hereafter
meet with other instances of differences of this kind
between the two sexes, especially with the lower ani-
mals; but they are rare in the higher classes. We
should, however, bear in mind that the sexes in the
higher classes generally follow the same habits of life ;
and supposing that the males alone varied in a manner
favouring their power of gaining subsistence, &c, and
transmitted such variations to their male offspring
alone, these would acquire an organization superior to
that of the females ; but it is probable that the females,
from having the same general constitution and from
being exposed to the same conditions, would sooner or
later vary in the same manner ; and as soon as this
occurred, the variations would be equally preserved
through natural selection in the two sexes, which would
thus ultimately become like each other. The case is
widely different with variations accumulated through
sexual selection ; for the habits of the two sexes in
relation to the reproductive functions are not the same,
and sexually-transmitted modifications serviceable to
the one sex would in it be preserved, whilst similar
modifications would often be quite useless to the other
sex, and consequently would in this latter soon be lost.

In the following chapters, I shall treat of the
secondary sexual characters in animals of all classes,

300 THE PEINCIPLES OF Part II.

and shall endeavour in each case to apply tbe prin-
ciples explained in the present chapter. The lowest
classes will detain us for a very short time, but the
higher animals, especially birds, must be treated at
considerable length. It should be borne in mind that
for reasons already assigned, I intend to give only a few
illustrative instances of the innumerable structures by
the aid of which the male finds the female, or, when
found, holds her. On the other hand, all structures
and instincts by which the male conquers other males,
and by which he allures or excites the female, will be
fully discussed, as these are in many ways the most
interesting.

Supplement on the proportional members of the two sexes
in animals belonging to various classes.

As no one, as far as I can discover, has paid atten-
tion to the relative numbers of the two sexes through-
out the animal kingdom, I will here give such materials
as I have been able to collect, although they are ex-
tremely imperfect. They consist in only a few instances
of actual enumeration, and the numbers are not very
large. As the proportions are known with certainty on
a large scale in the case of man alone, I will first give
them, as a standard of comparison.

Man. — In England during ten years (from 1857 to
1866) 707,120 children on an annual average have
been born alive, in the proportion of 104*5 males to
100 females. But in 1857 the male births through-
out England were as 105*2, and in 1865 as 104*0 to
100. Looking to separate districts, in Buckinghamshire
(where on an average 5000 children are annually born)

Chap. VIII. SEXUAL SELECTION. 301

the mean proportion of male to female births, during
the whole period of the above ten years, was as 102*8
to 100 ; whilst in N. Wales (where the average annual
births are 12,873) it was as high as 106*2 to 100.
Taking a still smaller district, viz., Rutlandshire (where
the annual births average only 739), in 1864 the male
births were as 114-6, and in 1862 as 97'0 to 100; but
even in this small district the average of the 7385
births during the whole ten years was as 104*5 to 100;
that is in the same ratio as throughout England.32
The proportions are sometimes slightly disturbed by
unknown causes ; thus Prof. Faye states " that in
" some districts of Norway there has been during a
" decennial period a steady deficiency of boys, whilst
" in others the opposite condition has existed." In
France during forty-four years the male to the female
births have been as 106*2 to 100 ; but during this
period it has occurred five times in one department,
and six times in another, that the female births have
exceeded the males. In Russia the average proportion
is as high as 108*9 to 100.33 It is a singular fact that
with Jews the proportion of male births is decidedly
larger than with Christians : thus in Prussia the propor-
tion is as 113, in Breslau as 114, and in Livonia as 120
to 100 ; the Christian births in these countries being
the same as usual, for instance, in Livonia as 104 to
100.34 It is a still more singular fact that in different
nations, under different conditions and climates, in
Naples, Prussia, Westphalia, France and England, the

32 ' Twenty-ninth Annual Report of the Registrar-General for 1866.'
In this report (p. xii) a special decennial table is given.

33 For Norway unci Russia, see abstract of Pro!'. Faye's researches,
in ' British and Foreign Medieo-Cliirurg. Review/ April, 1867, p. 343,
345. For France, the ' Annuaire pour lAn 1867,' p. 213,

34 In regard to the Jews, see M. Thury, ' La Loi de Production dcs
Sexes,' 1863, p. 25.

302 THE PRINCIPLES OF Part II.

excess of male over female births is less when they are
illegitimate than when legitimate.35

In various parts of Europe, according to Prof. Faye
and other authors, "a still greater preponderance of

males would be met with, if death struck both sexes

in equal proportion in the womb and during birth.
" But the fact is, that for every 100 still-born females,
" Ave have in several countries from 134*6 to 144*9
" still-born males." Moreover during the first four or
five years of life more male children die than females ;
" for example in England, during the first year, 126
" boys die for every 100 girls, — a proportion wrhich in
" France is still more unfavourable."36 As a consequence
of this excess in the death-rate of male children, and of
the exposure of men when adult to various dangers, and of
their tendency to emigrate, the females in all old-settled
countries, where statistical records have been kept,37 are
found to preponderate considerably over the males.

It has often been supposed that the relative ages
of the parents determine the sex of the offspring ;
and Prof. Leuckart38 has advanced what he considers

35 Babbage, • Edinburgh Journal of Science,' 1829, vol. i. p. 88 ; also
p. 90, on still-born children. On illegitimate children in England,
see ' Keport of Kegistrar-General for 1866,' p. xv.

36 'British and Foreign Medico-Chirurg. Keview,' April, 1867, p.
343. Dr. Stark also remarks (' Tenth Annual Keport of Births, Deaths,
&c, in Scotland,' 1867, p. xxviii) that "'lhese examples may suffice
" to shew that, at almost every stage of life, the males in Scotland
" have a greater liability to death and a higher death-rate than the
" females. The fact, however, of this peculiarity being most strongly
" developed at that infantile period of life when the dress, food, and
" general treatment of both sexes are alike, seems to prove that the
" higher male death-rate is an impressed, natural, and constitutional
" peculiarity due to sex alone."

37 "With the savage Guaranys of Paraguay, according to the accurate
Azara ('Voyages dans l'Ame'rique me'rid.' torn. ii. 1809, p. 60, 179),
the women in proportion to the men are as 14 to 13.

38 Leuckart (in Wagner, ' Handworterbuch der Phys.' B. iv. 1853,
s. 774.

Chap. VIII. SEXUAL SELECTION. 303

sufficient evidence, with respect to man and certain
domesticated animals, to shew that this is one impor-
tant factor in the result. So again the period of
impregnation has been thought to be the efficient cause ;
but recent observations discountenance this belief.
Again, with mankind polygamy has been supposed to
lead to the birth of a greater proportion of female
infants ; but Dr. J. Campbell 39 carefully attended to
this subject in the harems of Siam, and he concludes
that the proportion of male to female births is the
same as from monogamous unions. Hardly any animal
has been rendered so highly polygamous as our English
race-horses, and we shall immediately see that their
male and female offspring are almost exactly equal in
number.

Horses. — Mr. Tegetmeier has been so kind as to tabulate for me
from the ' Eacing Calendar' the births of race-horses during a period
of twenty-one years, viz. from 1846 to 1867 ; 1849 being omitted,
as no returns were that year published. The total births have
been 25,560,40 consisting of 12,763 males and 12,797 females, or in
the proportion of 99*7 males to 100 females. As these numbers are
tolerably large, and as they are drawn from all parts of England,
during several years, we may with much confidence conclude that
with the domestic horse, or at least with the race-horse, the two
sexes are produced in almost equal numbers. The fluctuations in
the proportions during successive years are closely like those which
occur with mankind, when a small and thinly-populated area is con-
sidered : thus in 1856 the male horses were as 107 '1, and in 1867
as only 92*6 to 100 females. In the tabulated returns the propor-
tions vary in cycles, for the males exceeded the females during six
successive years ; and the females exceeded the males during two

39 Anthropological Eeview, April, 1870, p. cyiii.

40 During the last eleven years a record has been kept of the number of
mares which have proved barren or prematurely slipped their foals ; and it
deserves notice, as shewing how infertile these highly-nurtured and rather
closely-interbred animals have become, that not far from one-third of the
mares failed to produce living foals. Thus during 1866, 809 male colts
and 816 female colts were born, and 743 mares failed to produce offspring.
During 1867, 836 males and 902 females were born, and 794 mares failed.

304 THE PRINCIPLES QF Part II.

periods each of four years : this, however, may be accidental ; at
least I can detect nothing; of the kind with man in the decennial
table in the Registrar's Report for 1866. I may add that certain
mares, and this holds good with certain cows and with women,
tend to produce more of one sex than of the other ; Mr. Wright of
Yeldersley House, informs me that one of his Arab mares, though
put seven times to different horses, produced seven fillies.

Dogs. — During a period of twelve years, from 1857 to 1868, the
births of a large number of greyhounds, throughout England, have
been sent to the ' Field ' newspaper ; and I am again indebted to
Mr. Tegetmeier for carefully tabulating the results. The recorded
births have been 6878, consisting of 3605 males and 3273 females,
that is, in the proportion of HO'l males to 100 females. The
greatest fluctuations occurred in 1864, when the proportion was as
95*3 males, and in 1867, as 116'3 males to 100 females. The above
average proportion of 110*1 to 100 is probably nearly correct in the
case of the greyhound, but whether it would hold with other domes-
ticated breeds is in some degree doubtful. Mr. Cupples has enquired
from several great breeders of dogs, and finds that all without
exception believe that females are produced in excess ; he suggests
that this belief may have arisen from females being less valued and
the consequent disappointment producing a stronger impression on
the mind.

Sheep. — The sexes of sheep are not ascertained by agriculturists
until several months after birth, at the period when the males are
castrated ; so that the following returns do not give the proportions
at birth. Moieover, I find that seveial great breeders in Scotland,
who annually raise some thousand sheep, are firmly convinced that
a larger proportion of males than of females die during the first one
or two years ; therefore the proportion of males would be somewhat
greater at birth than at the age of castration. This is a remarkable
coincidence with what occurs, as we have seen, with mankind, and
both cases probably depend on some common cause. I have re-
ceived returns from four gentlemen in England who have bred low-
land sheep, chiefly Leicesters, during the last ten or sixteen years ;
they amount altogether to 8965 births, consisting of 4407 males
and 4558 females ; that is in the proportion of 96*7 males to 100
females. With respect to Cheviot and black-faced sheep bred in
Scotland, I have received returns from six breeders, two of them on
a large scale, chiefly for the years 1867-1869, but some of the
returns extending back to 1862. The total number recorded
amounts to 50,685, consisting of 25,071 males and 25,614 females,
or in the proportion of 97'9 males to 100 females. If we take the
English and Scotch returns together, the total number amounts

Chap. VIII. SEXUAL SELECTION. 305

to 59,650, consisting of 29,478 males and 30,172 females, or as
97*7 to 100. So that with sheep at the age of castration the females
are certainly in excess of the males ; but whether this would hold
good at birth is doubtful, owing to the greater liability in the males
to early death.41

Of Cattle I have received returns from nine gentlemen of 982
births, too few to be trusted ; these consisted of 477 bull-calves and
505 cow-calves ; i.e. in the proportion of 944 males to 100 females.
The Rev. W. D. Fox informs me that in 1867 out of 34 calves born
on a farm in Derbyshire only one was a bull. Mr. Harrison Weir
writes to me that he has enquired from several breeders of Pigs, and
most of them estimate the male to the female births as about
7 to 6. This same gentleman has bred Babbits for many years,
and has noticed that a far greater number of bucks are produced
than does.

Of mammalia in a state of nature I have been able to learn very
little. In regard to the common rat, I have received conflicting
statements. Mr. E. Elliot of Laighwood, informs me that a rat-
catcher assured him that he had always found the males in
great excess, even with the young in the nest. In consequence
of this, Mr. Elliot himself subsequently examined some hundred
old ones, and found the statement true. Mr. F. Buckland ha:
bred a large number of white rats, and he also believes that the
males greatly exceed the females. In regard to Moles, it is said
that " the males are much more numerous than the females ;"42
and as the catching of these animals is a special occupation, the
statement may perhaps be trusted. Sir A. Smith, in describing
an antelope of S. Africa43 (Kobus ettijosiprymnus), remarks, that
in the herds of this and other species, the males are few in number
compared with the females : the natives believe that they are born
in this proportion ; others believe that the younger males are ex-
pelled from the herds, and Sir A. Smith says, that though he has
himself never seen herds consisting of young males alone, others
affirm that this does occur. It appears probable that the young
males when expelled from the herd, would be likely to fall a prey
to the many beasts of prey of the country.

s

41 I am much indebted to Mr. Cupples for having procured for me the
above returns from Scotland, as well as some of the following returns on
cattle. Mr. R. Elliot, of Laighwood, first called my attention to the pre-
mature deaths of the males, — a statement subsequently confirmed by Mr.
Aitchison and others. To this latter gentleman, and to Mr. Payan, I owe
my thanks for the larger returns on sheep.

42 Bell, < History .of British Quadrupeds,' p. 100.

43 * Illustrations of the Zoology of S. Africa,' 1849, pi. 29.

VOL. I. X

306 THE PRINCIPLES OF Part II.

BIKDS.

With respect to the Fowl, I have received only one account,
namely, that out of 1001 chickens of a highly-bred stock of Cochins,
reared during eight years by Mr. Stretch, 487 proved males and 514
females : i. e. as 94'7 to 100. In regard to domestic pigeons there
is good evidence that the males are produced in excess, or that their
lives are longer; for these birds invariably pair, and single males,
as Mr. Tegetmeier informs me, can always be purchased cheaper
than females. Usually the two birds reared from the two eggs
laid in the same nest consist of a male and female ; but Mr. Harrison
Weir, who has been so large a breeder, says that he has often bred
two cocks from the same nest, and seldom two hens ; moreover the
hen is generally the weaker of the two, and more liable to perish.

With respect to birds in a state of nature, Mr. Gould and others44
are convinced that the males are generally the more numerous; and
as the young males of many species resemble the females, the latter
would naturally appear to be the most numerous. Large numbers
of pheasants are reared by Mr. Baker of Leadenhall from eggs laid
by wild birds, and he informs Mr. Jenner Weir that four or five
males to one female are generally produced. An experienced ob-
server remarks45 that in Scandinavia the broods of the capercailzie
and black-cock contain more males than females; and that with the
Dal-ripa (a kind of ptarmigan) more males than females attend the
hks or places of courtship ; but this latter circumstance is accounted
for by some observers by a greater number of hen birds being killed
by vermin. From various facts given by White of Selborne,46 it
seems clear that the males of the partridge must be in considerable
excess in the south of England ; and I have been assured that this
is the case in Scotland. Mr. Weir on enquiring from the dealers
who receive at certain seasons large numbers of ruffs (Machetes
puynax) was told that the males are much the most numerous.
This same naturalist has also enquired for me from the bird-
catchers, who annually catch an astonishing number of various small
species alive for the London market, and he was unhesitatingly
answered by an old and trustworthy man, that with the chaffinch
the males are in large excess ; he thought as high as 2 males to

41 Brehm (' Illust. Thievleben,' B. iv. s. 990) comes to the same con-
clusion.

45 On the authority of L. Lloyd, 'Game Birds of Sweden,' 1867, p. 12,
132.

4,5 'Nat. Hist, of Selbourne,' letter xxix. edit, of 1825, vol. i. p. 139.

Chap. VIII. SEXUAL SELECTION. 307

1 female, or at least as high as 5 to 3.47 The males of the black-
bird, he likewise maintained, were by far the most numerous, whe-
ther caught by traps or by netting at night. These statements
may apparently be trusted, because the same man said that the
sexes are about equal with the lark, the twite (Linaria montana),
and goldfinch. On the other hand he is certain that with the
common linnet, the females preponderate greatly, but unequally
during different years ; during some years he has found the females
to the males as four to one. It should, however, be borne in mind,
that the chief season for catching birds does not begin till Sep-
tember, so that with some species partial migrations may have begun,
and the flocks at this period often consist of hens alone. Mr. Salvin
paid particular attention to the sexes of the humming-birds in
Central America, and he is convinced that with most of the species
the males are in excess ; thus one year he procured 204 specimens
belonging to ten species, and these consisted of 166 males and of
38 females. With two other species the females were in excess :
but the proportions apparently vary either during different seasons
or in different localities ; for on one occasion the males of Cam-
pylopterus hemileucurus were to the females as five to two, and
on another occasion48 in exactly the reversed ratio. As bearing on
this latter point, I may add, that Mr. Powys found in Corfu and
Epirus the sexes of the chaffinch keeping apart, and "the females
" by far the most numerous ;" whilst in Palestine Mr. Tristram
found " the male flocks appearing greatly to exceed the female in
"number."49 So again with the Quiscalus major, Mr. Gr. Taylor50
says, that in Florida there were " very few females in proportion to
" the males," whilst in Honduras the proportion was the other way,
the species there having the character of a polygamist.

FISH.

With Fish the proportional numbers of the sexes can be ascertained
only by catching them in the adult or nearly adult state; and there

47 Mr. Jenner Weir received similar information, on making enquiries
during the following year. To shew the number of chaffinches caught, I
may mention that in 1869 there was a match., between two experts; and
one man caught in a day 62, and another 40, male chaffinches. The greatest
number ever caught by one man in a single day was 70.
^ 18 'Ibis,' vol. ii. p. 260, as quoted in Gould's ' Trochilidac,' 1861, p. 52.
For the foregoing proportions, I am indebted to Mr. Salvin for a table of
his results.

•,9 'Ibis,' 1860, p. 137 ; and 1867, p. 369.

50 ' Ibis,' 1862, p. 137.

x 2

308 THE PRINCIPLES OF Part II.

are many difficulties in arriving at any just conclusion.51 Infertile
females might readily be mistaken for males, as Dr. Giintlier has
remarked to me in regard to trout. With some species the males
are believed to die soon after fertilising the ova. With many species
the males are of much smaller size than the females, so that a large
number of males would escape from the same net by which the
females were caught. M. Carbonnier,52 who has especially attended
to the natural history of the pike (Esox lucius) states that many
males, owing to their small size, are devoured by the larger females ;
and he believes that the males of almost all fish are exposed from
the same cause to greater danger than the females. Nevertheless
in the few cases in which the proportional numbers have been
actually observed, the males appear to be largely in excess. Thus
Mr. R. Buist, the superintendent of the Stormont field experi-
ments, says that in 1865, out of 70 salmon first landed for the
purpose of obtaining the ova, upwards of 60 were males. In
.1867 he again " calls attention to the vast disproportion of the
" males to the females. We had at the outset at least ten males
" to one female." Afterwards sufficient females for obtaining ova
were procured. He adds, " from the great proportion of the
" males, they are constantly fighting and tearing each other on the
" spawning-beds."53 This disproportion, no doubt, can be accounted
for in part, but whether wholly is very doubtful, by the males
ascending the rivers before the females. Mr. F. Buckland remarks
in regard to trout, that " it is a curious fact that the males prepon-
" derate very largely in number over the females. It invariably
" happens that when the first rush of fish is made to the net, there
" will be at least seven or eight males to one female found captive.
" I cannot quite account for this ; either the males are more numer-
" ous than the females, or the latter seek safety by concealment
" rather than flight." He then adds, that by carefully searching the
banks, sufficient females for obtaining ova can be found.54 Mr. H.
Lee informs me that out of 212 trout, taken for this purpose in Lord
Portsmouth's park, 150 were males and 62 females.

With the Cyprinida} the males likewise seem to be in excess ;
but several members of this Family, viz., the carp, tench, bream
and minnow, appear regularly to follow the practice, rare in the

51 Leuckart quotes Bloch (Wagner, ' Handworterbuch der Phys.' B. iv .
1853, s. 775), that with fish there are twice as many males as females.

52 Quoted in the 'Farmer,' March 18, 1869, p. 369.

53 'The Stormontfield Piscicultural Experiments,' 1866, p. 23. The
' Field ' newspaper, June 29th, 1867.

54 ' Land and Water,' 1868, p. 41.

Chap. VIII. SEXUAL SELECTION. 30 D

animal kingdom, of polyandry ; for the female whilst spawning is
always attended by two males, one on each side, and in the case of
the bream by three or four males. This fact is so well known, that
it is always recommended to stock a pond with two male tenches
to one female, or at least with three males to two females. With
the minnow, an excellent observer states, that on the spawning-
beds the males are ten times as numerous as the females ; when a
female comes amongst the males, " she is immediately pressed closely
" by a male on each side ; and when they have been in that situa-
tion for a time, are superseded by other two males." 55

INSECTS.

In this class, the Lepidoptera alone afford the means of judging
of the proportional numbers of the sexes ; for they have been col-
lected with special care by many good observers, and have been
largely bred from the egg or caterpillar state. I had hoped that
some breeders of silk-moths might have kept an exact record, but
after writing to France and Italy, and consulting various treatises,
I cannot find that this has ever been done. The general opinion
appears to be that the sexes are nearly equal, but in Italy as I hear
from Professor Canestrini, many breeders are convinced that the
females are produced in excess. The same naturalist, however,
informs me, that in the two yearly broods of the Ailanthus silk-
moth {Borribyx cynthia), the males greatly preponderate in the
first, whilst in the second the two sexes are nearly equal, or the
females rather in excess.

In regard to Butterflies in a state of nature, several observers
have been much struck by the apparently enormous preponderance
of the males.56 Thus Mr. Bates,57 in speaking of the species, no
less than about a hundred in number, which inhabit the Upper
Amazons, says that the males are much more numerous than the
females, even in the proportion of a hundred to one. In North
America, Edwards, who had great experience, estimates in the
genus Papilio the males to the females as four to one ; and Mr.

55 Yarrell, ' Hist. British Fishes,' vol. i. 18-86, p. 307; on the Cyprians
carjrio, p. 331 ; on the Tinea vulgaris, p. 331 ; on the Abramis brama, p.
336. See, for the minnow (Leuciscus phoxinus), ' Loudon's Mag. of Nat.
Hist.' vol. v. 1832, p. 682.

56 Leuckart quotes Meinecke (Wagner, ' Handworterbuch der Phys.'
B. iv. 1853, s. 775) that with Butterflies the males are three or four times
as numerous as the females.

57 ' The Naturalist on the Amazons,' vol. ii. 1863, p. 228, 347.

310 THE PRINCIPLES OF Part II.

Walsh, who informed me of this statement, says that with P.
turnus this is certainly the case. Jn South Africa, Mr. R. Trimen
found the males in excess in 19 species ;58 and in one of these,
which swarms in open places, he estimated the number of males as
fifty to one female. With another species, in which the males are
numerous in certain localities, he collected during seven years only
five females. In the island of Bourbon, M. Maillard states that
the males of one species of Papilio are twenty times as numerous
as the females.59 Mr. Trimen informs me that as far as he has
himself seen, or heard from others, it is rare for the females of any
butterfly to exceed in number the males ; but this is perhaps the
case with three South African species. Mr. Wallace60 states that
the females of Ornithoptera crcesus, in the Malay archipelago, are
more common and more easily caught than the males ; but this is
a rare butterfly. I may here add, that in Hyperythra, a genus of
moths, Guene'e says, that from four to five females are sent in
collections from India for one male.

When this subject of the proportional numbers of the sexes of
insects was brought before the Entomological Societ}^61 it was
generally admitted that the males of most Lepidoptera, in the
adult or imago state, are caught in greater numbers than the
females ; but this fact was attributed by various observers to the
more retiring habits of the females, and to the males emerging
earlier from the cocoon. This latter circumstance is well known to
occur with most Lepidoptera, as well as with other insects. So
that, as M. Personnat remarks, the males of the domesticated
Bombyx Yamamai, are lost at the beginning of the season, and
the females at the end, from the want of mates.62 I cannot how-
ever persuade myself that these causes suffice to explain the great
excess of males in the cases, above ' given, of butterflies which are
extremely common in their native countries. Mr. Stainton, who
has paid such close attention during many years to the smaller
moths, informs me that when he collected them in the imago state,
he thought that the males were ten times as numerous as the
females, but that since he has reared them on a large scale from the
caterpillar state, he is convinced that the females are the most

58 Four of these cases are given by Mr. Trimen in his ' Pdiopalocera
Africse Australis.'

50 Quoted by Trimen, ' Transact. Ent. Soc' vol. v. part iv. 1866, p. 330.

60 ' Transact. Linn. Soc' vol. xxv. p. 37.

61 ' Proc. Entomolog. Soc' Feb. 17th, 1868.

6" Quoted by Dr. Wallace in ' Proc. Ent. Soc' 3rd series, vol. v. 1867,
p. 487.

Chap. VIII. SEXUAL SELECTION. 311

numerous. Several entomologists concur in this view. Mr. Double-
day, however, and some others, take an oj3posite view, and are con-
vinced that they have^ reared from the egg and caterpillar states a
larger proportion of males than of females.

Besides the more active habits of the males, their earlier emerg-
ence from the cocoon, and their frequenting in some cases more
open stations, other causes may be assigned for an apparent or real
difference in the proportional numbers of the sexes of Lepidop-
tera, when captured in the imago state, and when reared from the
egg or caterpillar state. It is believed by many breeders in Italy,
as 1 hear from Professor Canestrini, that the female caterpillar of
the silk-moth suffers more from the recent disease than the male ;
and Dr. Staudinger informs me that in rearing Lepidoptera more
females die in the cocoon than males. With many species the
female caterpillar is larger than the male, and a collector would
naturally choose the finest specimens, and thus unintentionally
collect a larger number of females. Three collectors have told me
that this was their practice; but Dr. Wallace is sure that most
collectors take all the specimens which they can find of the rarer
kinds, which alone are worth the trouble of rearing. Birds when
surrounded by caterpillars would probably devour the largest ; and
Professor Canestrini informs me that in Italy some breeders believe,
though on insufficient evidence, that in the first brood of the
Ailanthus silk-moth, the wasps destroy a larger number of the
female than of the male caterpillars. Dr. Wallace further remarks
that female caterpillars, from being larger than the males, require
more time for their development and consume more food and mois-
ture ; and thus they would be exposed during a longer time to
danger from ichneumons, birds, &c, and in times of scarcity would
perish in greater numbers. Hence it appears quite possible that,
in a state of nature, fewer female Lepidoptera may reach maturity
than males ; and for our special object we are concerned with the
numbers at maturity, when the sexes are ready to propagate their
kind.

The manner in which the males of certain moths congregate in
extraordinary numbers round a single female, apparently indicates
a great excess of males, though this fact may perhaps be accounted
for by the earlier emergence of the males from their cocoons. Mr.
Stainton informs me that from twelve to fwenty males may often be
seen congregated round a female Elacliista rufocinerea. Jt is well
known that if a virgin Lasiocainpa quercus or iiaturnia carpini
be exposed in a cage, vast numbers of males collect round her,
and if confined in a room will even come down the chimney to her.

.312 THE PRINCIPLES OF Part II.

Mr. Doubleday believes that he has seen from fifty to a hundred
males of both these species attracted in the course of a single day
by a female under confinement. Mr. Trim en exposed in the Isle
of Wight a box in which a female of the Lasiocampa had been
confined on the previous day, and five males soon endeavoured
to gain admittance. M. "Verreaux, in Australia, having placed
the female of a small Bombyx in a box in his pocket, was fol-
lowed by a crowd of males, so that about 200 entered the house
with him.63

Mr. Doubleday has called my attention to Dr. Standi nger's64 list
of Lepidoptera, which gives the prices of the males and females of
300 species or well-marked varieties of (Ehopalocera) butterflies.
The prices for both sexes of the very common species are of
course the same ; but with 114 of the rarer species they differ ; the
males being in all cases, excepting one, the cheapest. On an ave-
rage of the prices of the 113 species, the price of the male to that
of the female is as 100 to 149 ; and this apparently indicates that
inversely the males exceed the females in number in the same
proportion. About 2000 species or varieties of moths (Heterocera)
are catalogued, those with wingless females being here excluded on
account of the difference in habits of the two sexes : of these 2000
species, 141 differ in price according to sex, the males of 130 being
cheaper, and the males of only 11 being dearer than the females.
The average price of the males of the 130 species, to that of the
females, is as 100 to 143. With respect to the butterflies in this
priced list, Mr. Doubleday thinks (and no man in England has had
more experience), that there is nothing in the habits of the species
which can account for the difference in the prices of the two sexes,
and that it can be accounted for only by an excess in the numbers of
the males. But I am bound to add that Dr. Staudinger himself, as
he informs me, is of a different opinion. He thinks that the less
active habits of the females and the earlier emergence of the males
will account for his collectors securing a larger number of males than
of females, and consequently for the lower prices of the former.
With respect to specimens reared from the caterpillar-state, Dr.
Staudinger believes, as previously stated, that a greater number of
females than of males die under confinement in the cocoons. He
adds that with certain species one sex seems to preponderate over
the other during certain years.

Of direct observations on the sexes of Lepidoptera, reared either

63 Blanchard, ' Metamorphoses, Mceurs des Insectes,' 1868, p. 225-226,
H4 ' Lepidopteren-Doubblettren Liste,' Berlin, No. x. 1866,

Chap. VIII. SEXUAL SELECTION. 313

from eggs or caterpillars, I have received only the few following
crises : —

Males, j Females.

The Rev. J. Hellins 65 of Exeter reared, during 1868,

imagos of 73 species, which consisted of .. .. 153 137

Mr. Albert Jones of Eltham reared, during 1868, im-
agos of 9 species, which consisted of 159 126

During 1869 he reared imagos from 4 species, consist-
ing of 114 112

Mr. Buckler of Emsworth, Hants, during 1869, reared

imagos from 74 species, consisting of 180 169

Dr. Wallace of Colchester reared from one brood of

Bombyx cynthia 52 48

Dr. Wallace raised, from cocoons of Bombyx Pernyisent

from China, during 1869 224 123

Dr. Wallace raised, during 1868 and 1869, from two

lots of cocoons of Bombyx yama-mai 52 46

Total 934 761

So that in these eight lots of cocoons and eggs, males were pro-
duced in excess. Taken together the proportion of males is as
122*7 to 100 females. But the numbers are hardly large enough
to be trustworthy.

On the whole, from the above various sources of evidence, all
pointing to the same direction, 1 infer that with mo.-t species of
Lepidoptera, the males in the imago state generally exceed the
females in number, whatever the proportions may be at their first
emergence from the egg.

\Y ith reference to the other Orders of insects, I have been able
to collect very little reliable information. With the stag-beetle
{Lucanus cervus) " the males appear to be much more numerous
" than the females ;" but when, as Cornelius remarked during 1867,
an unusual number of these b< etles appeared in one part of Ger-
many, the females appeared to exceed the males as six so ©ne.
With one of the Elateridas, the males are said to be much more
numerous than the females, and " two or three are often found
" united with one female ,66 so that here polyandry seems to prevail.

65 This naturalist has been so kind as to send me some results from
former years, in which the females seemed to preponderate ; but so many

of the figures were estimates, that I found it impossible to tabulate them. ^

66 Giinther's 'Record of Zoological Literature,' 1867, p. 260. On^th^'N
excess of female Lucanus, ibid. p. 250. On the males of Lucanus in Etftp,^*-
land, Westwood, ' Modern Class, of Insects,' vol. i. p. 187. On the SingonKi^ q°
ibid. p. 172. />

314 THE PRINCIPLES OF Part II.

With Siagonium (Staphylinidae), in which the males are furnished
with horns, " the females are far more numerous than the opposite
" sex." Mr. Janson stated at the Entomological Society that the
females of the bark-feeding Tomicus villosus are so common as to
be a plague, whilst the males are so rare as to be hardly known.
In other Orders, from unknown causes, but apparently in some in-
stances owing to parthenogenesis, the males of certain species have
never been discovered or are excessively rare, as with several of the
Cynipida?.67 In all the gall-making Cynipida? known to Mr. Walsh,
the females are four or five times as numerous as the males ; and so
it is, as he informs me, with the gall-making Cecidomyiia; (Diptera).
With some common species of Saw-flies (Tenthredime) Mr. ¥.
Smith has reared hundreds of specimens from larvae of all sizes,
but has never reared a single male : on the other hand Curtis says,68
that with certain species (Athalia), bred by him, the males to the
females were as six to one ; whilst exactly the reverse occurred with
the mature insects of the same species caught in the fields. With
the Neuroptera, Mr. Walsh states that in many, but by no means
in all, the species of the Odonatous groups (Ephemerina), there is a
great overplus of males : in the genus Heta?rina, also, the males are
generally at least four times as numerous as the females. In certain
species in the genus Gomphus the males are equally numerous,
whilst in two other species, the females are twice or thrice as
numerous as the males. In some European species of Psocus thou-
sands of females may be collected without a single male, whilst
with other species of the same genus both sexes are common.69 In
England, Mr. MacLachlan has captured hundreds of the female
Apatania muliebris, but has never seen the male ; and of Boreus
hyemalis only four or five males have been here seen.70 With most
of these species (excepting, as I have heard, with the Tenthredina3)
there is no reason to suppose that the females are subject to parthe-
nogenesis ; and thas we see how ignorant we are on the causes of the
apparent discrepancy in the proportional numbers of the two sexes.
In the other Classes of the Articulata I have been able to collect
still less information. With Spiders, Mr. Blackwall, who has care-
fully attended to this class during many years, writes to me that
the males from their more erratic habits are more commonly seen,

67 Walsh, in 'The American Entomologist,' vol. i. 1869, p. 103. F. Smith,
' Record of Zoological Literature,' 1867, p. 328.

68 ' Farm Insects,' p. 45-46.

69 ' Observations on N. American Neuroptera,' bv H. Hagen and B. D.
Walsh, 'Proc. Ent. Soc. Philadelphia,' Oct. 1863, p. 168, 223, 239.

70 ' Proc. Ent. Soc. London,' Feb. 17, 1868.

Chap. VJII. SEXUAL SELECTION. 315

and therefore appear to be the more numerous. This is actually the
case with a few species ; but he mentions several species in six
genera, in which the females appear to be much more numerous
than the males.71 The small size of the males in comparison with
the females, which is sometimes carried to an extreme degree, and
their widely different appearance, may account in some instances
for their rarity in collections.72

Some of the lower Crustaceans are able to propagate their kind
asexually, and this will account for the extreme rarity of the males.
With some other forms (as with Tanais and Cypris) there is reason
to believe, as Fritz Muller informs me, that the male is much shorter-
lived than the female, which, supposing the two sexes to be at first
equal in number, would explain the scarcity of the males. On the
other hand this same naturalist has invariably taken, on the shores
of Brazil, far more males than females of the Diastylida? and of
Cypridina ; thus with a species in the latter genus, 63 specimens
caught the same day, included 57 males ; but he suggests that this
preponderance may be due to some unknown difference in the habits
of the two sexes. With one of the higher Brazilian crabs, namely
a Gelasimus, Fritz Muller found the males to be more numerous
than the females. The reverse seems to be the case, according to
the large experience of Mr. C. Spence Bate, with six common British
crabs, the names of which he has given me.

On the Tower of Natural Selection to regulate the pro-
portional Numbers of the Sexes, and General Fertility. —
In some peculiar cases, an excess in the number of one
sex over the other might be a great advantage to a
species, as with the sterile females of social insects, or
with those animals in which more than one male is
requisite to fertilise the female, as with certain cirri-
pecles and perhaps certain fishes. An inequality be-
tween the sexes in these cases might have been acquired
through natural selection, but from their rarity they
need not here be further considered. In all ordinary

71 Another great authority in this class, Prof. Thorell of Upsala (' On
European Spiders,' 1869-70, part i. p. 205) speaks as if female spiders were
generally commoner than the males.

"- See, on this subject, Mr. Pickard-Cambridge, as quoted in 'Quarterly
Journal of Science,' 1868, p. 429.

316 THE PRINCIPLES OF Part II.

cases an inequality would be no advantage or disad-
vantage to certain individuals more than to others ; and
therefore it could hardly have resulted from natural
selection. We must attribute the inequality to the
direct action of those unknown conditions, which with
mankind lead to the males being born in a somewhat
larger excess in certain countries than in others, or
which cause the proportion between the sexes to differ
slightly in legitimate and illegitimate births.

Let us now take the case of a species producing from
the unknown causes just alluded to, an excess of one
sex — we will say of males — these being superfluous and
useless, or nearly useless. Could the sexes be equalised
through natural selection ? We may feel sure, from all
characters being variable, that certain pairs would pro-
duce a somewhat less excess of males over females than
other pairs. The former, supposing the actual number
of the offspring to remain constant, would necessarily
produce more females, and would therefore be more pro-
ductive. On the doctrine of chances a greater number
of the offspring of the more productive pairs would sur-
vive ; and these would inherit a tendency to procreate
fewer males and more females. Thus a tendency to-
wards the equalisation of the sexes would be brought
about. But our supposed species would by this process
be rendered, as just remarked, more productive ; and
this would in many cases be far from an advantage ;
for whenever the limit to the numbers which exist, de-
pends, not on destruction by enemies, but on the amount
of food, increased fertility will lead to severer competi-
tion and to most of the survivors being badly fed. In
this case, if the sexes were equalised by an increase in
the number of the females, a simultaneous decrease in
the total number of the offspring would be beneficial ;
and this, I believe, could be effected through natural

Chap. VIII. SEXUAL SELECTION. 317

selection in the manner hereafter to be described. The
same train of reasoning is applicable in the above,
as well as in the following case, if we assume that
females instead of males are produced in excess, for
such females from not uniting with males would be
superfluous and useless. So it would be with poly-
gamous species, if we assume the excess of females
to be inordinately great.

An excess of either sex, we will again say of the
males, could, however, apparently be eliminated through
natural selection in another and indirect manner, namely
by an actual diminution of the males, without any in-
crease of the females, and consequently without any
increase in the productiveness of the species. From
the variability of all characters, we may feel assured
that some pairs, inhabiting any locality, would produce
a rather smaller excess of superfluous males, but an
equal number of productive females. When the off-
spring from the more and the less male-productive
parents were all mingled together, none would have any
direct advantage over the others; but those that pro-
duced few superfluous males would have one great
indirect advantage, namely that their ova or embryos
would probably be larger and finer, or their young
better nurtured in the womb and afterwards. We see
this principle illustrated with plants; as those which
bear a vast number of seed produce small ones ; whilst
those which bear comparatively few seeds, often produce
large ones well-stocked with nutriment for the use of the
seedlings.73 Hence the offspring of the parents which
had wasted least force in producing superfluous males

'3 I have often been struck with the fact, that in several species of
Primula the seeds in the capsules which contained only a few were
very much larger than the numerous seeds in the more productive
capsules.

318 THE PKINCIPLES OF Part II.

would be the most likely to survive, and would inherit
the same tendency not to produce superfluous males,
whilst retaining their full fertility in the production of
females. So it would be with the converse case of the
female sex. Any slight excess, however, of either sex
could hardly be checked in so indirect a manner. Nor
indeed has a considerable inequality between the sexes
been always prevented, as we have seen in some of the
cases given in the previous discussion. In these cases
the unknown causes which determine the sex of the
embryo, and which under certain conditions lead to
the production of one sex in excess over the other, have
not been mastered by the survival of those varieties
which were subjected to the least waste of organised
matter and force by the production of superfluous indi-
viduals of either sex. Nevertheless we may conclude
that natural selection will always tend, though some-
times inefficiently, to equalise the relative numbers of
the two sexes.

Having said this much on the equalisation of the
sexes, it may be well to add a few remarks on the regula-
tion through natural selection of the ordinary fertility
of species. Mr. Herbert Spencer has shewn in an able
discussion 74 that with all organisms a ratio exists be-
tween what he calls individuation and genesis; whence
it follows that beings which consume much matter or
force in their growth, complicated structure or activity,
or which produce ova and embryos of large size, or
which expend much energy in nurturing their young,
cannot be so productive as beings of an opposite nature.
Mr. Spencer further shews that minor differences in fer-
tility will be regulated through natural selection. Thus
the fertility of each species will tend to increase, from

r-i <

Principles of Biology,' vol. ii. 18G7, chaps, ii.-xi.

Chap. VIII. SEXUAL SELECTION. 319

the more fertile pairs producing a larger number of off-
spring, and these from their mere number will have the
best chance of surviving, and will transmit their ten-
dency to greater fertility. The only check to a con-
tinued augmentation of fertility in each organism seems
to be either the expenditure of more power and the
greater risks run by the parents that produce a more
numerous progeny, or the contingency of very numerous
eggs and young being produced of smaller size, or less
vigorous, or subsequently not so well nurtured. To
strike a balance in any case between the disadvantages
which follow from the production of a numerous pro-
geny, and the advantages (such as the escape of at least
some individuals from various dangers) is quite beyond
our power of judgment.

When an organism has once been rendered extremely
fertile, how its fertility can be reduced through natural
selection is not so clear as how this capacity was first
acquired. Yet it is obvious that if individuals of a
species, from a decrease of their natural enemies, were
habitually reared in larger numbers than could be sup-
ported, all the members would suffer. Nevertheless the
offspring from the less fertile parents would have no
direct advantage over the offspring from the more fer-
tile parents, when all were mingled together in the
same district. All the individuals would mutually tend
to starve each other. The offspring indeed of the less
fertile parents would lie under one great disadvantage,
for from the simple fact of being produced in smaller
numbers, they would be the most liable to extermina-
tion. Indirectly, however, they wOuld partake of one
great advantage ; for under the supposed condition of
severe competition, when all were pressed for foofl, it is
extremely probable that those individuals which from
some variation in their constitution produced fewer eggs

320 THE PRINCIPLES OF SEXUAL SELECTION. Part II.

or young, would produce them of greater size or vigour ;
and the adults reared from such eggs or young would
manifestly have the best chance of surviving:, and
would inherit a tendency towards lessened fertility.
The parents, moreover, which had to nourish or provide
for fewer offspring would themselves be exposed to a
less severe strain in the struggle for existence, and
would have a better chance of surviving. By these
steps, and by no others as far as I can see, natural
selection under the above conditions of severe com-
petition for food, would lead to the formation of a new
race less fertile, but better adapted for survival, than
the parent-race.

Chap. IX. SEXUAL SELECTION. 321

CHAPTER IX.

Secondary Sexual Characters in the Lower Classes of

the Animal Kingdom.

These characters 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.

In the lowest classes the two sexes are not rarely united
in the same individual, and therefore secondary sexual
characters cannot be developed. In many cases in which
the two sexes are separate, both are permanently at-
tached to some support, and the one cannot search or
strup-o-le for the other. Moreover it is almost certain
that these animals have too imperfect senses and
much too low mental powers to feel mutual rivalry,
or to appreciate each other's beauty or other attrac-
tions.

Hence in these classes or sub-kingdoms, such as the
Protozoa, Coelenterata, Echinodermata, Scolecida, true
secondarv sexual characters do not occur ; and this fact
agrees with the belief that such characters in the
higher classes have been acquired through sexual selec-
tion, which depends on the will, desires, and choice of
either sex. Nevertheless some few apparent exceptions
occur ; thus, as I hear from Dr. Baird, the males of
certain Entozoa, or internal parasitic worms, differ
slightly in colour from the females ; but we have no
reason to suppose that such differences have been
augmented through sexual selection.

VOL. I. Y

322 SEXUAL SELECTION. Part II.

Many of the lower animals, whether hermaphrodites
or with the sexes separate, are ornamented with the
most brilliant tints, or are shaded and striped in an
elegant manner. This is the case with many corals
and sea-anemonies (Actiniae), with some jelly-fish (Me-
dusae, Porpita, &c), with some Planariae, Ascidians,
numerous Star-fishes, Echini, &e. ; but we may conclude
from the reasons already indicated, namely the union
of the two sexes in some of these animals, the per-
manently affixed condition of others, and the low
mental powers of all, that such colours do not serve
as a sexual attraction, and have not been acquired
through sexual selection. With the higher animals
the case is very different ; for with them when one sex
is much more brilliantly or conspicuously coloured
than the other, and there is no difference in the
habits of the two sexes which will account for this
difference, we have reason to believe in the influence
of sexual selection ; and this belief is strongly con-
firmed when the more ornamented individuals, which
are almost always the males, display their attractions
before the other sex. We may also extend this con-
clusion to both sexes, when coloured alike, if their
colours are plainly analogous to those of one sex alone
in certain other species, of the same group.

How, then, are we to account for the beautiful or
even gorgeous colours of many animals in the lowest
classes ? It appears very doubtful whether such colours
usually serve as a protection ; but we are extremely
liable to err in regard to characters of all kinds in
relation to protection, as will be admitted by every one
who has read Mr. Wallace's excellent essay on this
subject. It would not, for instance, at first occur to
any one that the perfect transparency of the Medusae,
or jelly-fishes, was of the highest service to them as a

Chap. IX. SEXUAL SELECTION. 323

protection ; but when we are reminded by Hackel that
not only the medusas but many floating mollusca, crus-
taceans, and even small oceanic fishes partake of this
same glass-like structure, we can hardly doubt that
they thus escape the notice of pelagic birds and other
enemies.

Notwithstanding our ignorance how far colour in
many cases serves as a protection, the most probable
view in regard to the splendid tints of many of the
lowest animals seems to be that their colours are the
direct result either of the chemical nature or the minute
structure of their tissues, independently of any benefit
thus derived. Hardly any colour is finer than that of
arterial blood ; but there is no reason to suppose that
the colour of the blood is in itself any advantage ; and
though it adds to the beauty of the maiden's cheek, no
one will pretend that it has been acquired for this pur-
pose. So again with many animals, especially the lower
ones, the bile is richly coloured ; thus the extreme
beauty of the Eolidae (naked sea-slugs) is chiefly due, as
I am informed by Mr. Hancock, to the biliary glands
seen through the translucent integuments ; this beauty
being probably of no service to these animals. The
tints of the decaying leaves in an American forest are
described by every one as gorgeous ; yet no one sup-
poses that these tints are of the least advantage to
the trees. Bearing in mind how many substances
closely analogous to natural organic compounds have
been recently formed by chemists, and which exhibit
the most splendid colours, it would have been a strange
fact if substances similarly coloured had not often
originated, independently of any useful end being
thus gained, in the complex laboratory of living
organisms.

y 2

324 SEXUAL SELECTION. Part II.

The sub-kingdom of the Mollusca. — Throughout this
great division (taken in its largest acceptation) of the
animal kingdom, secondary sexual characters, such as
we are here considering, never, as far as I can discover,
occur. Nor could they be expected in the three lowest
classes, namely in the Ascidians, Polyzoa, and Brachio-
pods (constituting the Molluscoida of Huxley), for most
of these animals are permanently affixed to a support
or have their sexes united in the same individual. In
the Lamellibranchiata, or bivalve shells, hermaphro-
ditism is not rare. In the next higher class of the
Gasteropoda, or marine univalve shells, the sexes are
either united or separate. But in this latter case the
males never possess special organs for finding, securing,
or charming: the females, or for fio-htino- with other
males. The sole external difference between the sexes
consists, as I am informed by Mr. Gwyn Jeffreys, in
the shell sometimes differing a little in form ; for
instance, the shell of the male periwinkle (Littorina
littorea) is narrower and has a more elongated spire than
that of the female. But differences of this nature, it
may be presumed, are directly connected with the act
of reproduction or with the development of the ova.

The Gasteropoda, though 'capable of locomotion and
furnished with imperfect eyes, do not appear to be en-
dowed with sufficient mental powers for the members
of the same sex to struggle together in rivalry, and
thus to acquire secondary sexual characters. Never-
theless with the pulmoniferous gasteropods, or land-
snails, the pairing is preceded by courtship ; for these
animals, though hermaphrodites, are compelled by their
structure to pair together. Agassiz remarks,1 " Qui-
" conque a eu l'occasion d'observer les amours cles lima-

1 ' De l'Espece et cle la Class.' &c, 1869, p. 106.

Chap. IX. MOLLUSCS. 325

" cons, ne saurait mettre en doute la seduction deployee
"dans les mouvements et les allures qui preparent et
" accomplissent le double embrassement de ces her-*
" maphrodites." These animals appear also susceptible
of some degree of permanent attachment : an accurate
observer, Mr. Lonsdale, informs me that he placed a
pair of land-shells (Helix pomatia), one of which was
weakly, into a small and ill-provided garden. After a
short time the strong and healthy individual disappeared,
and was traced by its track of slime over a wall into an
adjoining well-stocked garden. Mr. Lonsdale concluded
that it had deserted its sickly mate ; but after an
absence of twenty-four hours it returned, and apparently
communicated the result of its successful exploration,
for both then started along the same track and disap-
peared over the wall.

Even in the highest class of the Mollusca, namely the
Cephalopoda or cuttle-fishes, in which the sexes are
separate, secondary sexual characters of the kind which
we are here considering, do not, as far as I can discover,
occur. This is a surprising circumstance, as these
animals possess highly-developed sense-organs and have
considerable mental powers, as will be admitted by
everv one who has watched their artful endeavours to
escape from an enemy.2 Certain Cephalopoda, however,
are characterised by one extraordinary sexual character,
namely, that the male element collects within one of
the arms or tentacles, which is then cast off, and,
clinging by its sucking-discs to the female, lives for
a time an independent life. So completely does the
cast-off arm resemble a separate animal, that it was
described by Cuvieras a parasitic worm under the name

2 See, for instance, the account which I have given in my ' Journal
of Kesenrches,' 1845. p. 7.

326 SEXUAL SELECTION. Part II.

of Hectocotyle. But this marvellous structure may be
classed as a primary rather than as a secondary sexual
character.

Although with the Mollusca sexual selection does not
seem to have come into play ; yet many univalve and
bivalve shells, such as volutes, cones, scallops, &c, are
beautifully coloured and shaped. The colours do not
appear in most cases to be of any use as a protection ;
they are probably the direct result, as in the lowest
classes, of the nature of the tissues ; the patterns and
the sculpture of the shell depending on its manner of
growth. The amount of light seems to a certain extent
to be influential ; for although, as repeatedly stated by
Mr. G-wyn Jeffreys, the shells of some species living at a
profound depth are brightly coloured, yet we generally
see the lower surfaces and the parts covered by the
mantle less highly coloured than the upper and exposed
surfaces.3 In some cases, as with shells livino- ainono-st
corals or brightly-tinted sea-weeds, the bright colours
may serve as a protection. But many of the nudibranch
mollusca, or sea-slugs, are as beautifully coloured as
any shells, as may be seen in Messrs. Alder and Han-
cock's magnificent work ; and from information kindly
given me by Mr. Hancock, it is extremely doubtful
whether these colours usually serve as a protection.
With some species this may be the case, as with one
which lives on the green leaves of algae, and is itself
bright-green. But many brightly-coloured, white or
otherwise conspicuous species, do not seek concealment;
whilst again some equally conspicuous species, as well
as other dull-coloured kinds, live under stones and in

3 I have given ('Geolog. Observations on Volcanic Islands,' 1844,
p. 53) a curious instance of the influence of light on the colours of
a frondescent incrustation, deposited by the surf on the coast-rocks of
Ascension, and formed by the solution of triturated sea-shells.

Chap. IX. MOLLUSCS AND ANNELIDS. 327

dark recesses. So that with these nudibranch molluscs,
colour apparently does not stand in any close relation
to the nature of the places which they inhabit.

These naked sea-slugs are hermaphrodites, yet they
pair together, as do land-snails, many of which have
extremely pretty shells. It is conceivable that two
hermaphrodites, attracted by each others' greater beauty,
might unite and leave offspring: which would inherit
their parents' greater beauty. But with such lowly-
organised creatures this is extremely improbable. Nor
is it at all obvious how the offspring from the more
beautiful pairs of hermaphrodites would have any ad-
vantage, so as to increase in numbers, over the offspring
of the less beautiful, unless indeed vigour and beauty
generally coincided. We have not here a number of
males becoming mature before the females, and the
more beautiful ones selected by the more vigorous
females. If, indeed, brilliant colours were beneficial
to an hermaphrodite animal in relation to its general
habits of life, the more brightly-tinted individuals would
succeed best and would increase in number ; but this
would be a case of natural and not of sexual selection.

Sub-kingdom of the Vermes : Class, Annelida {or Sea-
worms). — In this class, although the sexes (when separate)
sometimes differ from each other in characters of such
importance that they have been placed under distinct
genera or even families, yet the differences do not
seem of the kind which can be safely attributed to
sexual selection. These animals, like those in the pre-
ceding classes, apparently stand .too low in the scale,
for the individuals of either sex to exert any choice in
selecting a partner, or for the individuals of the same
sex to struggle together in rivalry.

328 SEXUAL SELECTIOX. Part II.

Sub-kingdom of the Arthropoda: Class, Crustacea. —
In this great class we first meet with undoubted se-
condary sexual characters, often developed in a remark-
able manner. Unfortunately the habits of crustaceans
are very imperfectly known, and we cannot explain the
uses of many structures peculiar to one sex. With
the lower parasitic species the males are of small size,
and they alone are furnished with perfect swimming-
legs, antennae and sense-organs ; the females being
destitute of these organs, with their bodies often consist-
ing of a mere distorted mass. But these extraordinary
differences between the two sexes are no doubt related
to their widely different habits of life, and consequently
clo not concern us. In various crustaceans, belonging
to distinct families, the anterior antennas are furnished
with peculiar thread-like bodies, which are believed to
act as smelling-organs, and these are much more nume-
rous in the males than in the females. As the males,
without any unusual development of their olfactory
organs, would almost certainly be able sooner or later
to find the females, the increased number of the smell-
ing-threads has probably been acquired through sexual
selection, by the better provided males having been the
most successful in finding partners and in leaving off-
spring. Fritz Miiller has described a remarkable dimor-
phic species of Tanais, in which the male is represented
by two distinct forms, never graduating into each other.
In the one form the male is furnished with more
numerous smel ling-threads, and in the other form with
more powerful and more elongated chelae or pincers
which serve to hold the female. Fritz Miiller suo-gests
that these differences between the two male forms of the
same species must have originated in certain individuals
having varied in the number of the smelling-threads,
whilst other individuals varied in the shape and size of

Chap. IX.

CRUSTACEANS.

329

their chelae ; so that of the former, those which were best
able to find the female, and of the latter, those which were
best able to hold her when found,
have left the greater number of
progeny to inherit their respec-
tive advantages.4

In some of the lower crusta-
ceans, the right-hand anterior
antenna of the male differs
greatly in structure from the
left-hand one, the latter re-
sembling in its simple tapering
joints the antennae of the fe-
male. In the male the modi-
fied antenna is either swollen
in the middle or angularly bent,
or converted (fig. 3) into an
elegant, and sometimes wonder-
fully complex, prehensile organ.5
It serves, as I hear from Sir J.
Lubbock, to hold the female,
and for this same purpose one
of the two posterior legs (b) on
the same side of the body is
converted into a forceps. In
another family the inferior or
posterior antennae are " curiously zigzagged ': in the
males alone.

Fig. 3.

Labidocera Darwinii (from
Lubbock).

Part of right-hand anterior an-
tenna of male, forming a pre-
hensile organ.

Posterior pair of thoracic legs of
male.

Ditto of female.

4 ' Facts and Arguments for Darwin,' English translat. 1869, p. 20.
See the previous discussion on the olfactory threads. Sars has de-
scribed a somewhat analogous case (as quoted in 'Nature,' 1870,
p. 455) in a Norwegian crustacean, the Pontoporeia affinis.

5 See Sir J. Lubbock in 'Annals, and Mag. of Nat. Hist.' vol. si.
1853, pi. i. and x. ; and vol. xii. (185H) pi. vii. See also Lubbock in
' Transact. Ent. Soc' vol. iv. new series, 1856-1858, p. 8. With respect
to the zigzagged antenna? mentioned below, see Fritz Miiller, ' Facts
and Arguments for Darwin ' 1869, p. 40, foot-note.

330

SEXUAL SELECTION.

Part II.

In the higher crustaceans the anterior legs form a
pair of chelae or pincers, and these are generally
larger in the male than in the female. In many species
the chelae on the opposite sides of the body are of
unequal size, tbe right-hand one being, as I am in-

Fig. 4. Anterior part of body of Callianassa (from Milne-Edwards), showing the unequal
and differently -constructed right and left-hand chela? of the male.

N.B.— The artist by mistake has reversed the drawing, and made the left-hand chela
the largest.

Fig. 5.

Fig. 6.

Fig. 5. Second leg of male Orchestia Tucuratinga (from Fritz Miiller).
Fig. 6. Ditto of female.

formed by Mr. C. Spence Bate, generally, though not
invariably, the largest. This inequality is often much
greater in the male than in the female. The two chelae
also often differ in structure (figs. 4 and 5, 6), the smaller
one resembling that of the female. What advantage

Chap. IX. CRUSTACEANS. 331

is gained by their inequality in size on the opposite
sides of the body, and by the inequality being much
greater in the male than in the female ; and why, when
they are of equal size, both are often much larger in
the male than in the female, is not known. The chelae
are sometimes of such length and size that they cannot
possibly be used, as I hear from Mr. Spence Bate, for
carrying food to the mouth. In the males of certain fresh-
water prawns (Palsemon) the right leg is actually longer
than the whole body.6 It is probable that the great size
of one leg with its chelae may aid the male in fighting
with his rivals ; but this use will not account for their
inequality in the female on the opposite sides of the body.
In Gelasimus, according to a statement quoted by Milne-
Edwards,7 the male and female live in the same burrow,
which is worth notice, as shewing that they pair, and
the male closes the mouth of the burrow with one of its
chelae, which is enormously developed ; so that here it
indirectly serves as a means of defence. Their main
use, however, probably is to seize and to secure the
female, and this in some instances, as with Gammarus,
is known to be the case. The sexes, however, of the
common shore-crab (Carcinus meenas), as Mr. Spence
Bate informs me, unite directly after the female has
moulted her hard shell, and when she is so soft that she
would be injured if seized by the strong pincers of the
male ; but as she is caught and carried about by the
male previously to the act of moulting, she could then
be seized with impunity.

Fritz Miiller states that certain species of Melita are

6 See a paper by Mr. C. Spence Bate, with figures, iu ' Proc. Zoolog.
Soc.' ]868, p. 3(J3 ; and on the nomenclature of the genus, ibid. p. 585.
I am greatly indebted to Mr. Spence Bate for nearly all the above
statements with respect to the chelae of the higher crustaceans.

7 ' Hist. Nat. des Crust.' torn. ii. 1837, p. 50.

332 SEXUAL SELECTION. Part II.

distinguished from all other amphipods by the females
having "the coxal lamellae of the penultimate pair of
" feet produced into hook-like processes, of which the
" males lay hold with the hands of the first pair." The
development of these hook-like processes probably
resulted from those females which were the most
securely held during the act of reproduction, having
left the largest number of offspring. Another Bra-
zilian amphipod (Orchestia Darwinii, fig. 7) is de-
scribed by Fritz Miiller, as presenting a case of dimor-
phism, like that of Tanais ; for there are two male
forms, which differ in the structure of their chelae.8 As
chelae of either shape would certainly have sufficed to
hold the female, for both are now used for this purpose,
the two male forms probably originated, by some having
varied in one manner and some in another ; both forms
having derived certain special, but nearly equal advan-
tages, from their differently shaped organs.

It is not known that male crustaceans fisrht together
for the possession of the females, but this is probable ;
for with most animals when the male is larger than the
female, he seems to have acquired his greater size by
having conquered during many generations other males.
Now Mr. 8pence Bate informs me that in most of the
crustacean orders, especially in the highest or the
Brachyura, the male is larger than the female; the
parasitic genera, however, in which the sexes follow
different habits of life, and most of the Entomostraca
must be excepted. The chelae of many crustaceans are
weapons well adapted for fighting. Thus a Devil-crab
(Portunus puher) was seen by a son of Mr. Bate fighting
with a Carcinus mmnas, and the latter was soon thrown
on its back, and had every limb torn from its body.

Fritz Miiller, ' Facts and Arguments for Darwin,' 1869, p. 25-28.

Chap. IX.

CRUSTACEANS.

333

When several males of a Brazilian Gelasimus, a species
furnished with immense pincers, were placed together
by Fritz Miiller in a glass vessel, they mutilated and
killed each other. Mr. Bate put a large male Carcinus

;j'!r^r^rr

Kig. 7. Oichestia Darwinii (from Fritz IVIuller), showing the different ly-cm istriu ted

chehe of the two male forms.

mamas into a pan of water, inhabited by a female paired
with a smaller male ; the Litter was soon dispossessed,
but, as Mr. Bate adds, " if they fought, the victor)'

334 SEXUAL SELECTION. Part II.

" was a bloodless one, for I saw no wounds." This
same naturalist separated a male sand-skipper (so com-
mon on our sea- shores), Gammarus marinus, from its
female, both of which were imprisoned in the same
vessel with many individuals of the same species. The
female being thus divorced joined her comrades. After
an interval the male was again put into the same
vessel and he then, after swimming about for a time,
dashed into the crowd, and without anv fight-ins: at once
took away his wife. This fact shews that in the Amphi-
poda, an order low in the scale, the males and females
recognise each other, and are mutually attached.

The mental powers of the Crustacea are probably
higher than might have been expected. Any one who
has tried to catch one of the shore-crabs, so numerous
on many tropical coasts, will have perceived how wary
and alert they are. There is a large crab (Birgus
latro), found on coral islands, which makes at the
bottom of a deep burrow a thick bed of the picked
fibres of the cocoa-nut. It feeds on the fallen fruit of
this tree by tearing off the husk, fibre by fibre ; and
it always begins at that end where the three eye-
like depressions are situated. It then breaks through
one of these eyes by hammering with its heavy front
pincers, and turning round, extracts the albuminous
core with its narrow posterior pincers. But these actions
are probably instinctive, so that they would be per-
formed as well by a young as by an old animal.
The following case, however, can hardly be so con-
sidered : a trustworthy naturalist, Mr. Gardner,9 whilst
watching a shore-crab (Gelasimus) making its burrow,

9 'Travels in the Interior of Brazil,' 1846, p. 111. I have given, in
my ' Journal of Researches,' p. 463, an account of the habits of the

Birgos.

Chap. IX. CRUSTACEANS. 335

threw some shells towards the hole. One rolled in,
and three other shells remained within a few inches of
the mouth. In about live minutes the crab brought
out the shell which had fallen in, and carried it away
to the distance of a foot ; it then saw the three other
shells lying near, and evidently thinking that they
might likewise roll in, carried them to the spot where
it had laid the first. It would, I think, be difficult to
distinguish this act from one performed by man by the
aid of reason.

With respect to colour which so often differs in the
two sexes of animals belonging to the higher classes,
Mr. Spence Bate does not know of any well-marked
instances with our British crustaceans. In some cases,
however, the male and female differ slightly in tint,
but Mr. Bate thinks not more than may be accounted
for by their different habits of life, such as by the
male wandering more about and being thus more ex-
posed to the light. In a curious Bornean crab, which
inhabits sponges, Mr. Bate could always distinguish the
sexes by the male not having the epidermis so much
rubbed off. Dr. Power tried to distinguish by colour
the sexes of the species which inhabit the Mauritius, but
always failed, except with one species of Squilla, pro-
bably the S. stijlifera, the male of which is described as
being " of a beautiful blueish-green," with some of the
appendages cherry-red, whilst the female is clouded
with brown and grey, " with the red about her much
" less vivid than in the male." 10 In this case, we may
suspect the agency of sexual selection. With Sa-
phirina (an oceanic genus of Entomostraca, and there-
fore low in the scale) the males are furnished with

10 Mr. Oh. Fraser, in 'Proc. Zoolog. Soc.' 1869, p. 3. I am indebted
to Mr. Bate for the statement from Dr. Power.

336 SEXUAL SELECTION. Part II.

minute shields or cell-like bodies, which exhibit beau-
tiful changing colours ; these being absent in the
females, and in the case of one species in both sexes.11
It would, however, be extremely rash to conclude that
these curious organs serve merelv to attract the females.

© Ml

In the female of a Brazilian species of Gelasimus, the
whole body, as I am informed by Fritz Muller, is of a
nearly uniform greyish-brown. In the male the posterior
part of the cephalo-thorax is pure white, with the
anterior part of a rich green, shading into dark brown ;
and it is remarkable that these colours are liable to
change in the course of a few minutes — the white
becoming dirty grey or even black, the green " losing
much of its brilliancy." The males apparently are
much more numerous than the females. It deserves
especial notice that they do not acquire their bright
colours until they become mature. They differ also
from the females in the larger size of their chela?.
In some species of the genus, probably in all, the
sexes pair and. inhabit the same burrow. They are
also, as we have seen, highly intelligent animals.
From these various considerations it seems highly
probable that the male in this species has become
srailv ornamented in order to attract or excite the
female.

It has just been stated, that the male Gelasimus does
not acquire his conspicuous colours until mature and
nearly ready to breed. This seems the general rule in
the whole class with the many remarkable difference^
in structure between the two sexes. We shall here-
after find the same law prevailing throughout the great
sub-kingdom of the Yertebrata, and in all cases it is
eminently distinctive of characters which have been

11 Clans, ' Die freilebendeu Copepoden/ 1863, s. 35.

Chap. IX. SPIDERS. 337

acquired through sexual selection. Fritz Miiller12 gives
some striking instances of this law ; thus the male
sand-hopper (Orchestia) does not acquire his large
claspers, which are very differently constructed from
those of the female, until nearly full-grown ; whilst
young his claspers resemble those of the female. Thus,
again, the male Brachyscelus possesses, like all other
amphipods, a pair of posterior antennae ; the female,
and this is a most extraordinary circumstance, is desti-
tute of them, and so is the male as long as he remains
immature.

Class, Araclmida (Spiders). — The males are often
darker, but sometimes lighter than the females, as may
be seen in Mr. Blackwall's magnificent work.13 In
some species the sexes differ conspicuously from each
other in colour ; thus the female of Sjparassus sma-
ragdulus is dullish-green ; whilst the adult male has
the abdomen of a fine yellow, with three longitudinal
stripes of rich red. In some species of Thomisus the
two sexes closely resemble each other ; in others they
differ much; thus in T. citreus the legs and body of
the female are pale-yellow or green, whilst the front
legs of the male are reddish-brown : in T. Jiorieolens,
the legs of the female are pale-green, those of the
male being ringed in a conspicuous manner with various
tints. Numerous analogous cases could be given in the
genera Epeira/Nephila, Philodromus, Theridion, Liny-
phia, &c. It is often difficult to say which of the two
sexes departs most from the ordinary coloration of the
genus to which the species belong ; "but Mr. Blackwall

f

12 ' Facts and Arguments,' &c., p. 79.

13 'A History of the Spiders of Great Britain,' 18<»1-G4. For thc

following facts, see p. 102, 77, 8S.
VOL. I.

338 SEXUAL SELECTION. Part II.

thinks that, as a general rule, it is the male. Both
sexes whilst young, as I am informed by the same
author, usually resemble each other; and both often
undergo great changes in colour during their successive
moults before arriving at maturity. In other cases
the male alone appears to change colour. Thus the
male of the above-mentioned brightly-coloured Spa-
rassus at first resembles the female and acquires his
peculiar tints only when nearly adult. Spiders are
possessed of acute senses, and exhibit much intelli-
gence. The females often shew, as is well known, the
strongest affection for their eggs, which they carry
about enveloped in a silken web. On the whole it
appears probable that well-marked differences in colour
between the sexes have generally resulted from sexual
selection, either on the male or female side. But doubts
may be entertained on this head from the extreme
variability in colour of some species, for instance of
Theridion lineatum, the sexes of which differ when
adult ; this great variability indicates that their colours
have not been subjected to any form of selection.

Mr. Black wall does not remember to have seen the
males of any species fighting together for the posses-
sion of the female. Nor, judging from analogy, is this
probable; for the males are generally much smaller
than the females, sometimes to an extraordinary de-
gree.14 Had the males been in the habit of fighting
together, they would, it is probable, have gradually

14 Aug. Vinson ('Arane'ides des lies de la Ke'union,' pi. vi. figs. 1
and 2) gives a good instance of the small size of the male in Epeira
nigra. In this species, as I mny add, the male is testaceous and the
female black with legs banded with red. Other even more striking
cases of inequality in size between the sexes have been recorded
C Quarterly Journal of Science,' 1868, July, p. 429); but I have not seen
the original accounts.

C;iai\ IX. SPIDERS AND MYEIAPODA. 339

acquired greater size and strength. Mr. Blackwall Las
sometimes seen two or more males on the same web
with a single female ; but their courtship is too tedious
and prolonged an affair to be easily observed. The male
is extremely cautious in making his advances, as the
female carries her coyness to a dangerous pitch. De
Geer saw a male that " in the midst of his preparatory
" caresses was seized by the object of his attentions,
" enveloped by her in a web and then devoured, a
" sight which, as he adds, filled him with horror and
" indignation." 15

Westrino* has made the interesting discovery that
the males of several species of Theridion16 have the
power of making a striclulating sound (like that made
by many beetles and other insects, but feebler), whilst
the females are quite mute. The apparatus consists of
a serrated ridge at the base of the abdomen, against
which the hard hinder part of the thorax is rubbed ;
and of this structure not a trace could be detected in
the females. From the analogy of the Orthoptera and
Homoptera, to be described in the next chapter, we
may feel almost sure that the stridulation serves, as
Westring remarks, either to call or to excite the
female ; and this is the first case in the ascending scale
of the animal kingdom, known to me, of sounds emitted
for this purpose.

Class, Myriapoda. — In neither of the two orders in
this class, including the millipedes and centipedes,

15 Kirby and Spence, ' Introduction to Entomology,' vol. i. ISIS,
p. 280.

16 Tberidion (Asagena, Sund.) serratipes, 4-punctatum et guttatum ;
see Westrir.g, in Kroyer, ' Naturhist. Tidskrift,' vol. iv. 1842-1843,
p. 319; and vol. ii. 1846-1849, p. 342. See, also, for other species,
' Aruneui Svecicso,' p. 184.

z 2

340 SEXUAL SELECTION. Part II.

can I find any well-marked instances of sexual dif-
ferences such as more particularly concern us. In
Glomeris limbata, however, and perhaps in some few
other species, the males differ slightly in colour from
the females; but this Glomeris is a highly variable
species. In the males of the Diplopoda, the legs be-
longing to one of the anterior segments of the body, or
to the posterior segment, are modified into prehensile
hooks which serve to secure the female. In some
species of lulus the tarsi of the male are furnished
with membranous suckers for the same purpose. It is
a much more unusual circumstance, as we shall see
when w^e treat of Insects, that it is the female in
Lithobius which is furnished with prehensile appen-
dages at the extremity of the body for holding the
male. 17

17 Walckenaer et P. Gervais. 'Hist. Nat. cles Insectes : Apteres,'
torn. iv. 1847, p. 17, 19, 68.

Chap. X. SEXUAL SELECTION. 341

CHAPTER X.

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 Dowers 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 colours — Coleoptera, colours ; furnished with great horns,
apparently as an ornament ; battles ; stridulating organs generally
common to both sexes.

In the immense class of insects the sexes sometimes
differ in their organs for locomotion, and often in
their sense-organs, as in the pectinated and beauti-
fully plumose antennae of the males of many species.
In one of the Ephemerae, namely Chloeon, the male
has great pillared eyes, of which the female is entirely
destitute.1 The ocelli are absent in the females of
certain other insects, as in the Mutillida?, which are
likewise destitute of wings. But we are chiefly con-
cerned with structures by which one male is enabled to
conquer another, either in battle or courtship, through
his strength, pugnacity, ornaments, or music. The
innumerable contrivances, therefore, by which the male
is able to seize the female, may be briefly passed over.
Besides the complex structures at the apex of the abdo-
men, which ought perhaps to be ranked as primary

1 Sir J. Lubbock, 'Transact. Linnean Soc.' vol. xxv. 1866. p. 484.
With respect to the Mutillidje see Westwood, ' Modern Class, of Insects^'
vol. ii. p. 213.

342 SEXUAL SELECTION. Part II.

organs,2 " it is astonishing," as Mr. B. D. Walsh3 has
remarked, " how many different organs are worked in
" by nature, for the seemingly insignificant object of
" enabling the male to grasp the female firmly." The
mandibles or jaws are sometimes used for this purpose ;
thus the male Corydalis cor nidus (a neuropterous insect
in some degree allied to the Dragon-flies, &c.) has im-
mense curved jaws, many times longer than those of the
female ; and. they are smooth instead of being toothed,
by which means he is enabled to seize her without
injury.4 One of the stag-beetles of North America
(Lucanus elaphus) uses his jaws, which are much larger
than those of the female, for the same purpose, but
probably likewise for fighting. In one of the sand-wasps
(AmmojiMla) the jaws in the two sexes are closely
alike, but are used for widely different purposes ; the
males, as Professor Westwood observes, " are exceed-
" ingly ardent, seizing their partners round the neck
" with their sickle-shaped jaws ; " 5 whilst the females use

2 These organs in the male often differ in closely-allied species, and
afford excellent specific characters. But their importance, under a
functional point of view, as Mr. K. MacLachlan has remarked to me,
has probably been overrated. It has been suggested, that slight dif-
ferences in these organs would suffice to prevent the intercrossing of
well-marked varieties or incipient species, and would thus aid in their
development. That this can hardly be the case, we may infer from the
many recorded cases (see for instance, Bronn, ' Gesehichte der Natur/
B. ii. 1843, s. 164; and Westwood, 'Transact. Ent. Soc' vol. iii. 1842,
p. 195) of distinct species having been observed in union. Mr.
MacLachlan informs me vide 'Stett. Ent. Zeitung,' 1867. s. 155) that
when several species of Phryganidse, which present strongly-pronounced
differences of this kind, were confined together by Dr. Aug. Meyer,
they coupled, and one pair produced fertile ova.

s ' The Practical Entomologist,' Philadelphia, vol. ii. May,^ 1867,
p. 88.

4 Mr. Walsh, ibid. p. 107.

5 < Modern Classification of Insects/ vol. ii. 1840, p. 206, 205. Mr.
Walsh, who called my attention to this double use of the jaws, says
that he has repeatedly observed this fact.

Chap. X.

INSECTS.

o i o

these organs for burrowing in sand-banks and making
their nests.

The tarsi of the front-legs are dilated in many male
beetles, or are furnished with broad cushions of hairs ;
and in many genera of water-beetles they are armed
with a round flat sucker, so that the male may adhere
to the slippery body of the female. It is a much more
unusual circumstance that the females of some water-
beetles (Dytiscus) have their
elytra deeply grooved, and
in Acilius sulcatus thickly set
with hairs, as an aid to the
male. The females of some
other water-beetles (Hydro-
porus) have their elytra
punctured for the same ob-
ject.6 In the male of Crabro
cribrarius (fig. 8.), it is the
tibia which is dilated into a
broad horny plate, with mi-
nute membraneous dots, giv-
ing to it a singular appear-
ance like that of a riddle.7
In the male of Pen the (a

genUS Of beetles) a few Of FiS- 8. Crabro cribrarius. Upper figure
,-. . , -,-, . . ' r.,1 male; lower figure, female.

the middle joints ot the an-

tennge are dilated and furnished on the inferior surface

6 We have here a curious ana inexplicable cdse of dimorphism, for
some of the females of four European species of Dytiscus, and of certain
species of Iiydroporus, have their elytra smooth ; and no intermediate
gradations between sulcated or punctured "and quite smooth elytra
have been observed. See Dr. H. Schaum, as quoted in the ' Zoologist,'
vol. v.-vi. 1847-48, p. 1896. Also Kirby and Spence, ' Introduction to
Entomology/ vol. hi. 1826, p. 305.

7 Westwood, 'Modern Class.' vol. ii. p. 193. The following state-
ment about Pentlie, and others in inverted commas, are taken from
Mr. Walsh, ' Practical Entomologist,' Philadelphia, vol. ii. p. 88.

3-14

SEXUAL SELECTION.

Part II.

with cushions of hair, exactly like those on the tarsi of
the Carabidse, " and obviously for the same end." In
male dragon-flies, "the appendages at the tip of the tail
" are modified in an almost infinite variety of curious

" patterns to enable them to embrace
<( the neck of the female." Lastly in
the males of many insects, the legs
are furnished with peculiar spines,
knobs or spurs ; or the whole leg is
bowed or thickened, but this is by
no means invariably a sexual cha-
racter ; or one pair, or all three
pairs are elongated, sometimes to
an extravagant length.8

In all the orders, the sexes of
many species present differences, of
which the meaning is not under-
stood. One curious case is that of
a beetle (fig. 9), the male of which
has the left mandible much en-
larged ; so that the mouth is greatly
distorted. In another Carabidous
beetle, the Eurygnathus,9 we have
the unique case, as far as known to
Mr. Wollaston, of the head of the
female beino; much broader and
larger, though in a variable degree,
than that of the male. Any number

Fig. 9. Taphrodei-es distortus f j cageg 1(J be „iven< TlieV

(much enlarged). Upper > n *

figure, male; lower figure, abound in the Lepidoptera : one

of the most extraordinary is that
certain male butterflies have their fore-legs more or

s Kirby and Spence, ' Introduct.' &c, vol. iii. p. 332-336.
9 'Insecta Maclerensia,' 1854, p. 20.

Chap. X. INSECTS. 345

less atrophied, with the tibiee and tarsi reduced to mere
rudimentary knobs. The wings, also, in the two sexes
often differ in neuration,10 and sometimes considerably
in outline, as in the Aricoris epitus, which was shown
to me in the British Museum by Mr. A. Butler. The
males of certain South American butterflies have tufts
of hair on the margins of the wings, and. horny excres-
cences on the discs of the posterior pair.11 In several
British butterflies, the males alone, as shewn by Mr.
Wonfor. are in parts clothed with peculiar scales.

The purpose of the luminosity in the female glow-
worm is likewise not understood ; for it is very doubtful
whether the primary use of the light is to guide the
male to the female. It is no serious objection to this
latter belief that the males emit a feeble light; for
secondary sexual characters proper to one sex are often
developed in a slight degree in the other sex. It is a
more valid objection that the larvse shine, and in some
species brilliantly : Fritz Miiller informs me that the
most luminous insect which he ever beheld in Brazil,
was the larva of some beetle. Both sexes of certain
luminous species of Elater emit light. Kirby and
Spence suspect that the phosphorescence serves to
frighten and drive away enemies.

Difference in Size between the Sexes. — With insects
of all kinds the males are commonly smaller than the
females ; 12 and this difference can often be detected,
even in the larval state. So considerable is the difference

10 E. Doubleday, 'Annals and Mag. of Nat. Hist.' vol. i. 1848, p. 371*.
I may add that the wings in certain Hymen optera (see Shuckard,
Tossorial Hymenup.' 18:37, p. 39-43) differ in neuration according to
sex.

11 H. W. Bates, in ' Journal of Proc. Linn. Soc." vol. vi. 1862,
Mr. Wonfor 's observations are quoted in ' Popidar Science,
1868, p. 343. /<^Q°S H6

12 Kirliv and Spence, ' Introduction to Entomologv,' vol/^H\*Zo9. A.^ 4

LIBRAR

34(5 SEXUAL SELECTION. Pakt II.

between the male and female cocoons of the silk-moth
(Bombyx mori), that in France they are separated by
a particular mode of weighing.13 In the lower classes
of the animal kingdom, the greater size of the females
seems generally to depend on their developing an enor-
mous number of ova ; and this may to a certain extent
hold good with insects. But Dr. Wallace has suggested
a much more probable explanation. He finds, after
carefully attending to the development of the cater-
pillars of Bombyx cynthia and yamamai, and especially
of some dwarfed caterpillars reared from a second brood
on unnatural food, "that in proportion as the indivi-
" dual moth is finer, so is the time required for its
" metamorphosis longer ; and for this reason the female,
" which is the larger and heavier insect, from having to
" carry her numerous eggs, will be preceded by the
" male, which is smaller and has less to mature." 14 Now
as most insects are short-lived, and as they are exposed
to many dangers, it wTould manifestly be advantageous
to the female to be impregnated as soon as possible.
This end would be gained by the males being first
matured in large numbers ready for the advent of the
females ; and this again would naturally follow, as
Mr. A. R. Wallace has remarked,15 through natural
selection ; for the smaller males would be first matured,
and thus would procreate a large number of offspring
which would inherit the reduced size of their male
parents, whilst the larger males from being matured
later would leave fewer offspring.

There are, however, exceptions to the rule of male
insects being smaller than the females; and some of

13 Kobinet, ' Vers a Soie,' 1848, p. 207.

14 ' Transact. Ent. Soc.' 3rd series, vol. v. p. 486.

15 < Journal of Proc. Ent. Soc' Feb. 4th, 1867, p. lxxi.

Chap. X. INSECTS. 347

these exceptions are intelligible. Size and strength
would be an advantage to the males, which fight for the
possession of the female ; and in these cases the males,
as with the stag-beetle (Lucanus), are larger than the
females. There are, however, other beetles which are
not known to fio-ht Wether, of which the males exceed
the females in size ; and the meaning of this fact is not
known ; but in some of these cases, as with the huge
Dynastes and Megasoma, we can at least see that there
would be no necessity for the males to be smaller than
the females, in order to be matured before them, for
these beetles are not short-lived, and there would be
ample time for the pairing of the sexes. So, again,
male dragon-flies (Libellulidse) are sometimes sensibly
larger, and never smaller, than the females ; 16 and
they do not, as Mr. MacLachlan believes, generally
pair with the females, until a week or fortnight has
elapsed, and until they have assumed their proper
masculine colours. But the most curious case, shewing
on what complex and easily-overlooked relations, so
trifling a character as a difference in size between the
sexes may depend, is that of the aculeate Hymenoptera ;
for Mr. F. Smith informs me that throughout nearly
the whole of this large group the males, in accor-
dance with the general rule, are smaller than the
females and emerge about a week before them ; but
amongst the Bees, the males of Apis mellifica, Anthidium
manicatum and Anthophora aeervorum, and amongst the
Fossores, the males of the Methoca ichneumonides, are
larger than the females. The explanation of this ano-
maly is that a marriage-flight is absolutely necessary

16 For this and other statements on the size of the sexes, see Kirhy
and Spenoe, ibid. vol. iii. p. 300 ; on the duration of life in insects,
see p. 314.

348 SEXUAL SELECTION. Part II.

with these species, and the males require great strength
and size in order to carry the females through the air.
Increased size has here been acquired in opposition to
the usual relation between size and the period of de-
velopment, for the males, though larger, emerge before
the smaller females.

We will now review the several Orders, selecting such
facts as more particularly concern us. The Lepidoptera
(Butterflies and Moths) will be retained for a separate
chapter.

Order, Thysanura.—The members of this Order are
lowly organised for their class. They are wingless,
dull-coloured, minute insects, with ugly, almost mis-
shapen heads and bodies. The sexes do not differ ; but
they offer one interesting fact, by shewing that the males
pay sedulous court to their females even low down in the
animal scale. Sir J. Lubbock17 in describing the Smyn-
thurm liiteus, says : " it is very amusing to see these
" little creatures coquetting together. The male, which
" is much smaller than the female, runs round her, and
" they butt one another, standing face to face, and
" moving backward and forward like two playful lambs.
" Then the female pretends to run away and the male
" runs after her with a queer appearance of anger, gets
" in front and stands facing her again ; then she turns
" coyly round, but he, quicker and more active, scuttles
"round too, and seems to whip her with his antennae;
" then for a bit they stand face to face, play with their
" antennae, and seem to be all in all to one another."

Order, Diptera (Flies). — The sexes differ little in
colour. The greatest difference, known to Mr. F. Walker,

" ' Transact. Linnean Soc.' vol. xxvi. 1868, p. 296.

Chap. X. DIPTERA AND HEMTPTERA. 349

is in the genus Bibio, in which the males are blackish
or quite black, and the females obscure brownish-orange.
The genus Elaphomvia, discovered by Mr. Wallace 18 in
New Guinea, is highly remarkable, as the males are
furnished with horns, of which the females are quite
destitute. The horns spring from beneath the eyes, and
curiously resemble those of stags, being either branched
or palmated. They equal in length the whole of the
body in one of the species. They might be thought
to serve for fighting, but as in one species they are
of a beautiful pink colour, edged with black, with a
pale central stripe, and as these insects have altogether
a very elegant appearance, it is perhaps more pro-
bable that the horns serve as ornaments. That the
males of some Diptera fight together is certain ; for
Prof. Westwoed 19 has several times seen this with some
species of Tipula or Harry-long-legs. Many observers
believe that when gnats (Culickke) dance in the air in
a body, alternately rising and falling, the males are
courting the females. The mental faculties of the
Diptera are probably fairly well developed, for their
nervous system is more highly developed than in most
other Orders of insects.20

Order, He?niptera (Field-Bugs). — Mr. J. W. Douglas,
who has particularly attended to the British species, has
kindly aiven me an account of their sexual differences.
The males of some species are furnished with wings,
whilst the females are windless ; the sexes differ in the
form of the body and elytra ; in the second joints of
their antennae and in their tarsi ; but as the signification

18 « The Malay Archipelago,' vol. ii. 1809, p. 313.

19 ' Modern Classification of Insects,' vol. ii. 1840, p. 526.

30 See Mr. B. T. Lowne's very interesting work, ' On the Anatomy of
the Blow-Fly, Mnsca vomitoria,' 1870, p. 14.

350 SEXUAL SELECTION. Part II.

of these differences is quite unknown, they may be here
passed over. The females are generally larger and more
robust than the males. With British, and, as far as
Mr. Douglas knows, with exotic species, the sexes do
not commonly differ much in colour; but in about six
British species the male is considerably darker than the
female, and in about four other species the female is
darker than the male. Both sexes of some species are
beautifully marked with vermilion and black. It is
doubtful whether these colours serve as a protection.
If in any species the males had differed from the females
in an analogous manner, we might have been justified
in attributing such conspicuous colours to sexual selec-
tion with transference to both sexes.

Some species of Keduvidae make a stridulating noise ;
and, in the case of Pirates stridulus, this is said21 to
be effected by the movement of the neck within the
pro-thoracic cavity. According to Westring, Beduvius
personatus also stridulates. But I have not been able
to learn any particulars about these insects ; nor have I
any reason to suppose that they differ sexually in this
respect.

Order, Homojrfera. — Every one who has wandered in
a tropical forest must have been astonished at the din
made by the male Cicadaa. The females are mute ;
as the Grecian poet Xenarchus says, " Happy the
" Cicadas live, since they all have voiceless wives."
The noise thus made could be plainly heard on board
the " Beagle," when anchored at a quarter of a mile
from the shore of Brazil ; and Captain Hancock says
it can be heard at the distance of a mile. The Greeks
formerly kept, and the Chinese now keep, these insects

21 Westwocd, ' Modern Class, of Insects,' vol. ii. p. 473.

Chap. X. HOMOPTEKA. 351

in caires for the sake of their sods:, so that it must be
pleasing to the ears of some men.22 The Cicadida3
usually sing during the day ; whilst the Fulgoridaa
appear to be night-songsters. The sound, according
to Landois,23 who has recently studied the subject, is
produced by the vibration of the lips of the spiracles,
which are set into motion by a current of air emitted
from the tracheae. It is increased by a wonderfully
complex resounding apparatus, consisting of two cavi-
ties covered by scales. Hence the sound may truly
be called a voice. In the female the musical apparatus
is present, but very much less developed than in the
male, and is never used for producing sound.

With respect to the object of the music, Dr. Hartman
in speaking of the Cicada sejrfemdecim of the United
States, says,24 " the drums are now (June 6th and 7th,
" 1851) heard in all directions. This I believe to be the
" marital summons from the males. Standing in thick
" chestnut sprouts about as high as my head, where
" hundreds were around me, I observed the females
" coming around the drumming males." He adds, " this
" season (Aug. 1868) a dwarf pear-tree in my garden
" produced about fifty larvae of Cic. pruinosa; and I
" several times noticed the females to alight near a
" male while he was uttering his clanging notes." Fritz
Miiller writes to me from S. Brazil that he has often
listened to a musical contest between two or three
males of a Cicada, having a particularly loud voice, and
seated at a considerable distance from each other. As

22 These particulars are taken from Westwood's ' Modern Class, of
Insects' v«l. ii. 1840, p. 422. See, also, on the Fulgoridse, Kirby and
Spciice, Introduce' vol. ii. p. 401.

23 ' Zeitschrift fiir wissenschait. Zcolog.' B. xvii. 1867. s. 152-158.

21 lam indebted to Mr. Walsh for having sent me this extract from
a ' Journal of the Doings of Cicada septemdecim,' by Dr. Hartman.

352 SEXUAL SELECTION. Part II.

soon as the first had finished his sonp\ a second im-
mediately began ; and after he had concluded, another
began, and so on. As there is so much rivalry between
the males, it is probable that the females not only dis-
cover them by the sounds emitted, but that, like female
birds, they are excited or allured by the male with the
most attractive voice.

I have not found any well-marked cases of orna-
mental differences between the sexes of the Homoptera.
Mr. Douglas informs me that there are three British
species, in which the male is black or marked with black
bands, whilst the females are pale-coloured or obscure.

Order, Orthoptera. — The males in the three salta-
torial families belonging to this Order are remark-
able for their musical powers, namely the Achetidse or
crickets, the Locustidae for which there is no exact
equivalent name in English, and the Acridiidge or grass-
hoppers. The stridulation produced by some of the
Locustidae is so loud that it can be heard durino- the
night at the distance of a mile ;25 and that made by
certain species is not unmusical even to the human
ear, so that the Indians on the Amazons keep them
in wicker ca^es. All observers agree that the sounds
serve either to call or excite the mute females. But it
has been noticed26 that the male migratory locust of
Russia (one of the Acridiidae) whilst coupled with the
female, stridulates from anger or jealousy when ap-
proached by another male. The house-cricket when
surprised at night uses its voice to warn its fellows.27 In
North America the Katy-did (Platyphyllum concavum,

25 L. Guikling, ' Transact. Linn. Soc' vol. xv. p. 154.

26 Koppen, as quoted in the ' Zoological Record ' for 1867, p. 46'K

27 Gilbert White, ' Nat. Hist, of Seiborne,' vol. ii. 1825, p. 262.

Chap. X.

ORTHOPTERA.

353

one of the Locustidae) is described28 as mounting on the
upper branches of a tree, and in the evening beginning
" his noisy babble, while rival notes issue from the neigh-
" bouring trees, and the groves resound with the call of
" Katy-did-she-did, the live-long night." Mr. Bates, in
speaking of the European field-cricket (one of the Ache-
tidae), says, " the male has been observed to place itself
" in the evening at the entrance of its' burrow, and
" stridulate until a female approaches, when the louder
" notes are succeeded by a more subdued tone, whilst
" the successful musician caresses with his antennae
"the mate he has won."29
Dr. Scudder was able to
excite one of these insects
to answer him, by rubbing
on a file with a quill.30
In both sexes a remark-
able auditory apparatus
has been discovered by
Von Siebold, situated in
the front legs.31

In the three Families

the SOUnds are differently Fig. 10. Gryllus campestris (from Landois).

•nrnrlnnprl Tn tllP TYIfllpS of Right-band figure, under side of part of the
prOUUCea. ±U Uie Illdieb Ul wing-nervure, much magnified, showing

the Achetidae both wing

the teeth, st.
Left-band figure, upper surface of wing-cover,
with the projecting, smooth nervure, r.,
across which the teeth (si) are scraped.

covers have the same
structure ; and this in the
field-cricket (Gryllus campestris, fig. 10) consists, as de

23 Harris, 'Insects of New England,' 1842, p. 128.

29 « xhe Naturalist on the Amazons,' vol. i. 1863, p. 252. Mr. Bates
gives a very interesting discussion on the gradations in the musical
apparatus of the three "families. See also Westwood, 'Modern Class.'
vol. ii. p. 445 and 453.

30 'Proc. Boston Soc. of Nat. Hist.' vol. xi. April, 18G8.

31 ' Nouveau Manuel d'Anat. Comp.' (French translat.), torn. i. 1850
p. 567.

VOL. I. 2 A

354 SEXUAL SELECTION. Part II.

scribed by Landois,32 of from 131 to 138 sharp, trans-
verse ridges or teeth (st) on the under side of one of the
nervures of the wing-cover. This toothed nervure is
rapidly scraped across a projecting, smooth, hard nervure
(r) on the upper surface of the opposite wing. First

one wing is rubbed over the other,
and then the movement is reversed.
Both wings are raised a little at the
same time, so as to increase the re-
sonance. In some species the wing-
covers of the males are furnished at
the base with a talc-like plate.33 I
have here given a drawing (fig. 11)
Fig. ii. Teeth of Nervure of the teeth on the under side of the

of Gryllus domesticus ? ,i • j? ri 11

(from Landois). nervure oi another species 01 Uryllus,

viz. G. domesticus.

In the Locustidse the opposite wing-covers differ in
structure (fig. 12), and cannot, as in the last family,
be indifferently used in a reversed manner. The left
wing, which acts as the bow of the fiddle, lies over the
right wing which serves as the fiddle itself. One of
the nervures (a) on the under surface of the former is
finely serrated, and is scraped across the prominent
nervures on the upper surface of the opposite or right
wing. In our British Pliasgonura viridissima it ap-
peared to me that the serrated nervure is rubbed
against the rounded hind corner of the opposite wing,
the edge of which is thickened, coloured brown, and
very sharp. In the right wing, but not in the left,
there is a little plate, as transrjarent as talc, surrounded
by nervures, and called the speculum. In Ephippig&r
vitium, a member of this same family, we have a curious

3:- ' Zeitsehrift fur wissenschaft. Zoolog.' B. xvii. 18G7, s. 117.
:i3 Westwood, ' Modern Class, of Insects,' vol. i. p. 440.

Chap. X.

ORTHOPTERA.

355

subordinate modification ; for the wing-covers are greatly
reduced in size, but " the posterior part of the pro-thorax
" is elevated into a kind of dome over the wing-covers,
" and which has probably the effect of increasing the
" sound." 34

Fig. 12. (Jblorocoilus Tanana (from Bates), a, b. Lobes of opposite wing-covers.

We thus see that the musical apparatus is more
differentiated or specialised in the Locustidse, which
includes I believe the most powerful performers in
the Order, than in the Achetidse, in which both wing-
covers have the same structure and the same function.35
Landois, however, detected in one of the Locustidse,
namelv in Decticus, a short and narrow row of small

31 Westwood, ' Modern Cla&s. of Insects,' vol. i. p. 453.
35 Landois, ibid. s. 121, 122.

2 a 2

356 SEXUAL SELECTION. Part II.

teeth, mere rudiments, on the inferior surface of the
right wing-cover, which underlies the other and is
never used as the bow. I observed the same rudi-
mentary structure on the under side of the right wing-
cover in Phasgonura viridissima. Hence we may with
confidence infer that the Locustidaa are descended from
a form, in which, as in the existing Achetidas, both
wing-covers had serrated nervures on the under surface,
and could be indifferently used as the bow ; but that
in the Locustidae the two wing-covers gradually became
differentiated and perfected, on the principle of the divi-
sion of labour, the one to act exclusively as the bow and
the other as the fiddle. By what steps the more simple
apparatus in the Achetidae originated, we do not know,
but it is probable that the basal portions of the wing-
covers overlapped each other formerly as at present, and
that the friction of the nervures produced a grating
sound, as I find is now the case with the wing-covers
of the females.36 A grating sound thus occasionally
and accidentally made by the males, if it served them
ever so little as a love-call to the females, might readily
have been intensified through sexual selection by fitting
variations in the roughness of the nervures having been
continually preserved.

In the last and third Family, namely the Acridiidae
or grasshoppers, the stridulation is produced in a very
difYerent manner, and is not so shrill, according to Dr.
Scudder, as in the preceding Families. The inner sur-
face of the femur (fig. 13, r) is furnished with a longi-
tudinal row of minute, elegant, lancet-shaped, elastic
teeth, from 85 to 93 in number ;37 and these are scraped

36 Mr. Walsh also informs me that he has noticed that the female of
the Platyphyttum concavu.m, "when captured makes a feeble grating
" noise by shuffling her wing-covers together."

37 Landois, ibid. s. 113.

Chap. X.

ORTHOPTERA.

357

u

Fig. 13. Hind-leg of Stenobothrus pratorura:
r, the striduiating ridge ; lower figure, the
teeth, forming the ridge, much magnified
(from Landois).

across the sharp, projecting nervures on the wing-covers,

which are thus made to vibrate and resound. Harris3

says that when one of

the males begins to play,

he first " bends the shank

" of the hind-leg beneath

" the thigh, where it is

" lodged in a furrow de-

" signed to receive it,

" and then draws the lee

a

briskly up and down.

He does not play both
" fiddles together, but al-
" ternately first upon one
" and then on the other."
In many species, the base
of the abdomen is hollowed out into a great cavity
which is believed to act as a resounding board. In
Pneumora (fig. 14), a S. African genus belonging to
this same family, we meet with a new and remarkable
modification : in the males a small notched ridge pro-
jects obliquely from each side of the abdomen, against
which the hind femora are rubbed.39 As the male is
furnished with wings, the female being wingless, it is
remarkable that the thighs are not rubbed in the usual
manner against the wing-covers ; but this may perhaps
be accounted for by the unusually small size of the hind-
legs. I have not been able to examine the inner
surface of the thighs, which, judging from analogy,
would be finely serrated. The species of Pneumora
have been more profoundly modified for the sake of
stridulation than any other orthopterous insect; for

3S 'Insects of New England,' 1842, p. 133.

M Westwood, ' Modern Classification,' vol. i. p. 462.

358

SEXUAL SELECTION.

Part II.

in the male the whole body has been converted into a
musical instrument, being distended with air, like a
great pellucid bladder, so as to increase the resonance.
Mr. Trimen informs me that at the Cape of Good Hope
these insects make a wonderful noise during the night

Fig. 14. Pneumora (from specimens in the British Museum). Upper figure, male ;

lower figure, female.

There is one exception to the rule that the females
in these three Families are destitute of an efficient
musical apparatus ; for both sexes of Ephippiger (Lo-
custidae) are said40 to be thus provided. This case may

40 Westwood, ibid. vol. i. p. 453.

Chap. X. ORTHOPTERA. 359

be compared with that of the reindeer, in which species
alone both sexes possess horns. Although the female
orthoptera are thus almost invariably mute, yet Landois41
found rudiments of the stridulating organs on the fe-
mora of the female Acridiidae, and similar rudiments on the
under surface of the wing-covers of the female Achetidae ;
but he failed to find any rudiments in the females
of Decticus, one of the Locustidae. In the Homoptera
the mute females of Cicada, have the proper musical
apparatus in an undeveloped state ; and we shall here-
after meet in other divisions of the animal kingdom with
innumerable instances of structures proper to the male
being present in a rudimentary condition in the female.
Such cases appear at first sight to indicate that both
sexes were primordially constructed in the same manner,
but that certain organs were subsequently lost by the
females. It is, however, a more probable view, as pre-
viously explained, that the organs in question were
acquired by the males and partially transferred to the
females.

Landois has observed another interesting fact, namely
that in the females of the Acridiidae, the stridulating
teeth on the femora remain throughout life in the same
condition in which they first appear in both sexes
during the larval state. In the males, on the other
hand, they become fully developed and acquire their
perfect structure at the last moult, when the insect is
mature and ready to breed.

From the facts now given, we see that the means
by which the males produce their sounds are extremely
diversified in the Orthoptera, and .are altogether dif-
ferent from those employed by the Homoptera. But
throughout the animal kingdom we incessantly find the

41 Landois, ibid. s. 115, 116, 120, 122.

360 SEXUAL SELECTION. Part II.

same object gained by the most diversified means; this
being due to the whole organisation undergoing in
the course of ages multifarious changes ; and as part
after part varies, different variations are taken advantage
of for the same general purpose. The diversification of
the means for producing sound in the three families
of the Orthoptera and in the Homoptera, impresses the
mind with the high importance of these structures to
the males, for the sake of calling or alluring the females.
We need feel no surprise at the amount of modification
which the Orthoptera have undergone in this respect, as
we now know, from Dr. Scudder's remarkable discovery,42
that there has been more than ample time. This
naturalist has lately found a fossil insect in the Devonian
formation of New Brunswick, which is furnished with
" the well-known tympanum or stridulating apparatus
" of the male Locustidae." This insect, though in most
respects related to the Neuroptera, appears to connect,
as is so often the case with very ancient forms, the
two Orders of the Neuroptera and Orthoptera which are
now generally ranked as quite distinct.

I have but little more to say on the Orthoptera.
Some of the species are very pugnacious : when two
male field-crickets (GryJIus camjiestris) are confined
together, they fight till one kills the other ; and the
sj)ecies of Mantis are described as manoeuvring with
their sword-like front-limbs, like hussars with their
sabres. The Chinese keep these insects in little bamboo
cages and match them like game-cocks.43 With respect
to colour, some exotic locusts are beautifully orna-
mented ; the posterior wings being marked with red,

42 ' Transact. Ent. Soc' 3rd series, vol. ii. (' Journal of Proceedings,
p. 117.)

43 Wefctwood, 'Modern Class, of Insects,' vol. i. p. 427; for crickets,
p. 445.

Chap. X. NEUROPTERA. 361

blue, and black; but as throughout the Order the
two sexes rarely differ much in colour, it is doubtful
whether they owe these bright tints to sexual selection.
Conspicuous colours may be of use to these insects
as a protection, on the principle to be explained in the
next chapter, by giving notice to their enemies that
they are unpalatable. Thus it has been observed44
that an Indian brightly-coloured locust was invariably
rejected when offered to birds and lizards. Some cases,
however, of sexual differences in colour in this Order
are known. The male of an American cricket45 is de-
scribed as being as white as ivory, whilst the female
varies from almost white to greenish-yellow or dusky.
Mr. Walsh informs me that the adult male of Spectrum
femoratum (one of the Phasinidae) " is of a shining
" brownish-yellow colour; the adult female being of
" a dull, opaque, cinereous-brown; the young of both
" sexes being green." Lastly, I may mention that the
male of one curious kind of cricket 46 is furnished with
" a long membranous appendage, which falls over the
" face like a veil ; " but whether this serves as an orna-
ment is not known.

Order, Neuroptera. — Little need here be said, except
in regard to colour. In the Ephenieridae the sexes
often differ slightly in their obscure tints;47 but it is
not probable that the males are thus rendered attrac-
tive to the females. The Libellulidae or dragon-flies
are ornamented with splendid green, blue, yellow, and

44 Mr. Ch. Home, in < Proc. Ent. Soc' May ^3, 1869, p. xii.

45 The Oecanthus nivalis. Harris, 'Insects of New EDgland,' 1842,
p. 124.

46 Platybknmus : Westwood, ' Modern. Class.' vol. i. p. 447.

4? B. D. Walsh, the Pseudo-neuroptera of Illinois, in 'Proc. Ent. Soc.
of Philadelphia,' 1802, p. 3G1.

362 SEXUAL SELECTION. Part II.

vermilion metallic tints; and the sexes often differ.
Thus, the males of some of the Agrionidse, as Prof.
Westwood remarks,48 "are of a rich blue with black
" wings, whilst the females are fine green with colourless
"wings." But in Agrion Ramburii these colours are
exactly reversed in the two sexes.49 In the extensive
N. American genus of Hetaerina, the males alone have
a beautiful carmine spot at the base of each wing. In
Anax Junius the basal part of the abdomen in the male
is a vivid ultra-marine blue, and in the female grass-
green. In the allied genus Gomphus, on the other
hand, and in some other genera, the sexes differ but
little in colour. Throughout the animal kingdom,
similar cases of the sexes of closely-allied forms either
differing greatly, or very little, or not at all, are of
frequent occurrence. Although with many Libellulidae
there is so wide a difference in colour between the sexes,
it is often difficult to say which is the most brilliant ;
and the ordinary coloration of the two sexes is exactly
reversed, as we have just seen, in one species of Agrion.
It is not probable that their colours in any case have
been gained as a protection. As Mr. MacLachlan, who
has closely attended to this family, writes to me, dragon-
flies — the tyrants of the insect-world — are the least
liable of any insect to be attacked by birds or other
enemies. He believes that their bright colours serve
as a sexual attraction. It deserves notice, as bearing
on this subject, that certain dragon-flies appear to be
attracted by particular colours : Mr. Patterson observed50
that the species of Agrionidse, of which the males are
blue, settled in numbers on the blue float of a fishing

48 ' Modern Class.' vol. ii. p. 37.

49 Walsh, ibid. p. 381. I am indebted to this naturalist for the
following facts on Hetsorina, Anax, and Gomphus.

50 ' Transact. Ent. Soc' vol. i. 1836, p. lxxxi.

Chap. X. NEUEOPTEKA. 363

line ; whilst two other species were attracted by shining
white colours.

It is an interesting fact, first observed by Schelver,
that the males, in several genera belonging to two sub-
families, when they first emerge from the pupal state
are coloured exactly like the females ; but that their
bodies in a short time assume a conspicuous milky-blue
tint, owing to the exudation of a kind of oil, soluble in
ether and alcohol. Mr. MacLachlan believes that in the
male of Libellula depressa this change of colour does not
occur until nearly a fortnight after the metamorphosis,
when the sexes are ready to pair.

Certain species of Neurothemis present, according
to Brauer51 a curious case of dimorphism, some of the
females having their wings netted in the usual manner ;
whilst other females have them " very richly netted as in
" the males of the same species." Brauer " explains
" the phenomenon on Darwinian principles by the
" supposition that the close netting of the veins is a
" secondary sexual character in the males." This
latter character is generally developed in the males
alone, but being, like every other masculine character,
latent in the female, is occasionally developed in them.
We have here an illustration of the manner in which
the two sexes of many animals have probably come to
resemble each other, namely by variations first appear-
ing in the males, being preserved in them, and then
transmitted to and developed in the females ; but in
this particular genus a complete transference is occa-
sionally and abruptly effected. Mr. MacLachlan in-
forms me of another case of dimorphism occurring in
several species of Agrion in which a certain number of
individuals are found of an orange colour, and these are

51 See abstract in the 'Zoological Record ' fur 18G7, p. 4^0.

364 SEXUAL SELECTION. Part II.

invariably females. This is probably a case of reversion,
for in the true Libellulae, when the sexes differ in
colour, the females are always orange or yellow, so
that supposing Agrion to be descended from some pri-
mordial form having the characteristic sexual colours
of the typical Libellulre, it would not be surprising that
a tendency to vary in this manner should occur in the
females alone.

Although many dragon-flies are such large, powerful,
and fierce insects, the males have not been observed
by Mr. MacLachlan to fight together, except, as he
believes, in the case of some of the smaller species of
Agrion. In another very distinct group in this Order,
namely in the Termites or white auts, both sexes at
the time of swarming may be seen running about, " the
" male after the female, sometimes two chasing one
t( female, and contending with great eagerness who shall
" win the prize." 52

Order, Hymenoptera. — That inimitable observer, M.
Fabre,53 in describing the habits of Cerceris, a wasp-
like insect, remarks that " fights frequently eusue
" between the males for the possession of some parti*
" cular female, who sits an apparently unconcerned
** beholder of the struggle for supremacy, and when the
" victory is decided, quietly flies away in company
" with the conqueror." Westwood M says that the
males of one of the saw-flies (Tenthredinse) " have been
" found fighting together, with their mandibles locked."
As M. Fabre speaks of the males of Cerceris striving
to obtain a particular female, it may be well to bear in

52 Kirby and Spence, ' Introduct. to Entomology,' vol. ii. 1818, p. 35.

53 See an interesting article, " The Writings of Fabre," in ' Nat. Hist.
Review,' April, 1802, p. 122.

54 'Journal of Proe. of Entomolcg. Soc.' Sept. 7th, 18G3, p. 169.

Chap. X. HYMENOPTERA. 365

mind that insects belonging to this Order have the
power of recognising each other after long intervals of
time, and are deeply attached. For instance, Pierre
Huber, whose accuracy no one doubts, separated some
ants, and when after an interval of four months they
met others which had formerly belonged to the same
community, they mutually recognised and caressed each
other with their antennae. Had they been strangers
they would have fought together. Again, when two
communities engage in a battle, the ants on the same
side in the general confusion sometimes attack each
other, but they soon perceive their mistake, and the
one ant soothes the other.55

In this Order slight differences in colour, according
to sex, are common, but conspicuous differences are
rare except in the family of Bees; yet both sexes of
certain groups are so brilliantly coloured — for instance
in Chrysis, in which vermilion and metallic greens
prevail — that we are tempted to attribute the result
to sexual selection. In the Ichneumonidaa, according to
Mr. Walsh,56 the males are almost universally lighter
coloured than the females. On the other hand, in the
Tenthredinidse the males are generally darker than the
females. In the Siricidse the sexes frequently differ;
thus the male of Sirex juvencus is banded with orange,
whilst the female is dark purple ; but it is difficult to
say which sex is the most ornamented. In Tremex
columbte the female is much brighter coloured than the
male. With ants, as I am informed by Mr. F. Smith,
the males of several species are black, the females
being testaceous. In the family Of Bees, especially in

55 P. Huber, ' Recherches sur les Moeurs des Fourmis,' 1810, p. 150,
165.

56 ' Pi-oc. Eutomolog. Soc. of Philadelphia,' I860, p. 238-239.

366 SEXUAL SELECTION". Part II.

the solitary species, as I hear frorn the same distin-
guished entomologist, the sexes often differ in colour.
The males are generally the brightest, and in Bombus
as well as in Apathus, much more variable in colour
than the females. In Anthophora retusa the male ' is
of a rich fulvous-brown, whilst the female is quite
black: so are the females of several species of Xylocopa,
the males being bright yellow. In an Australian bee
(Lestis bombylans), the female is of an extremely brilliant
steel-blue, sometimes tinted with vivid green ; the male
being of a bright brassy colour clothed with rich fulvous
pubescence. As in this group the females are provided
with excellent defensive weapons in their stings, it is
not probable that they have come to differ in colour
from the males for the sake of protection.

Mutilla Europsea emits a stridulating noise; and ac-
cording to Groureau 57 both sexes have this power. He
attributes the sound to the friction of the third and
preceding abdominal segments ; and I find that these
surfaces are marked with very fine concentric ridges,
but so is the projecting thoracic collar, on which the
head articulates ; and this collar, when scratched with
the point of a needle, emits the proper sound. It is
rather surprising that both sexes should have the
power of stridulating, as the male is winged and the
female wingless. It is notorious that Bees express
certain emotions, as of anger, by the tone of their
humming, as do some dipterous insects ; but I have
not referred to these sounds, as they are not known to
be in any way connected with the act of courtship.

Order, Coleoptera (Beetles). — Many beetles are
coloured so as to resemble the surfaces which thev

57 Quoted by Westwood, 'Modern Class, of Insects,' vol. ii. p. 214.

Chap. X. COLEOPTERA. 367

habitually frequent. Other species are ornamented
with gorgeous metallic tints, — for instance, many Cara-
bidae, which live on the ground and have the power
of defending themselves by an intensely acrid secretion,
— the splendid diamond-beetles which are protected by
an extremely hard covering, — many species of Chry-
somela, such as C. cerealis, a large species beautifully
striped with various colours, and in Britain confined
to the bare summit of Snowdon, — and a host of other
species. These splendid colours, which are often
arranged in stripes, spots, crosses and other elegant
patterns, can hardly be beneficial, as a protection, except
in the case of some flower-feeding species; and we
cannot believe that they are purposeless. Hence the
suspicion arises, that they serve as a sexual attraction ;
but we have no evidence on this head, for the sexes
rarely differ in colour. Blind beetles, which cannot of
course behold each other's beauty, never exhibit, as I
hear from Mr. Waterhouse, jun., bright colours, though
they often have polished coats : but the explanation of
their obscurity may be that blind insects inhabit caves
and other obscure stations.

Some Longicorns, however, especially certain Pri-
onidte, offer an exception to the common rule that the
sexes of beetles do not differ in colour. Most of these
insects are large and splendidly coloured. The males in
the genus Pyrodes,58 as I saw in Mr. Bates' collection, are

53 Pyrodes pulcherriinus, in which the sexes differ conspicuously, has
been described by Mr. Bates in ' Transact. Ent. Soc' 1869, p. 50. I
will specify the few other cases in -which I have heard of a difference
in colour between the sexes of beetles. Kirby and Spence (' Introduct.
to Entomology,' vol. iii. p. 301) mention a Cantharis, Meloe, Rhagium,
and the Leptura testacca ; the male of the latter being testaceous, with
a black thorax, and the female of a dull red all over. These two
latter beetles belong to the Order of Longicorns. Messrs. R. Trimen
and Waterhouse, junr., inform me of two Lamellicorns, viz., a Peri-

368

SEXUAL SELECTION.

Part II.

generally redder but rather duller than the females, the
latter being coloured of a more or less splendid golden
green. On the other hand, in one species the male is
golden-green, the female being richly tinted with red
and purple. In the genus Esmeralda the sexes differ so
greatly in colour that they have been ranked as distinct
species : in one species both are of a beautiful shining
green, but the male has a red thorax. On the whole,
as far as I could judge, the females of those Prionidse,
in which the sexes differ, are coloured more richly
than the males; and this does not accord with the
common rule in regard to colour when acquired through
sexual selection.

Fig. 10. Chalco^oma atlas. Upper figure, male (reduced) ; lower figure, female

(nat. size).

tricliia and Trichius, the male of the latter being more obscurely
coloured than the female. In Tillus elongatus the male is black, and
the female always, as it is believed, of a dark blue colour with a red
thorax. The male, also, of Orsodacna atra, as I hear from Mr. Walsh,
is black, the female (the so-called 0. ruficollis) having a rufous thorax.

Chap. X.

COLEOPTERA.

369

Copris isidis. (Left-hand figures, males.)

Phanseus faun us.

Fig. 18.

Dipolicus canton,

Fig. 19.
VOL, I.

Onthopliagus rangifer, enlarged.

2 B

370 SEXUAL SELECTION. Part II.

A most remarkable distinction between the sexes of
many beetles is presented by the great horns which
rise from the head, thorax, or clypeus of the males ;
and in some few cases from the under surface of the
body. These horns, in the great family of the Lamelli-
corns, resemble those of various quadrupeds, such as
stags, rhinoceroses, &c, and are wonderful both from
their size and diversified shapes. Instead of describing
them, I have given figures of the males and females of
some of the more remarkable forms. (Figs. 15 to 1 9.)
The females generally exhibit rudiments of the horns
in the form of small knobs or ridges; but some are
destitute of even a rudiment. On the other hand, the
horns are nearly as well developed in the female as in
the male of Phanseus Jancifer ; and only a little less
well developed in the females of some other species of
the same genus and of Copris. In the several sub-
divisions of the family, the differences in structure of
the horns do not run parallel, as I am informed by
Mr. Bates, with their more important and characteristic
differences ; thus within the same natural section of the
genus Onthophagus, there are species which have either
a single cephalic horn, or two distinct horns.

In almost all cases, the horns are remarkable from
their excessive variability; so that a graduated series
can be formed, from the most highly developed males
to others so degenerate that they can barely be distin-
guished from the females. Mr Walsh59 found that in
Phaneeus camifex the horns were thrice as long in some
males as in others. Mr. Bates, after examining above
a hundred males of Onthophagus rangifer (fig. 19),
thought that he had at last discovered a species in

39 'Proc. Entomolog. Soc. of Philadelphia,' 1864, p. 228.

Chap. X. COLEOPTERA. 371

which the horns did not vary ; but further research
proved the contrary.

The extraordinary size of the horns, and their widely
different structure in closely-allied forms, indicate that
they have been formed for some important purpose;
but their excessive variability in the males of the same
species leads to the inference that this purpose cannot
be of a definite nature. The horns do not show marks
of friction, as if used for any ordinary work. Some
authors suppose r,° that as the males wander much more
than the females, they require horns as a defence
against their enemies; but in many cases the horns
do not seem well adapted for defence, as they are not
sharp. The most obvious conjecture is that they are
used by the males for fighting together ; but they
have never been observed to fight ; nor could Mr. Bates,
after a careful examination of numerous species, find
any sufficient evidence in their mutilated or broken
condition of their having been thus used. If the males
had been habitual fighters, their size would probably
have been increased through sexual selection, so as to
have exceeded that of the female ; but Mr. Bates, after
comparing the two sexes in above a hundred species of
the Copridse, does not find in well-developed individuals
any marked difference in this respect. There is, more-
over, one beetle, belonging to the same great division
of the Lamellicorns, namely Lethrus, the males of which
are known to fight, but they are not provided with
horns, though their mandibles are much larger than
those of the female.

The conclusion, which best agrees with the fact of
the horns having been so immensely yet not fixedly
developed, — as shewn by their extreme variability in

60 Kirby and Spence, ' Introduct. Entomolog.' vol. iii. p. 300.

2 b 2

372

SEXUAL SELECTION.

Part II.

the same species and by their extreme diversity in
closely-allied species — is that they have been acquired
as ornaments. This view will at first appear extremely
improbable; but we shall hereafter find with many
animals, standing much higher in the scale, namely
fishes, amphibians, reptiles and birds, that various
kinds of crests, knobs, horns and combs have been
developed apparently for this sole purpose.

The males of Onitis furcifer (fig. 20) are furnished
with singular projections on their anterior femora, and

with a great fork or pair of horns on
the lower surface of the thorax. This
situation seems extremely ill adapted
for the display of these projections,
and they may be of some real ser-
vice ; but no use can at present be
assigned to them. It is a highly
remarkable fact, that although the
Fig. 2o. onitis furcifer male, mal eg do t ^^bit even a trace of

viewed from beneath.

horns on the upper surface of the
body, yet in the females a rudiment of a single horn on
the head (fig. 21, a), and of a crest (6) on the thorax,
are plainly visible. That the slight thoracic crest in the

&

Fig. 21. Left-hand figure, male of Onitis furcifer, viewed laterally. Pdght-hand figure,
female. a. Rudiment of cephalic horn. b. Trace of thoracic horn or crest.

female is a rudiment of a projection proper to the male,
though entirely absent in the male of this particular
species, is clear : for the female of Bubas bison (a form

Chap. X. COLEOPTEEA. 373

which comes next to Onitis) has a similar slight crest
on the thorax, and the male has in the same situation a
great projection. So again there can be no doubt that
the little point (a) on the head of the female Onitis
furcifer, as well of the females of two or three allied
species, is a rudimentary representative of the cephalic
horn, which is common to the males of so many lamel-
licorn beetles, as in Phaneeus, fig. 17. The males indeed
of some unnamed beetles in the British Museum, which
are believed actually to belong to the genus Onitis, are
furnished with a similar horn. The remarkable nature
of this case will be best perceived by an illustration :
the Ruminant quadrupeds run parallel with the laniel-
licorn beetles, in some females possessing horns as large
as those of the male, in others having them much
smaller, or existing as mere rudiments (though this is
as rare with ruminants as it is common writh Lamelli-
corns), or in having none at all. Now if a new species
of deer or sheep were discovered with the female
bearing distinct rudiments of horns, whilst the head
of the male was absolutely smooth, we should have a
case like that of Onitis furcifer.

In this case the old belief of rudiments having been
created to complete the scheme of nature is so far from
holding good, that all ordinary rules are completely
broken through. The view which seems the most pro-
bable is that some early progenitor of Onitis acquired,
like other Lamellicorns, horns on the head and thorax,
and then transferred them, in a rudimentary condition,
as with so many existing species, to the female, by whom
they have ever since been retained. The subsequent
loss of the horns by the male may have resulted through
the principle of compensation from the development of
the projections on the lower surface, whilst the female
has not been thus affected, as she is not furnished with

374 SEXUAL SELECTION. Part II.

these projections, and consequently has retained the
rudiments of the horns on the upper surface. Although
this view is supported by the case of Bleclius imme-
diately to be given, yet the projections on the lower
surface differ greatly in structure and development in
the males of the several species of Onitis, and are even
rudimentary in some ; nevertheless the upper surface
in all these species is quite destitute of horns. As
secondary sexual characters are so eminently variable, it
is possible that the projections on the lower surface may
have been first acquired by some progenitor of Onitis and
produced their effect through compensation, and then
have been in certain cases almost completely lost.

All the cases hitherto given refer to the Lam ell i-
corns, but the males of some few other beetles, be-
longing to two widely distinct groups, namely, the
Curculioniclse and Staphylinidse, are furnished with
horns, — in the former on the lower surface of the body,61
in the latter on the upper surface of the head and
thorax. In the Staphylinidse the horns of the males
in the same species are extraordinarily variable, just
as we have seen with the Lamellicorns. In Siagonium

9

£}-, o v -v s ■ t

Fig. 22. Bledius taurus, magnified. Left-hand figure, male; right-hand figure, female.

we have a case of dimorphism, for the males can be
divided into two sets, differing greatly in the size of
their bodies, and in the development of their horns,
without any intermediate gradations. In a species of
Bledius (fig. 22), also belonging to the Staphylinidse,
male specimens can be found in the same locality, as

61 Kirby and Spence, ibid. vol. iii. p. 329.

Chap. X. COLEOPTERA. 375

Professor Westwood states, " in which the central horn
'* of the thorax is very largp, but the horns of the head
" quite rudimental ; and others, in which the thoracic
" horn is much shorter, whilst the protuberances on
" the head are long." 62 Here, then, we apparently
have an instance of compensation of growth, which
throws light on the curious case just given of the loss of
the upper horns by the males of Onitis furcifer.

Law of Battle. — Some male beetles, which seem ill
fitted for fighting, nevertheless engage in conflicts for
the possession of the females. Mr. Wallace 63 saw two
males of Leptorliynchus angustatus, a linear beetle with
a much elongated rostrum, " fighting for a female, who
" stood close by busy at her boring. They pushed at
" each other with their rostra, and clawed and thumped,
" apparently in the greatest rage." The smaller male,
however, " soon ran away, acknowledging himself van-
" quished." In some few cases the males are well
adapted for fighting, by possessing great toothed man-
dibles, much larger than those of the females. This
is the case with the common stag-beetle {Lucanus
cervus), the males of which emerge from the pupal state
about a week before the other sex, so that several may
often be seen pursuing the same female. At this period
they engage in fierce conflicts. When Mr. A. H.
Davis64 enclosed two males with one female in a box,
the larger male severely pinched the smaller one, until
he resigned his pretensions. A friend informs me

02 ' Modern Classification of Insects,' vol. i. p. 172. On the same
page there is an account of Siagouium. In the British Museum I
noticed one male specimen of Siagonium in an intermediate condition,
so that the dimorphism is not strict.

63 ' The Malay Archipelago,' vol. ii. 1869, p. 27G.

64 ' Entomological Magazine,' vol. i. 1833, p. 82. See also on the
conflicts of tliis species, Kirby and Spence, ibid. vol. iii. p. 311; and
Westwood, ibid. vol. i. p. 187.

376 SEXUAL SELECTIOX. Part II.

that when a boy he often put the males together to see
them fight, and he noticed that they were much bolder
and fiercer than the females, as is well known to be the
case with the higher animals. The males would seize
hold of his finger, if held in front, but not so the females.
With many of the Lucanidse, as well as with the abo^e-
mentioned Leptorhynchus, the males are larger and
more powerful insects than the females. The two sexes
of Lethrus cejplialotes (one of the Lamellicorns) inhabit the
same burrow ; and the male has larger mandibles than
the female. If, during the breeding-season, a strenge
male attempts to enter the burrow, he is attacked ; the
female does not remain passive, but closes the mouth of
the burrow, and encourages her mate by continually
pushing him on from behind. The action does not
cease until the aggressor is killed or runs away.65 The
two sexes of another lamellicorn beetle, the Ateuchus
cicatricosus live in pairs, and seem much attached to
each other; the male excites the female to roll the
balls of dung in which the ova are deposited ; and if
she is removed, he becomes much agitated. If the
male is removed, the female ceases all work, and as
M. Brulerie 66 believes, would remain on the spot until
she died.

The great mandibles of the male Lucanidse are ex-
tremely variable both in size and structure, and in this
respect resemble the horns or the head and thorax
of many male Lamellicorns and Staphylinidse. A per-
fect series can be formed from the best-provided to the
worst-provided or degenerate males. Although the
mandibles of the common stag-beetle, and probably of

65 Quoted from Fischer, in ' Diet. Class. d'Hist. Nat.' torn. s. p. 324.

66 'Ann. Soe. Entomolog. France,' 1866, as quoted in 'Journal of
Travel,' by A. Murray, 186S, p. 135.

Chap. X.

COLEOPTERA.

377

many other species, are used as efficient weapons for
fighting, it is doubtful whether their great size can
thus be accounted for. We have
seen that with the Lueanus ela-
plms of N. America they are used
for seizing the female, As they
are so conspicuous and so ele-
gantly branched, the suspicion
has sometimes crossed my mind
that they may be serviceable to
the males as an ornament, in the
same manner as the horns on the
head and thorax of the various
above described species. The
male Chiasoc/nathus grantii of S.
Chile — a splendid beetle belong-
ing to the same family — has enor-
mously-developed mandibles (fig.
23) ; he is bold and pugnacious ;
when threatened on any side he
faces round, opening his great
jaws, and at the same time stridu-
lating loudly ; but the mandibles
were not strong enough to pinch
my finger so as to cause actual
pain.

Sexual selection, which implies
the possession of considerable per-
ceptive powers and of strong pas-
sions, seems to have been more
effective with the Lamellicorns
than with any other family of the
Ooleoptera or beetles. With some
species the males are provided with weapons for fight-
ing ; some live in pairs and show mutual affection ;

Fig. 23. Chiasopnathus grantii,
reduced. Upper figure, male ;
lower figure, female.

378 SEXUAL SELECTION. Part II.

many have the power of stridulating when excited ; many
are furnished with the most extraordinary horns, appa-
rently for the sake of ornament ; some which are diurnal
in their habits are gorgeously coloured ; and, lastly,
several of the largest beetles in the world belong to this
family, which was placed by Linnaeus and Fabricius at
the head of the Order of the Coleoptera.67

Stridulating organs. — Beetles belonging to many
and widely distinct families possess these organs. The
sound can sometimes be heard at the distance of several
feet or even yards,68 but is not comparable with that
produced by the Orthoptera. The part which may be
called the rasp generally consists of a narrow slightly-
raised surface, crossed by very fine, parallel ribs, some-
times so fine as to cause iridescent colours, and having
a very elegant appearance under the microscope. In
some cases, for instance, with Typhoeus, it could be
plainly seen that extremely minute, bristly, scale-like
prominences, which cover the whole surrounding sur-
face in approximately parallel lines, give rise to the
ribs of the rasp by becoming confluent and straight, and
at the same time more prominent and smooth. A hard
ridge on any adjoining part of the body, which in some
cases is specially modified for the purpose, serves as the
scraper for the rasp. The scraper is rapidly moved across
the rasp, or conversely the rasp across the scraper.

These organs are situated in widely different posi-
tions. In the carrion-beetles (Necrophorus) two parallel
rasps (r, fig. 24) stand on the dorsal surface of the fifth
abdominal segment, each rasp being crossed, as described
by Landois,69 by from 126 to 140 fine ribs. These

6' Westwood, ' Modern Class.' vol. i. p. 184.

68 Wollaston, On certain musical Curculioiiidse, 'Annals and Mag. of
Nat. Hist.' vol. vi. 1860, p. 14.

,;,J ' Zeitschrift fur wiss~ Zoolog.' B. xvii. 1867, s. 127.

Chap. X.

COLEOPTERA.

9

ribs are scraped by the posterior margins of the elytra, a
small portion of which projects beyond the general out-
line. In many Crioceridse, and in Chjthra ^-punctata

Fig. 21. Necrophorus (from Landois). r. The two rasps.

the rasp highly magnified.

Left-hand figure, part of

(one of the Chrysoinelidoe), and in some Tenebrionidae,
&c.,70 the rasp is seated on the dorsal apex of the abdo-
men, on the pygidium or pro-pygidium, and is scraped
as above by the elytra. In Heterocerus, which belongs
to another family, the rasps are placed on the sides of
the first abdominal segment, and are scraped by ridges
on the femora.71 In certain Curculionidae and Cara-
bidae,7^ the parts are completely reversed in position,

70 I am greatly indebted to Mr. G. E. Crotch for having sent me
numerous prepared specimens of various beetles belonging to these three
families and others, as well as for valuable information of all kinds. He
believes that the power of stridulation in the Clythra has not been pre-
viously observed. I am also much indebted to Mr. E. W. Janson, for
information and specimens. I may add that my son, Mr. F. Darwin,
finds that Dermestes murium stridulates, but he searched in vain for the
apparatus. Scolytus has lately been described by Dr. Chapman as a
stridulator, in the 'Entomologist's Monthly Magazine,' vol. vi. p. 130.

71 Schiodte, translated in ' Annals and Mag. of Nat. Hist.' vol. xx.
1867, p. 37.

72 Westring has described (Kroyer, ' Naturhist. Tidskrift,' B. ii. 184S-
49, p. 334) the stridulating organs in these two, as well as in other
families. In the Carabidse I have examined EldpTirus uliginosus and
Blethisa multipunctata, sent to me by Mr. Crotch. In Blethisa the
transverse ridges on the farrowed border of the abdominal segment do
not come into play, as far as I could judge, in scraping the rasps on the
elytra.

380

SEXUAL SELECTION.

Part II.

for the rasps are seated "on the inferior surface of the
elytra, near their apices, or along their outer margins,
and the edges of the abdominal segments serve as the
scrapers. In Pelobius hermanni (one of Dytiscidse or
water-beetles) a strong ridge runs parallel and near to
the sutural margin of the elytra, and is crossed by ribs,
coarse in the middle part, but becoming gradually finer
at both ends, especially at the upper end ; when this
insect is held under water or in the air, a stridulating
noise is produced by scraping the extreme horny margin
of the abdomen against the rasp. In a great number
of long-horned beetles (Longicornia) the organs are alto-
gether differently situated, the rasp being on the meso-
thorax, which is rubbed against the pro-thorax ; Landois
counted 238 very fine ribs on the rasp of Ceramlyx
hews.

Many Lamellicorns have the power of stridulating,
and the organs differ greatly in position. Some species
r stridulate very loudly, so that when

Mr. F. Smith caught a Trox sabu-
losus, a gamekeeper who stood by
thought that he had caught a
mouse ; but I failed to discover the
proper organs in this beetle. In
Geotrupes and Typhaeus a narrow
ridge runs obliquely across (r, fig.
25) the coxa of each hind-leg,
having in Gr. stercorarius 84 ribs,
which are scraped by a specially-
projecting part of one of the abdo-

Fig. 25. Hind-leg of Geotrupes . , T , ,

stercorarius (from Landois). niinal segments. In the nearly

allied Copris lunar is, an excessively
narrow fine rasp runs along the
sutural margin of the elytra, with another short rasp
near the basal outer margin ; but in some other Coprini

8

Easp. c Coxa, /. Femur.
t. Tibia, tr. Tarsi.

Chap. X. COLEOPTERA. 381

the rasp is seated, according to Leconte,73 on the dorsal
surface of the abdomen. In Oryctes it is seated on the
pro-pygidium, and in some other Dynastini, according to
the same entomologist, on the under surface of the
elytra. Lastly, Westring states that in Omctloplia hrun-
nea the rasp is placed on the pro-sternum, and the scraper
on the meta-sternum, the parts thus occupying the under
surface of the body, instead of the upper surface as in
the Lonsncorns.

We thus see that the stridulatinsr organs in the dif-
ferent coleopterous families are wonderfully diversified
in position, but not much in structure. Within the
same family some species are provided with these
organs, and some are quite destitute of them. This
diversity is intelligible, if we suppose that originally
various species made a shuffling or hissing noise by the
rubbing together of the hard and rough parts of their
bodies which were in contact ; and that from the noise
thus produced being in some way useful, the rough
surfaces were gradually developed into regular stri-
dulating organs. Some beetles as they move, now
produce, either intentionally or unintentionally, a shuf-
fling noise, without possessing any proper organs for the
purpose. Mr. Wallace informs me that the Euchirus
longimanus (a Lamellicorn, with the anterior legs won-
derfully elongated in the male) " makes, whilst moving,
" a low hissing sound by the protrusion and contraction
" of the abdomen ; and when seized it produces a
" grating sound by rubbing its hind-legs against the
" edges of the elytra." The hissing sound is clearly
due to a narrow rasp running along "the sutural margin
of each elytron ; and I could likewise make the grating

73 I am indebted to Mr. Walsh, of Illinois, for having sent me
extracts from Leconte's 'Introduction to Entomology,' p. 101, 143.

382 SEXUAL SELECTION. Part II.

sound bv rubbing the sliasrreened surface of the femur
against the granulated margin of the corresponding
elytron ; but I could not here detect any proper rasp ;
nor is it likely that I could have overlooked it in so
large an insect. After examining Cychrus and reading
what Westring has written in his two papers about this
beetle, it seems very doubtful whether it possesses any
true rasp, though it has the power of emitting a sound.

From the analogy of the Orthoptera and Homoptera,
I expected to find that the stridulating organs in the
Coleoptera differed according to sex ; but Landois, who
has carefully examined several species, observed no
such difference ; nor did Westring ; nor did Mr. G-. R.
Crotch in preparing the numerous specimens which
he had the kindness to send me for examination. Any
slight sexual difference, however, would be difficult to
detect, on account of the great variability of these organs.
Thus in the first pair of the Necrojohorus humator and of
the Pelobius which I examined, the rasp was consider-
ably larger in the male than in the female ; but not so
with succeeding specimens. In Geotrujoes stercorarius
the rasp appeared to me thicker, opaquer, and more
prominent in three males than in the same number of
females ; consequently my son, Mr. F. Darwin, in order
to discover whether the sexes differed in their power of
stridulating, collected 57 liviug specimens, which he
separated into two lots, according as they made, when
held in the same manner, a greater or lesser noise. He
then examined their sexes, but found that the males
were very nearly in the same proportion to the females
in both lots. Mr. F. Smith has kept alive numerous
specimens of Mononychus pseudacori (Curculionidae), and
is satisfied that both sexes stridulate, and apparently in
an equal degree.

Nevertheless the power of stridulating is certainly a

Chap. X. COLEOPTERA. 383

sexual character in some few Coleoptera. Mr. Crotch
has discovered that the males alone of two species of
Heliopathes (Tenebrionidse) possess striclulating organs.
I examined five males of H. gibhus, and in all these
there was a well-developed rasp, partially divided into
two, on the dorsal surface of the terminal abdominal
segment ; whilst in the same number of females there
was not even a rudiment of the rasp, the membrane of
this segment being transparent and much thinner than
in the male. In H. cribratostriatus the male has a
similar rasp, excepting that it is not partially divided
into two portions, and the female is completely desti-
tute of this organ ; but in addition the male has on
the apical margins of the elytra, on each side of the
suture, three or four short longitudinal ridges, which
are crossed by extremely fine ribs, parallel to and re-
sembling those on the abdominal rasp ; whether these
ridges serve as an independent rasp, or as a scraper
for the abdominal rasp, I could not decide : the female
exhibits no trace of this latter structure.

Again, in three species of the Lamellicorn genus
Oryctes, we have a nearly parallel case. In the females
of 0. gryplius and nasicornis the ribs on the rasp of the
pro-pygidium are less continuous and less distinct than
in the males ; but the chief difference is that the whole
upper surface of this segment, when held in the proper
light, is seen to be clothed with hairs, which are absent
or are represented by excessively fine down in the males.
It should be noticed that in all Coleoptera the effective
part of the rasp is destitute of hairs. In 0. senegal-
ensis the difference between the sexes is more strongly
marked, and this is best seen when the proper segment
is cleaned and viewed as a transparent object. In the
female the whole surface is covered with little separate
crests, bearing spines ; whilst in the male these crests

384 SEXUAL SELECTION. Part II.

become, in proceeding towards the apex, more and more
confluent, regular, and naked ; so that three-fourths of
the segment is covered with extremely fine parallel
ribs, which are quite absent in the female. In the
females, however, of all three species of Oryctes, when
the abdomen of a softened specimen is pushed back-
wards and forwards, a slight grating or stridulating
sound can be produced.

In the case of the Heliopathes and Oryctes there can
hardly be a doubt that the males stridulate in order to
call or to excite the females ; but with most beetles the
stridulation apparently serves both sexes as a mutual
call. This view is not rendered improbable from beetles
stridulating under various emotions ; we know that birds
use their voices for many purposes besides singing to
their mates. The great Chiasognathus stridulates in
anger or defiance ; many species do the same from dis-
tress or fear, when held so that they cannot escape ;
Messrs. Wollaston and Crotch were able, bv striking
the hollow stems of trees in the Canary Islands, to dis-
cover the presence of beetles belonging to the genus
Acalles by their stridulation. Lastly the male Ateu-
chus stridulates to encourage the female in her work,
and from distress when she is removed.74 Some natu-
ralists believe that beetles make this noise to frighten
away their enemies ; but I cannot think that the quadru-
peds and birds which are able to devour the larger
beetles with their extremely hard coats, would be fright-
ened by so slight a grating sound. The belief that
the stridulation serves as a sexual call is supported
by the fact that death-ticks (Anobium tessellation) are
well known to answer each other's ticking, or, as I have

7* M. P. de la Brulerie, as quoted in 'Journal of Travel,' A. Murray,
vol. i. 1868, p. 135.

Chap. X. COLEOPTEEA. 385

myself observed, a tapping noise artificially made ; and
Mr. Doubelday informs me that lie has twice or thrice
observed a female ticking,75 and in the course of an hour
or two has found her united with a male, and on one
occasion surrounded by several males. Finally, it seems
probable that the two sexes of many kinds of beetles
were at first enabled to find each other by the slight
shuffling noise produced by the rubbing together of the
adjoining parts of their hard bodies ; and that as the
males or females which made the greatest noise suc-
ceeded best in finding partners, the rugosities on various
parts of their bodies were gradually developed by means
of sexual selection into true stridulating organs.

"5 Mr. Doubleday informs me that " the noise is produced by the
" insect raising itself on its legs as high as it can, and then striking its
" thorax five or six times, in rapid succession, against the substance
" upon which it is sitting." For references on this subject see Landois,
' Zeitschrift fiir wissen. Zoolog.' B. xvii. s. 131. Olivier says (as quoted
by Kirby and Spence, 'Introduct.' vol. ii. p. 395) that the female of
Pimelia striata produces a rather loud sound by striking her abdomen
against any hard substance, "and that the male, obedient to this call,

soon attends her and they pair."

..

VOL. I. 2 c

386 SEXUAL SELECTION. Part II.

CHAPTEK XL

Insects, continued. — Order Lepidoptera.

Courtship of "butterflies — Battles — Ticking noise — Colours com-
mon 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 — Per-
ceptive powers of the Lepidoptera — Variability — Causes of the
difference in colour between the males and females — Mimickry,
female butterflies more brilliantly coloured than the males —
Bright colours of caterpillars — Summary and concluding re-
marks on the secondary sexual characters of insects — Birds and
insects compared.

In this great Order the most interesting point for us is
the difference in colour between the sexes of the same
species, and between the distinct species of the same
genus. Nearly the whole of the following chapter will
be devoted to this subject ; but I will first make a few
remarks on one or two other points. Several males may
often be seen pursuing and crowding round the same
female. Their courtship appears to be a prolonged affair,
for I have frequently watched one or more males pirouet-
ting round a female until I became tired, without seeing
the end of the courtship. Although butterflies are such
weak and fragile creatures, they are pugnacious, and an
Emperor butterfly x has been captured with the tips of
its wings broken from a conflict with another male.
Mr. Collingwood in speaking of the frequent battles

1 Apatura Iris : ' The Entomologist's "Weekly Intelligencer,' 1859,
p. 139. For the Bornean Butterflies see C. Collingwood, 'Bamblea of
a Naturalist,' 1868, p. 183.

Chap. XL BUTTERFLIES AND MOTHS. 387

between the butterflies of Borneo says, "They whirl
" round each other with the greatest rapidity, and appear
" to be incited by the greatest ferocity." One case is
known of a butterfly, namely the Ageronia feronia,
which makes a noise like that produced by a toothed
wheel passing under a spring catch, and which could be
heard at the distance of several yards. At Kio de Janeiro
this sound was noticed by me, only when two were
chasing each other in an irregular course, so that it is
probably made during the courtship of the sexes ; but I
neglected to attend to this point.2

Every one has admired the extreme beauty of many
butterflies and of some moths ; and we are led to ask,
how has this beauty been acquired ? Have their colours
and diversified patterns simply resulted from the direct
action of the physical conditions to which these insects
have been exposed, without any benefit being thus de-
rived ? Or have successive variations been accumulated
and determined either as a protection or for some un-
known purpose, or that one sex might be rendered
attractive to the other ? And, again, what is the mean-
ing of the colours being widely different in the males
and females of certain species, and alike in the two
sexes of other species? Before attempting to answer
these questions a body of facts must be given.

With most of our English butterflies, both those which
are beautiful, such as the admiral, peacock, and painted
lady (Vanessse), and those which are plain-coloured,
such as the meadow-browns (Hipparchiae), the sexes
are alike. This is also the case with the magnificent
Heliconidse and Danaidse of the tropics. But in certain

2 See my ' Journal of Eesearches,' 1845, p. 33. Mr. Doubleday has
detected (' Proc. Ent. Soc.' March 3rd, 1845, p. 123) a peculiar mem-
branous sac at the base of the front wings, which is probably con-
nected with the production of the sound.

2 c 2

388 SEXUAL SELECTION. Part II.

other tropical groups, and with some of our English
butterflies, as the purple emperor, orange-tip, &c. (Apa-
tura Iris and Anthocharis cardamines), the sexes differ
either greatly or slightly in colour. No language suffices
to describe the splendour of the males of some tropical
species. Even within the same genus we often find spe-
cies presenting an extraordinary difference between the
sexes, whilst others have their sexes closely alike. Thus
in the South American genus Epical ia, Mr. Bates, to
whom I am much indebted for most of the following
facts and for looking over this whole discussion, informs
me that he knows twelve species, the two sexes of which
haunt the same stations (and this is not always the case
with butterflies), and therefore cannot have been dif-
ferently affected by external conditions.3 In nine of
these species the males rank amongst the most brilliant
of all butterflies, and differ so greatly from the compa-
ratively plain females that they were formerly placed
in distinct genera. The females of these nine species
resemble each other in their general type of coloration,
and likewise resemble both sexes in several allied genera,
found in various parts of the world. Hence in accord-
ance with the descent-theory we may infer that these
nine species, and probably all the others of the genus,
are descended from an ancestral form which was coloured
in nearly the same manner. In the tenth species the
female still retains the same general colouring, but the
male resembles her, so that he is coloured in a much
less gaudy and contrasted manner than the males of the
previous species. In the eleventh and twelfth species,
the females depart from the type of colouring which

3 See also Mr. Bates' paper in ' Proc. Ent. Soc. of Philadelphia,'
1865, p. 206. Also Mr. Wallace on the same subject, in regard to
Diadema, in ' Transact. Entomolog. Soc. of London,' I860, p. 278.

Chap. XL BUTTERFLIES AND MOTHS. 389

is usual with their sex in this genus, for they are gaily
decorated in nearly the same manner as the males, but
in a somewhat less degree. Hence in these two species
the bright colours of the males seem to have been trans-
ferred to the females ; whilst the male of the tenth
species has either retained or recovered the plain
colours of the female as well as of the parent-form of
the genus ; the two sexes being thus rendered in both
cases, though in an opposite manner, nearly alike. In
the allied genus Eubagis, both sexes of some of the
species are plain-coloured and nearly alike ; whilst
with the greater number the males are decorated with
beautiful metallic tints, in a diversified manner, and
differ much from their females. The females through-
out the genus retain the same general style of colouring,
so that they commonly resemble each other much more
closely than they resemble their own proper males.

In the genus Papilio, all the species of the iEneas
group are remarkable for their conspicuous and strongly
contrasted colours, and they illustrate the frequent ten-
dency to gradation in the amount of difference between
the sexes. In a few species, for instance in P. ascanius,
the males and females are alike ; in others the males
are a little or very much more superbly coloured than
the females. The genus Junonia allied to our Vanessaa
offers a nearly parallel case, for although the sexes of
most of the species resemble each other and are desti-
tute of rich colours, yet in certain species, as in J. oenone,
the male is rather more brightly coloured than the
female, and in a few (for instance J. andremiaja) the
male is so different from the female that he might be
mistaken for an entirely distinct species.

Another striking case was pointed out to me in the
British museum by Mr. A. Butler, namely one of
the Tropical American Theclae, in which both sexes

390 SEXUAL SELECTION. Part II.

are nearly alike and wonderfully splendid ; in another,
the male is coloured in a similarly gorgeous manner,
whilst the whole upper surface of the female is of a dull
uniform brown. Our common little English blue butter-
flies of the genus Lycsena, illustrate the various differ-
ences in colour between the sexes, almost as well,
though not in so striking a manner, as the above exotic
genera. In Lycsena agestis both sexes have wings of a
brown colour, bordered with small ocellated orange
spots, and are consequently alike. In L. oegon the
wings of the male are of a fine blue, bordered with
black ; whilst the wings of the female are brown, with
a similar border, and closely resemble those of L. agestis.
Lastly, in L. avion both sexes are of a blue colour and
nearly alike, though in the female the edges of the
wings are rather duskier, w7ith the black spots plainer ;
and in a bright blue Indian species both sexes are still
more closely alike.

I have given the foregoing cases in some detail in
order to shew, in the first place, that when the sexes of
butterflies differ, the male as a general rule is the most
beautiful, and departs most from the usual type of colour-
ing of the group to which the species belongs. Hence in
most groups the females of the several species resemble
each other much more closely than do the males. In
some exceptional cases, however, to which I shall here-
after allude, the females are coloured more splendidly
than the males. In the second place these cases have
been given to bring clearly before the mind that within
the same genus, the two sexes frequently present every
gradation from no difference in colour to so great a dif-
ference that it was long before the two were placed by
entomologists in the same genus. In the third place,
we have seen that when the sexes nearly resemble
each other, this apparently may be due either to the

Chap. XI. BUTTERFLIES AND MOTHS. 391

male having transferred his colours to the female,
or to the male having retained, or perhaps reco-
vered, the primordial colours of the genus to which the
species belongs. It also deserves notice that in those
groups in which the sexes present any difference of
colour, the females usually resemble the males to a cer-
tain extent, so that when the males are beautiful to an
extraordinary degree, the females almost invariably ex-
hibit some degree of beauty. From the numerous cases
of gradation in the amount of difference between the
sexes, and from the prevalence of the same general type
of coloration throughout the whole of the same group,
we may conclude that the causes, whatever they may
be, which have determined the brilliant colouring of the
males alone of some species, and of both sexes in a more
or less equal degree of other species, have generally
been the same.

As so many gorgeous butterflies inhabit the tropics, it
has often been supposed that they owe their colours to
the great heat and moisture of these zones ; but Mr.
Bates 4 has shewn by the comparison of various closely-
allied groups of insects from the temperate and tropical
regions, that this view cannot be maintained; and the
evidence becomes conclusive when brilliantly-coloured
males and plain-coloured females of the same species
inhabit the same district, feed on the same food, and
follow exactly the same habits of life. Even when
the sexes resemble each other, we can hardly believe
that their brilliant and beautifully-arranged colours are
the purposeless result of the nature of the tissues, and
the action of the surrounding conditions.

With animals of all kinds, whenever colour has been
modified for some special purpose, this has been, as far

4 ' The Naturalist on the Amazons,' vol. i. 1863, p. 19.

392 SEXUAL SELECTION. Part II.

as we can judge, either for protection or as an attraction
between the sexes. With many species of butterflies
the upper surfaces of the wings are obscurely coloured,
and this in all probability leads to their escaping ob-
servation and danger. But butterflies when at rest
would be particularly liable to be attacked by their
enemies; and almost all the kinds when resting raise
their wings vertically over their backs, so that the lower
sides alone are exposed to view. Hence it is this side
which in many cases is obviously coloured so as to
imitate the surfaces on which these insects commonly
rest. Dr. Bossier, I believe, first noticed the similarity
of the closed wings of certain Yanessaa and other butter-
flies to the bark of trees. Many analogous and striking
facts could be given. The most interesting one is that
recorded by Mr. Wallace5 of a common Indian and
Sumatran butterfly (Kallima), which disappears like
magic when it settles in a bush ; for it hides its head
and antennae between its closed wings, and these in
form, colour, and veining cannot be distinguished from
a withered leaf together with the footstalk. In some
other cases the lower surfaces of the wings are brilliantly
coloured, and yet are protective ; thus in Theda rubi
the wings when closed are of an emerald green and re-
semble the young leaves of the bramble, on which this
butterfly in the spring may often be seen seated.

Although the obscure tints of the upper or under
surface of many butterflies no doubt serve to conceal
them, yet we cannot possibly extend this view to
the brilliant and conspicuous colours of many kinds,
such as our admiral and peacock Vanessae, our white

5 See the interesting article in the * Westminster Eeview,' July, 1867,
p. 10. A woodcut of the Kallima is given by Mr. Wallace in ' Hard-
wicke's ' Science Gossip,' Sept. 1867, p. 196.

Chap. XI. BUTTERFLIES AND MOTHS. 393

cabbage-butterflies (Pieris), or the great swallow-tail
Papilio which haunts the open fens — for these butter-
flies are thus rendered visible to every living creature.
With these species both sexes are alike ; but in the
common brimstone butterfly (Gonepteryx rhamni), the
male is of an intense yellow, whilst the female is much
paler; and in the orange-tip (Anthoeharis cardamines)
the males alone have the bright orange tips to their
wings. In these cases the males and females are
equally conspicuous, and it is not credible that their
difference in colour stands in any relation to ordinary
protection. Nevertheless it is possible that the con-
spicuous colours of many species may be in an indirect
manner beneficial, as will hereafter be explained, by
leading their enemies at once to recognise them as
unpalatable. Even in this case it does not certainly
follow that their bright colours and beautiful patterns
were acquired for this special purpose. In some other
remarkable cases, beauty has been gained for the sake
of protection, through the imitation of other beautiful
species, which inhabit the same district and enjoy an
immunity from attack by being in some way offensive
to their enemies.

The female of our orange-tip buttterny, above re-
ferred to, and of an American species (Anth. genutia)
probably shew us, as Mr. Walsh has remarked to me,
the primordial colours of the parent - species of the
genus ; for both sexes of four or five widely-distributed
species are coloured in nearly the same manner. We
may infer here, as in several previous cases, that it is
the males of Anth. cardamines and genutia which have
departed from the usual type of colouring of their genus.
In the Anth. sara from California, the orange-tips have
become partially developed in the female ; for her wings
are tipped with reddish-orange, but paler than in the

394 • SEXUAL SELECTION. Part II.

male, and slightly different in some other respects. In
an allied Indian form, the Ijjhias glauci^e, the orange-
tips are fully developed in both sexes. In this Iphias
the under surface of the wings marvellously resembles,
as pointed out to me by Mr. A. Butler, a pale-coloured
leaf; and in our English orange-tip, the under surface
resembles the flower-head of the wild parsley, on which
it may be seen going to rest at night.6 The same
reasoning power which compels us to believe that the
lower surfaces have here been coloured for the sake of
protection, leads us to deny that the wings have been
tipped, especially when this character is confined to the
males, with bright orange for the same purpose.

Turning now to Moths : most of these rest motion-
less with their wings depressed during the whole or
greater part of the day ; and the upper surfaces of their
wings are often shaded and coloured in an admirable
manner, as Mr. Wallace has remarked, for escaping
detection. With most of the • Boinbycidse and Noc-
tuidse,7 when at rest, the front-wings overlap and
conceal the hind-wings; so that the latter might be
brightly coloured without much risk ; and they
are thus coloured in many species of both families.
During the act of flight, moths wrould often be able
to escape from their enemies ; nevertheless, as the
hind-wings are then fully exposed to view, their bright
colours must generally have been acquired at the
cost of some little risk. But the following fact shews
us how cautious we ought to be in drawing conclu-
sions on this head. The common yellow under-wings

6 See the interesting observations by Mr. T. "W. Wood, ' The Stu-
dent,' Sept. 1868, p. 81.

7 Mr. Wallace in ' Hardwicke's Science Gossip,' Sept. 1S67, p. 193.

Chap. XI. BUTTERFLIES AND MOTHS. 395

(Triphaena) often fly about during the day or early
evening, and are then conspicuous from the colour of
their hind-wings. It would naturally be thought that
this would be a source of danger ; but Mr. J. Jenner
Weir believes that it actually serves them as a means
of escape, for birds strike at these brightly coloured and
fragile surfaces, instead of at the body. For instance,
Mr. Weir turned into his aviary a vigorous specimen of
Triphaena pronuba, which was instantly pursued by
a robin ; but the bird's attention being caught by the
coloured wings, the moth was not captured until after
about fifty attempts, and small portions of the wings
were repeatedly broken off. He tried the same experi-
ment, in the open air, with a T. fimbria and swallow;
but the large size of this moth probably interfered
with its capture.8 We are thus reminded of a state-
ment made by Mr. Wallace,9 namely, that in the Bra-
zilian forests and Malayan islands, many common and
highly-decorated butterflies are weak flyers, though fur-
nished with a broad expanse of wings ; and they " are
"often captured with pierced and broken wings, as if
'• they had been seized by birds, from which they had
" escaped : if the wings had been much smaller in pro-
" portion to the body, it seems probable that the insect
" would more frequently have been struck or pierced in
"a vital part, and thus the increased expanse of the
" wings may have been indirectly beneficial."

Display. — The bright colours of butterflies and of
some moths are specially arranged^for display, whether
or not they serve in addition as a protection. Bright

8 See also, on this subject, Mr. Weir's paper in ' Transact. Ent. Soc'
1869, p. 23. A/ O"os Hl

9 ' Westminster Review,' July, 1867, p. 16.

lLjiLIBR f

396 SEXUAL SELECTION. Part II.

colours would not be visible during: the nig^ht: and
there can be no doubt that moths, taken as a body, are
much less gaily decorated than butterflies, all of which
are diurnal in their habits. But the moths in certain
families, such as the Zygaenidae, various Sphingidaa,
Uraniidae, some Arctiidse and Saturniidaa, fly about
during the clay or early evening, and many of these
are extremely beautiful, being far more brightly
coloured than the strictly nocturnal kinds. A few
exceptional cases, however, of brightly-coloured noc-
turnal species have been recorded.10

There is evidence of another kind in regard to display.
Butterflies, as before remarked, elevate their wings
when at rest, and whilst basking in the sunshine often
alternately raise and depress them, thus exposing to full
view both surfaces; and although the lower surface is
often coloured in an obscure manner as a protection,
yet in many species it is as highly coloured as the
upper surface, and sometimes in a very different man-
ner. In some tropical species the lower surface is even
more brilliantly coloured than the upper.11 In one
English fritillary, the Argijnnis aglaia, the lower sur-
face alone is ornamented with shining silver discs.
Nevertheless, as a general rule, the upper surface,
which is probably the most fully exposed, is coloured
more brightly and in a more diversified manner than
the lower. Hence the lower surface generally affords

10 For instance, Lithosia ; but Prof. Westwood (' Modern Class, of
Insects,' vol. ii. p. 390) seems surprised at this case. On the relative
colours of diurnal and nocturnal Lepidoptera, see ibid. p. H33 and 392 ;
also Harris, ' Treatise on the Insects of New England,' 1842, p. 315.

11 Such differences between the upper and lower surfaces of the
wings of several species of Papilio, may be seen in the beautiful plates
to Mr. Wallace's Memoir on the Papilionidse of the Malayan liegiou,
in ' Transact. Linn. Soc' vol. xxv. part i. 1865.

Chap. XI. BUTTERFLIES AND MOTHS. 397

to entomologists the most useful character for detecting
the affinities of the various species.

Now if we turn to the enormous group of moths,
which do not habitually expose to full view the under
surface of their wings, this side is very rarely, as I hear
from Mr. Stainton, coloured more brightly than the
upper side, or even with equal brightness. Some ex-
ceptions to the rule, either real or apparent, must be
noticed, as that of Hypopyra, specified by Mr. Wormald.12
Mr. E. Trimen informs me that in Guenee's great work,
three moths are figured, in which the under surface is
much the most brilliant. For instance, in the Australian
Gastrophora the upper surface of the fore-wing is pale
greyish-ochreous, while the lower surface is magnificently
ornamented by an ocellus of cobalt-blue, placed in the
midst of a black mark, surrounded by orange-yellow,
and this by bluish-white. But the habits of these
three moths are unknown ; so that no explanation
can be given of their unusual style of colouring. Mr.
Trimen also informs me that the lower surface of the
wings in certain other Geometrse 13 and quadrifid Noctuae
are either more variegated or more brightly-coloured
than the upper surface ; but some of these species have
the habit of " holding their wings quite erect over their
" backs, retaining them in this position for a considerable
" time," and thus exposing to view the under surface.
Other species when settled on the ground or herbage
have the habit of now and then suddenly and slightly
lifting up their wings. Hence the lower surface of the
wings being more brightly-coloured- than the upper sur-

12 ' Proc. Eut. Soc' March 2nd, 1868.

13 See also an account of the S. American genus Erateina (one of
the Geometrse) in ' Transact. Ent. Soc.' new series, vol. v. pi. xv. and
xvi.

398 SEXUAL SELECTION. Part II.

face in certain moths is not so anomalous a circum-
stance as it at first appears. The Saturniidse include
some of the most beautiful of all moths, their wings
being decorated, as in our British Emperor moth, with
fine ocelli ; and Mr. T. W. Wood w observes that they
resemble butterflies in some of their movements ; " for
" instance, in the gentle waving up and down of the
" wings, as if for display, which is more characteristic
" of diurnal than of nocturnal Lepidoptera."

It is a singular fact that no British moths, nor as
far as I can discover hardly any foreign species, which
are brilliantly coloured, differ much in colour according
to sex ; though this is the case with many brilliant but-
terflies. The male, however, of one American moth, the
Satumia Io, is described as having its fore-wings deep
yellow7, curiously marked with purplish-red spots ; whilst
the wings of the female are purple-brown, marked with
grey lines,15 The British moths which differ sexually in
colour are all brown, or various tints of dull yellow, or
nearly white. In several species the males are much
darker than the females,16 and these belong to groups
which generally fly about during the afternoon. On the
other hand, in many genera, as Mr. Stainton informs me,

14 ' Proc. Ent. Soc. of London,' July 6, 1868, p. xxvii.

15 Harris, ' Treatise/ &c, edited by Flint, 1862, p. 395.

16 For instance, I observe in my son's cabinet that the males are
darker than the females in the Lasiocampa quercus, Odonestis potatoria,
Hypogymna dispar, Dasycliira pudibunda, and Cycnia mendlca. In this
latter species the difference in colour between the two sexes is strongly
marked ; and Mr. Wallace informs me that we here have, as he believes,
an instance of protective mimickry confined to one sex, as will hereafter
be more fully explained. The white female of the Cycnia resembles
the very common Spilosoma menthrasti, both sexes of which are white ;
and Mr. Stainton observed that this latter moth was rejected with utter
disgust by a whole brood of young turkeys, which were fond of eating
other moths ; so that if the Cycnia was commonly mistaken by British
birds for the Spilosoma, it would escape being devoured, and its white
deceptive colour would thus be highly beneficial.

Chap. XL BUTTERFLIES AND MOTHS. 399

the males have the hind-wings whiter than those of
the female — of which fact Agrotis exclamationis offers a
good instance. The males are thus rendered more
conspicuous than the females, whilst flying about in
the dusk. In the Ghost Moth (Hejnalus humuli) the
difference is more strongly marked; the males being
white and the females yellow with darker markings.
It is difficult to conjecture what the meaning can be
of these differences between the sexes in the shades of
darkness or lightness ; but we can hardly suppose that
they are the result of mere variability with sexually-
limited inheritance, independently of any benefit thus
derived

From the foregoing statements it is impossible to
admit that the brilliant colours of butterflies and of
some few moths, have commonly been acquired for the
sake of protection. We have seen that their colours
and elegant patterns are arranged and exhibited as
if for display. Hence I am led to suppose that the
females generally prefer, or are most excited by the
more brilliant males ; for on any other supposition
the males would be ornamented, as far as we can
see, for no purpose. We know that ants and certain
lamellicorn beetles are capable of feeling an attachment
for each other, and that ants recognise their fellows
after an interval of several months. Hence there is no
abstract improbability in the Lepidoptera, which pro-
bably stand nearly or quite as high in the scale as these
insects, having sufficient mental capacity to admire
bright colours. They certainly discover flowers by
colour, and, as I have elsewhere shewn, the plants
which are fertilised exclusively by the wind never have
a conspicuously-coloured corolla. The Humming-bird
Sphinx may often be seen to swoop down from a distance
on a bunch of flowers in the midst of green foliage }

400 SEXUAL SELECTION. Part II.

and I have been assured by a friend, that these moths
repeatedly visited flowers painted on the walls of a room
in the South of France. The common white butterfly,
as I hear from Mr. Doubleday, often flies down to a bit
of paper on the ground, no doubt mistaking it for one of
its own species. Mr. Collingwood 17 in speaking of the
difficulty of collecting certain butterflies in the Malay
Archipelago, states that " a dead specimen pinned upon
" a conspicuous twig will often arrest an insect of the
" same species in its headlong flight, and bring it down
" within easy reach of the net, especially if it be of the
" opposite sex."

The courtship of butterflies is a prolonged affair. The
males sometimes fight together in rivalry ; and many
may be seen pursuing or crowding round the same
female. If, then, the females do not prefer one male to
another, the pairing must be left to mere chance, and
this does not appear to me a probable event. If, on the
other hand, the females habitually, or even occasionally,
prefer the more beautiful males, the colours of the latter
will have been rendered brighter by degrees, and will
have been transmitted to both sexes or to one sex,
according to which law of inheritance prevailed. The
process of sexual selection will have been much facili-
tated, if the conclusions arrived at from various kinds of
evidence in the supplement to the ninth chapter can be
trusted ; namely that the males of many Lepidoptera,
at least in the imago state, greatly exceed in number
the females.

Some facts, however, are opposed to the belief that
female butterflies prefer the more beautiful males ; thus,
as I have been assured by several observers, fresh females
may frequently be seen paired with battered, faded or

17 ' Eambles of a Naturalist in the Chinese Seas,' 1868, p. 182.

Chap. XL BUTTERFLIES AND MOTHS. 401

dingy males ; but this is a circumstance which could
hardly fail often to follow from the males emerging
from their cocoons earlier than the females. With moths
of the family of the Bombyeiclaa, the sexes pair imme-
diately after assuming the imago state ; for they cannot
feed, owing to the rudimentary condition of their mouths.
The females, as several entomologists have remarked
to me, lie in an almost torpid state, and appear not to
evince the least choice in regard to their partners.
This is the case with the common silk-moth (B. mori),
as I have been told by some continental and English
breeders. Dr. Wallace, who has had such immense
experience in breeding Bomhtjx cynthia, is convinced
that the females evince no choice or preference. He
has kept above 300 of these moths living together, and
has often found the most vigorous females mated with
stunted males. The reverse apparently seldom occurs ;
for, as he believes, the more vigorous males pass over the
weakly females, being attracted by those endowed with
most vitality. Although we have been indirectly in-
duced to believe that the females of many species prefer
the more beautiful males, I have no reason to suspect,
either with moths or butterflies, that the males are
attracted by the beauty of the females. If the more
beautiful females had been continually preferred, it is
almost certain, from the colours of butterflies being so
frequently transmitted to one sex alone, that the females
would often have been rendered more beautiful than
their male partners. But this does not occur except in
a few instances ; and these can be explained, as we
shall presently see, on the principle of mimickry and
protection.

As sexual selection primarily depends on variability,
a few words must be added on this subject. In respect
vol. i. 2d

402 SEXUAL SELECTION. Part II.

to colour there is no difficulty, as any number of highly
variable Lepicloptera could be named. One good in-
stance will suffice. Mr. Bates shewed me a whole series
of specimens of Pcqoilio sesostris and children^ ; in the
latter the males varied much in the extent of the beau-
tifully enamelled green patch on the fore-wings, and
in the size of the white mark, as well as of the splendid
crimson stripe on the hind-wings ; so that there was
a great contrast between the most and least gaudy
males. The male of Pajnlio sesostris, though a beautiful
insect, is much less so than P. chilclrense. It likewise
varies a little in the size of the green patch on the fore-
wings, and in the occasional appearance of a small
crimson stripe on the hind- wings, borrowed, as it would
seem, from its own female ; for the females of this and
of many other species in the ^neas group possess this
crimson stripe. Hence between the brightest specimens
of P. sesostris and the least bright of P. children^, there
was but a small interval ; and it was evident that as far
as mere variability is concerned, there would be no
difficulty in permanently increasing by means of selec-
tion the beauty of either species. The variability is
here almost confined to the male sex ; but Mr. Wallace
and Mr. Bates have shewn 18 that the females of some
other species are extremely variable, the males being
nearly constant. As I have before mentioned the Ghost
Moth (Hepialus humuli) as one of the best instances in
Britain of a difference in colour between the sexes of
moths, it may be worth adding 19 that in the Shetland

18 Wallace on the Papilionidae of the Malayan Region, in ' Transact.
Linn. Soc.' vol. xxv. 1865, p. 8, 36. A striking case of a rare variety,
strictly intermediate between two other well-marked female varieties,
is given by Mr. Wallace. See also Mr. Bates, in ' Proc. Entomolog.
Soc.' Nov. 19th, 1866, p. xl.

19 Mr. R. MacLacblan, ' Transact. Ent. Soc' vol. ii. part 6th, 3rd
series, 1866, p. 459.

Chap. XL BUTTERFLIES AND MOTHS. 403

Islands, males are frequently found which closely
resemble the females. In a future chapter I shall have
occasion to shew that the beautiful eye-like spots or
ocelli, so common on the wings of many Lepidoptera,
are eminently variable.

On the whole, although many serious objections may
be urged, it seems probable that most of the species of
Lepidoptera which are brilliantly coloured, owe their
colours to sexual selection, excepting in certain cases,
presently to be mentioned, in which conspicuous colours
are beneficial as a protection. From the ardour of the
male throughout the animal kingdom, he is generally
willing to accept any female ; and it is the female which
usually exerts a choice. Hence if sexual selection has
here acted, the male, when the sexes differ, ought to be
the most brilliantly coloured ; and this undoubtedly is
the ordinary rule. When the sexes are brilliantly co-
loured and resemble each other, the characters acquired
by the males appear to have been transmitted to both
sexes. But will this explanation of the similarity and
dissimilarity in colour between the sexes suffice ?

The males and females of the same species of butterfly
are known 20 in several cases to inhabit different stations,
the former commonly basking in the sunshine, the latter
haunting gloomy forests. It is therefore possible that
different conditions of life may have acted directly on
the two sexes ; but this is not probable,21 as in the adult
state they are exposed during a very short period to
different conditions ; and the larvae" of both are ex-
posed to the same conditions. Mr. Wallace believes

20 H. W. Bates, 'The Naturalist on the Amazons/ vol. ii. 1863,
p. 228. A. E. Wallace, in ' Transact. Linn. Soc' vol. xxv. 1865, p. 10.

21 On this whole subject see ' The Variation of Animals and Plants
under Domestication,' vol. ii. 1868, chap, xxiii.

2 d 2

404 SEXUAL SELECTION. Part II.

that the less brilliant colours of the female have been
specially gained in all or almost all cases for the sake
of protection. On the contrary it seems to me more
probable that the males alone, in the large majority of
cases, have acquired their bright colours through sexual
selection, the females having been but little modified.
Consequently the females of distinct but allied species
ought to resemble each other much more closely than
do the males of the same species ; and this is the general
rule. The females thus approximately show us the pri-
mordial colouring of the parent-species of the group to
which they belong. They have, however, almost always
been modified to a certain extent by some of the succes-
sive steps of variation, through the accumulation of
which the males were rendered beautiful, having been
transferred to them. The males and females of allied
though distinct species will also generally have been
exposed during their prolonged larval state to different
conditions, and may have been thus indirectly affected ;
though with the males any slight change of colour thus
caused will often have been completely masked by the
brilliant tints gained through sexual selection. When
we treat of Birds, I shall have to discuss the whole
question whether the differences in colour between the
males and females have been in part specially gained
by the latter as a protection ; so that I will here only
give unavoidable details.

In all cases when the more common form of equal
inheritance by both sexes has prevailed, the selection of
bright-coloured males would tend to make the females
bright-coloured ; and the selection of dull-coloured fe-
males wmdd tend to make the males dull. If both pro-
cesses were carried on simultaneously, they would tend
to neutralise each other. As far as I can see, it would
be extremely difficult to change through selection the

Chap. XI. BUTTERFLIES AND MOTHS. 405

one form of inheritance into the other. Bat by the
selection of successive variations, which were from the
first sexually limited in their transmission, there would
not be the slightest difficulty in giving bright colours to
the males alone, and at the same time or subsequently,
dull colours to the females alone. In this latter manner
female butterflies and moths may, as I fully admit, have
been rendered inconspicuous for the sake of protection,
and widely different from their males.

Mr. Wallace 22 has argued with much force in favour
of his view that when the sexes differ, the female has
been specially modified for the sake of protection ; and
that this has been effected by one form of inheritance,
namely, the transmission of characters to both sexes,
having been changed through the agency of natural
selection into the other form, namely, transmission to
one sex. I was at first strongly inclined to accept this
view ; but the more I have studied the various classes
throughout the animal kingdom, the less probable it
has appeared. Mr. Wallace urges that both sexes of
the Heliconidae, Danaidse, Acrseidm are equally brilliant
because both are protected from the attacks of birds
and other enemies, by their offensive odour ; but that
in other groups, which do not possess this immunity,
the females have been rendered inconspicuous, from
having more need of protection than the males. This
supposed difference in the " need of protection by the
" two sexes ' is rather deceptive, and requires some
discussion. It is obvious that brightly-coloured indi-
viduals, whether males or females, would equally attract,
and obscurely-coloured individuals equally escape, the

22 A. E. Wallace, in 'The Journal of Travel,' vol. i. 1868, p. 88.
'Westminster Keview,' July, 1867, p. 37. See also Messrs. Wallace
and Bates in ' Proc. Ent. Soc' Nov. 19th, 1S66, p. xxxix.

406 SEXUAL SELECTION. Part II.

attention of their enemies. But we are concerned
with the effects of the destruction or preservation of
certain individuals of either sex, on the character of
the race. With insects, after the male has fertilised
the female, and after the latter has laid her eggs,
the greater or less immunity from danger of either sex
could not possibly have any effect on the offspring.
Before the sexes have performed their proper functions,
if they existed in equal numbers and if they strictly
paired (all other circumstances being the same), the
preservation of the males and females would be equally
important for the existence of the species and for the
character of the offspring. But with most animals, as
is known to be the case with the domestic silk-moth,
the male can fertilise two or three females ; so that the
destruction of the males would not be so injurious to
the species as that of the females. On the other hand,
Dr. Wallace believes that with moths the progeny from
a second or third fertilisation is apt to be weakly, and
therefore would not have so good chance of surviving.
When the males exist in much greater numbers than the
females, no doubt many males might be destroyed with
impunity to the species ; but I cannot see that the
results of ordinary selection for the sake of protection
would be influenced by the sexes existing in unequal
numbers ; for the same proportion of the more con-
spicuous individuals, whether males or females, would
probably be destroyed. If indeed the males presented
a greater range of variation in colour, the result would
be different ; but we need not here follow^ out such com-
plex details. On the whole I cannot perceive that an
inequality in the numbers of the two sexes would in-
fluence in any marked manner the effects of ordinary
selection on the character of the offspring.

Female Lepidoptera require, as Mr. Wallace insists,

Chap. XI. BUTTEEFLIES AND MOTHS. 407

some days to deposit tlieir fertilised ova and to search
for a proper place; during this period (whilst the life
of the male was of no importance) the brighter-coloured
females would be exposed to clanger and would be
liable to be destroyed. The duller-coloured females on
the other hand would survive, and thus would in-
fluence, it might be thought, in a marked manner the
character of the species, — either of both sexes or of
one sex, according to which form of inheritance pre-
vailed. But it must not be forgotten that the males
emerge from the cocoon-state some days before the
females, and during this period, whilst the unborn
females were safe, the brighter-coloured males would be
exposed to danger ; so that ultimately both sexes wrould
probably be exposed during a nearly equal length of
time to clanger, and the elimination of conspicuous
colours would not be much more effective in the one
than the other sex.

It is a more important consideration that female
Lepicloptera, as Mr. Wallace remarks, and as is known
to every collector, are generally slower flyers than
the males. Consequently the latter, if exposed to
greater danger from being conspicuously coloured,
might be able to escape from their enemies, whilst the
similarly-coloured females would be destroyed ; and thus
the females would have the most influence in modi-
fying the colour of their progeny.

There is one other consideration : bright colours, as
far as sexual selection is concerned, are commonly of
no service to the females ; so that if the latter varied
in brightness, and the variations were sexually limited
in their transmission, it would depend on mere chance
whether the females had their bright colours increased ;
and this would tend throughout the Order to diminish
the number of species with brightly-coloured females

408 SEXUAL SELECTION. Part II.

in comparison with the species having brightly-coloured
males. On the other hand, as bright colours are sup-
posed to be highly serviceable to the males in their
love-struggles, the brighter males (as we shall see
in the chapter on Birds) although exposed to rather
greater danger, would on an average procreate a greater
number of offspring than the duller males. In this
case, if the variations were limited in their transmission
to the male sex, the males alone would be rendered
more brilliantly coloured ; but if the variations were
not thus limited, the preservation and augmentation of
such variations would depend on whether more evil was
caused to the species by the females being rendered
conspicuous, than good to the males by certain indivi-
duals being successful over their rivals.

As there can hardly be a doubt that both sexes of
many butterflies and moths have been rendered dull-
coloured for the sake of protection, so it may have
been with the females alone of some species in which
successive variations towards dullness first appeared
in the female sex and were from the first limited in
their transmission to the same sex. If not thus limited,
both sexes would become dull-coloured. We shall
immediately see, when we treat of mimickry, that
the females alone of certain butterflies have been ren-
dered extremely beautiful for the sake of protection,
without any of the successive protective variations
having been transferred to the male, to whom they
could not possibly have been in the least degree injuri-
ous, and therefore could not have been eliminated
through natural selection. Whether in each particular
species, in which the sexes differ in colour, it is the
female which has been specially modified for the sake
of protection ; or whether it is the male which has been
specially modified for the sake of sexual attraction, the

Chap. XI. BUTTEEFLIES AND MOTHS. 409

female having retained her primordial colouring only
slightly changed through the agencies before alluded
to ; or whether again both sexes have been modified,
the female for protection and the male for sexual attrac-
tion, can only be definitely decided when we know the
life-history of each species.

Without distinct evidence, I am unwilling to admit
that a double process of selection has long been going
on with a multitude of species, — the males having been
rendered more brilliant by beating their rivals ; and the
females more dull-coloured by having escaped from their
enemies. We may take as an instance the common brim-
stone butterfly (G-onepteryx), which appears early in the
spring before any other kind. The male of this species
is of a far more intense yellow than the female, though
she is almost equally conspicuous ; and in this case it
does not seem probable that she specially acquired
her pale tints as a protection, though it is probable
that the male acquired his bright colours as a sexual
attraction. The female of Anthocharis cardamines does
not possess the beautiful orange tips to her wings with
which the male is ornamented ; consequently she closely
resembles the white butterflies (Pieris) so common in
our gardens ; but we have no evidence that this resem-
blance is beneficial. On the contrary, as she resembles
both sexes of several species of the same genus inhabit-
ing various quarters of the world, it is more probable
that she has simply retained to a large extent her
primordial colours.

Various facts support the conclusion that with the
greater number of brilliantly-coloured Lepidoptera, it
is the male which has been modified ; the two sexes
having come to differ from each other, or to resemble
each other, according to which form of inheritance has
prevailed. Inheritance is governed by so many un-

410 SEXUAL SELECTION. Part II.

known laws or conditions, that they seem to us to be
most capricious in their action ; 23 and we can so far
understand how it is that with closely-allied species the
sexes of some differ to an astonishing degree, whilst
the sexes of others are identical in colour. As the
successive steps in the process of variation are neces-
sarily all transmitted through the female, a greater
or less number of such steps might readily become
developed in her; and thus we can understand the
frequent gradations from an extreme difference to no
difference at all between the sexes of the species within
the same group. These cases of gradation are much
too common to favour the supposition that we here see
females actually undergoing the process of transition
and losing their brightness for the sake of protection ;
for we have every reason to conclude that at any
one time the greater number of species are in a fixed
condition. With respect to the differences between the
females of the species in the same genus or family, we
can perceive that they depend, at least in part, on the
females partaking of the colours of their respective
males. This is well illustrated in those groups in which
the males are ornamented to an extraordinary degree ;
for the females in these groups generally partake to a
certain extent of the splendour of their male partners.
Lastly, we continually find, as already remarked, that
the females of almost all the species in the same genus,
or even family, resemble each other much more closely
in colour than do the males ; and this indicates that
the males have undergone a greater amount of modifi-
cation than the females.

23 < The Variation of Animals and Plants under Domestication,
vol. ii. chap. xii. p. 17.

Chap. XL BUTTERFLIES AND MOTHS. 411

Mimickry. — This principle was first made clear in an
admirable paper by Mr. Bates,24 who tlms threw a flood
of light on many obscure problems. It had previously
been observed that certain butterflies in S. America
belonging to quite distinct families, resembled the Heli-
conidee so closely in every stripe and shade of colour
that they could not be distinguished except by an
experienced entomologist. As the HeliconidaB are
coloured in their usual manner, whilst the others depart
from the usual colouring of the groups to which they
belong, it is clear that the latter are the imitators, and
the Heliconidse the imitated. Mr. Bates further observed
that the imitating species are comparatively rare, whilst
the imitated swarm in large numbers ; the two sets
living mingled together. From the fact of the Heli-
conidae being conspicuous and beautiful insects, yet
so numerous in individuals and species, he concluded
that they must be protected from the attacks of birds
by some secretion or odour ; and this hypothesis has
now been confirmed by a considerable body of curious
evidence.25 From these considerations Mr. Bates in-
ferred that the butterflies which imitate the protected
species had acquired their present marvellously decep-
tive appearance, through variation and natural selection,
in order to be mistaken for the protected kinds and
thus to escape being devoured. No explanation is
here attempted of the brilliant colours of the imi-
tated, but only of the imitating butterflies. We must
account for the colours of the former in the same
general manner, as in the cases previously discussed in
this chapter. Since the publication of Mr. Bates' paper,
similar and equally striking facts have been observed

24 ' Transact. Linn. Soc.' vol. xxiii. 1862, p. 495.

25 ' Proc. Ent. Soc' Dec. 3rd, I860, p. xlv.

412 SEXUAL SELECTION. Part II.

by Mr. Wallace26 in the Malayan region, and by
Mr. Trimen in South Africa.

As some writers 27 have felt much difficulty in under-
standing how the first steps in the process of mimickry
could have been effected through natural selection, it
may be well to remark that the process probably has
never commenced with forms widely dissimilar in colour.
But with two species moderately like each other, the
closest resemblance if beneficial to either form could
readily be thus gained ; and if the imitated form was
subsequently and gradually modified through sexual
selection or any other means, the imitating form would
be led along the same track, and thus be modified to
almost any extent, so that it might ultimately assume
an appearance or colouring wholly unlike that of the
other members of the group to which it belonged. As
extremely slight variations in colour would not in many
cases suffice to render a species so like another pro-
tected species as to lead to its preservation, it should
be remembered that many species of Lepidoptera are
liable to considerable and abrupt variations in colour.
A few instances have been given in this chapter ; but
under this point of view Mr. Bates' original paper on
mimickry, as well as Mr. Wallace's papers, should be
consulted.

In the foregoing cases both sexes of the imitating
species resemble the imitated ; but occasionally the

26 ' Transact. Linn. Soc' vol. xxv. 1865, p. 1 ; also ' Transact. Ent.
Soc' vol. iv. (3rd series), 1867, p. 301.

2' See an ingenious article entitled, " Difficulties of the Theory of
Natural Selection," in the ' Month,' 1869. The writer strangely sup-
poses that I attribute the variations in colour of the Lepidoptera, by
which certain species belonging to distinct families have come to
resemble others, to reversion to a common progenitor ; but there is no
more reason to attribute these variations to reversion than in the case
of any ordinary variation.

Chap. XI. BUTTERFLIES AND MOTHS. 41

o

female alone mocks a brilliantly-coloured and protected
species inhabiting the same district. Consequently the
female differs in colour from her own male, and, which
is a rare and anomalous circumstance, is the more
brightly-coloured of the two. In all the few species of
Pieridae, in which the female is more conspicuously
coloured than the male, she imitates, as I am informed
by Mr. Wallace, some protected species inhabiting the
same region. The female of Diadema anomala is rich
purple-brown with almost the whole surface glossed with
satiny blue, and she closely imitates the Eujoloea mida-
mus, "one of the commonest butterflies of the East;"
whilst the male is bronzy or olive-brown, with only a
slight blue gloss on the outer parts of the wings.28
Both sexes of this Diadema and of D. bolina follow
the same habits of life, so that the differences in colour
between the sexes cannot be accounted for by exposure
to different conditions;29 even if this explanation were
admissible in other instances.30

The above cases of female butterflies which are more
brightly-coloured than the males, shew us, firstly, that
variations have arisen in a state of nature in the female
sex, and have been transmitted exclusively, or almost ex-
clusively, to the same sex ; and, secondly, that this form
of inheritance has not been determined through natural
selection. For if we assume that the females, before
they became brightly coloured in imitation of some pro-
tected kind, were exposed during each season for a longer
period to danger than the males ; or if we assume that

28 Wallace, "Notes on Eastern Butterflies," 'Transact. Ent. Soc.'
1869, p. 287.

29 Wallace, in 'Westminster Eeview,' July, 1867, p. 37; and in
' Journal of Travel and Nat. Hist.' vol. i. 1868, p. 88.

30 See remarks by Messrs. Bates and Wallace, in ' Proc. Ent. Soc.'
Nov. 19, 1866, p. xxxix.

414 SEXUAL SELECTION. Part II.

they could not escape so swiftly from their enemies,
we can understand how they alone might originally
have acquired through natural selection and sexually-
limited inheritance their present protective colours.
But except on the principle of these variations having
been transmitted exclusively to the female offspring,
we cannot understand why the males should have re-
mained dull-coloured ; for it would surely not have
been in any way injurious to each individual male to
have partaken by inheritance of the protective colours
of the female, and thus to have had a better chance
of escaping destruction. In a group in which brilliant
colours are so common as with butterflies, it cannot be
supposed that the males have been kept dull-coloured
through sexual selection by the females rejecting the
individuals which wrere rendered as beautiful as them-
selves. We may, therefore, conclude that in these cases
inheritance bv one sex is not due to the modification
through natural selection of a tendency to equal inherit-
ance by both sexes.

It may be well here to give an analogous case in
another Order, of characters acquired only by the female,
though not in the least injurious, as far as we can judge,
to the male. Amongst the Phasinicke, or spectre-insects,
Mr. Wallace states that " it is often the females alone
" that so strikingly resemble leaves, while the males show
" only a rude approximation." Now, whatever may be
the habits of these insects, it is highly improbable that
it could be disadvantageous to the males to escape de-
tection by resembling leaves.31 Hence we may conclude

31 See Mr. Wallace in 'Westminster Keview,' July, 1867, p. 11 and
37. The male of no butterfly, as Mr. Wallace informs me, is known to
differ in colour, as a protection, from the female ; and he asks me how
I can explain this fact on the principle that one sex alone has varied
and has transmitted its variations exclusively to the same sex, without

Chap. XL BUTTERFLIES AND MOTHS. 415

that the females alone in this latter as in the previous
cases originally varied in certain characters ; these cha-
racters having been preserved and augmented through
ordinary selection for the sake of protection and from
the first transmitted to the female offspring alone.

Bright Colours of Caterpillars. — Whilst reflecting on
the beauty of many butterflies, it occurred to me that
some caterpillars were splendidly coloured, and as
sexual selection could not possibly have here acted,
it appeared rash to attribute the beauty of the mature
insect to this agency, unless the bright colours of their
larvae could be in some manner explained. In the first
place it may be observed that the colours of caterpillars
do not stand in any close correlation with those of the
mature insect. Secondly, their bright colours do not

the aid of selection to check the variations being inherited by the
other sex. No doubt if it could be shewn that the females of very
many species had been rendered beautiful through protective miinickry,
but that this has never occurred with the males, it would be a serious
difficulty. But the number of cases as yet known hardly suffices for a
fair judgment. We can see that the males, from having the power of
flying more swiftly, and thus escaping danger, would not be so likely
as the females to have had their colours modified for the sake of protec-
tion ; but this would not in the least have interfered with their receiving
protective colours through inheritance from the females. In the second
place, it is probable that sexual selection would actually tend to prevent
a beautiful male from becoming obscure, for the less brilliant individuals
would be less attractive to the females. Supposing that the beauty of
the male of any species had been mainly acquired through sexual
selection, yet if this beauty likewise served as a protection, the acquisi-
tion would have been aided by natural selection. But it would be
quite beyond our power to distinguish between the two processes of
sexual and ordinary selection. Hence it is not likely that we should
be able to adduce cases of the males having been rendered brilliant
exclusively through protective mimickry, though this is comparatively
easy with the females, which have rarely or never been rendered beau-
tiful, as far as we can judge, for the sake of sexual attraction, although
they have often received beauty through inheritance from their male
parents.

416 SEXUAL SELECTION. Part II.

serve in any ordinary manner as a protection. As an
instance of this, Mr. Bates informs me that the most
conspicuous caterpillar which he ever beheld (that of a
Sphinx) lived on the large green leaves of a tree on the
open llanos of South America ; it was about four inches
in length, transversely banded with black and yellow,
and with its head, legs, and tail of a bright red. Hence
it caught the eye of any man who passed by at the
distance of many yards, and no doubt of every passing
bird.

I then applied to Mr. Wallace, who has an innate
genius for solving difficulties. After some consideration
he replied : " Most caterpillars require protection, as
" may be inferred from some kinds being furnished
" with spines or irritating hairs, and from many being
" coloured green like the leaves on which they feed,
'•' or curiously like the twigs of the trees on which they
" live." I may add as another instance of protection,
that there is a caterpillar of a moth, as I am informed
by Mr. J. Mansel Weale, which lives on the mimosas in
South Africa, and fabricates for itself a case, quite un-
distinguishable from the surrounding thorns. From
such considerations Mr. Wallace thought it probable
that conspicuously-coloured caterpillars were protected
by having a nauseous taste ; but as their skin is ex-
tremely tender, and as their intestines readily protrude
from a wound, a slight peck from the beak of a bird
would be as fatal to them as if they had been devoured.
Hence, as Mr. Wallace remarks, " distastefulness alone
" would be insufficient to protect a caterpillar unless
" some outward sign indicated to its would-be destroyer
" that its prey was a disgusting morsel." Under these
circumstances it would be highly advantageous to a
caterpillar to be instantaneously and certainly recog-
nised as unpalatable by all birds and other animals.

Chap. XI. SUMMAEY ON INSECTS. 417

Thus the most gaudy colours would be serviceable, and
might have been gained by variation and the survival
of the most easily-recognised individuals.

This hypothesis appears at first sight very bold ; but
when it was brought before the Entomological Society 32
it was supported by various statements ; and Mr. J.
Jenner Weir, who keeps a large number of birds in an
aviary, has made, as he informs me, numerous trials,
and finds no exception to the rule, that all caterpillars
of nocturnal and retiring habits with smooth skins,
all of a green colour, and all which imitate twigs, are
greedily devoured by his birds. The hairy and spinose
kinds are invariably rejected, as were four conspicuously-
coloured species. When the birds rejected a caterpillar,
they plainly shewed, by shaking their heads and cleans
ing their beaks, that they were disgusted by the taste.
Three conspicuous kinds of caterpillars and moths were
also given by Mr. A. Butler to some lizards and frogs,
and were rejected ; though other kinds were eagerly
eaten. Thus the probable truth of Mr. Wallace's view
is confirmed, namely, that certain caterpillars have been
made conspicuous for their own good, so as to be easily
recognise 1 by their enemies, on nearly the same prin-
Ciple that certain poisons are coloured by druggists for
the good of man. This view will, it is probable, be
hereafter extended to many animals, which are coloured
in a conspicuous manner.

Summary and Concluding Remarks on Insects. —
Looking back to the several Orders; we have seen that
the sexes often differ in various characters, the meaning

33

32 « Proc. Eutomolog. Soc' Dec. 3rd, 1866, p. xlv., and March 4th,

1867, p. Ixxs.

33 See Mr. J. Jenner Weir's paper on insects and insectivorous birds,
in ' Transact. Ent. Soc." 1869, p. 21 ; also Mr. Butler's paper, ibid p. 27.

VOL. I. ^ E

418 SEXUAL SELECTION. Part II.

of which is not understood. The sexes, also, often differ
in their organs of sense or locomotion, so that the males
may quickly discover or reach the females, and still
oftener in the males possessing diversified contrivances
for retaining the females when found. But we are not
here much concerned with sexual differences of these
kinds.

In almost all the Orders, the males of some species,
even of weak and delicate kinds, are known to be highly
pugnacious ; and some few are furnished with special
weapons for fighting with their rivals. But the law of
battle does not prevail nearly so widely with insects as
with the higher animals. Hence probably it is that the
males have not often been rendered larger and stronger
than the females. On the contrary they are usually
smaller, in order that they may be developed within a
shorter time, so as to be ready in large numbers for the
emergence of the females.

In two families of the Homoptera the males alone
possess, in an efficient state, organs which may be called
vocal ; and in three families of the Orthoptera the males
alone possess stridulating organs. In both cases these
organs are incessantly used during the breeding-season,
not only for calling the females, but for charming or
exciting them in rivalry with other males. No one
who admits the agency of natural selection, will dispute
that these musical instruments have been acquired
through sexual selection. In four other Orders the
members of one sex, or more commonly of both sexes,
are provided with organs for producing various sounds,
which apparently serve merely as call-notes. Even
when both sexes are thus provided, the individuals
which were able to make the loudest or most continuous
noise would gain partners before those which were less
noisy, so that their organs have probably been gained

Chap. XI. SUMMAKY ON INSECTS: 419

through sexual selection. It is instrutive to reflect
on the wonderful diversity of the means for producing
sound, possessed by the males aloue or by both sexes
in no less than six Orders, and which were possessed
by at least one insect at an extremely remote geolo-
gical epoch. We thus learn how effectual sexual selec-
tion has been in leading to modifications of structure,
which sometimes, as with the Homoptera, are of an im-
portant nature.

From the reasons assigned in the last chapter, it is
probable that the great horns of the males of many
lamellicorn, and some other beetles, have been ac-
quired as ornaments. So perhaps it may be with cer-
tain other peculiarities confined to the male sex. From
the small size of insects, we are apt to undervalue their
appearance. If we could imagine a male Chalcosoma
(fig. 15) with its polished, bronzed coat of mail, and
vast complex horns, magnified to the size of a horse or
even of a dog, it would be one of the most imposing
animals in the world.

The colouring of insects is a complex and obscure
subject. When the male differs slightly from the female,
and neither are brilliantly coloured, it is probable that
the two sexes have varied in a slightly different manner,
with the variations transmitted to the same sex, without
any benefit having been thus derived or evil suffered.
When the male is brilliantly coloured and differs con-
spicuously from the female, as with some dragon-flies
and many butterflies, it is probable that he alone has
been modified, and that he owes his colours to sexual
selection ; whilst the female has retained a primordial
or very ancient type of colouring, slightly modified by
the agencies before explained, and has therefore not
been rendered obscure, at least in most cases, for the
sake of protection. But the female alone has soine-

2 e 2

420 SEXUAL SELECTION. Fart II.

times been coloured brilliantly so as to imitate other
protected species inhabiting the same district. When
the sexes resemble each other and both are obscurely
coloured, there is no doubt that they have been in a
multitude of cases coloured for the sake of protection.
So it is in some instances when both are brightly
coloured, causing them to resemble surrounding objects
such as flowers, or other protected species, or indirectly
by giving notice to their enemies that they are of an
unpalatable nature. In many other cases in which the
sexes resemble each other and are brilliantly coloured,
especially when the colours are arranged for display, we
may conclude that they have been gained by the male
sex as an attraction, and have been transferred to both
sexes. We are more especially led to this conclusion
whenever the same type of coloration prevails through-
out a group, and we find that the males of some species
differ widely in colour from the females, whilst both
sexes of other species are quite alike, with intermediate
gradations connecting these extreme states.

In the same manner as bright colours have often
been partially transferred from the males to the females,
so it has been with the extraordinary horns of many
lamellicorn and some other beetles. So, again, the
vocal or instrumental organs proper to the males of
the Homoptera and Orthoptera have generally been
transferred in a rudimentary, or even in a nearly perfect
condition to the females ; yet not sufficiently perfect to
be used for producing sound. It is also an interesting
fact, as bearing on sexual selection, that the stridulating
organs of certain male Orthoptera are not fully deve-
loped until the last moult ; and that the colours of cer-
tain male dragon-flies are not fully developed until
some little time after their emergence from the pupal
state, and when they are ready to breed.

Chap. XI. SUMMAEY ON INSECTS. 421

Sexual selection implies that the more attractive
individuals are preferred by the opposite sex ; and as
with insects, when the sexes differ, it is the male which,
with rare exceptions, is the most ornamented and
departs most from the type to which the species
belongs ; — and as it is the male which searches eagerly
for the female, we must suppose that the females habit-
ually or occasionally prefer the more beautiful males,
and that these have thus acquired their beauty. That
in most or all the orders the females have the power
of rejecting any particular male, we may safely infer
from the many singular contrivances possessed by the
males, such as great jaws, adhesive cushions, spines,
elongated legs, &c, for seizing the female ; for these
contrivances shew that there is some difficulty in the
act. In the case of unions between distinct species,
of which many instances have been recorded, the
female must have been a consenting party. Judging
from what we know of the perceptive powers and
affections of various insects, there is no antecedent im-
probability in sexual selection having come largely into
action ; but we have as yet no direct evidence on this
head, and some facts are opposed to the belief. Never-
theless, when we see many males pursuing the same
female, we can hardly believe that the pairing is left to
blind chance — that the female exerts no choice, and
is not influenced by the gorgeous colours or other
ornaments, with which the male alone is decorated.

If we admit that the females of the Homoptera and
Orthoptera appreciate the musical tones emitted by their
male partners, and that the various instruments for this
purpose have been perfected through sexual selection,
there is little improbability in the females of other
insects appreciating beauty in form or colour, and con-
sequently in such characters having been thus gained

422 SEXUAL SELECTION. Part II.

by the males. But from the circumstance of colour
being so variable, and from its having been so often
modified for the sake of protection, it is extremely
difficult to decide in how large a proportion of cases
sexual selection has come into play. This is more
especially difficult in those Orders, such as the Orthop-
tera, Hymenoptera, and Coleoptera, in which the two
sexes rarely differ much in colour ; for we are thus cut
off from our best evidence of some relation between the
reproduction of the species and colour. With the
Coleoptera, however, as before remarked, it is in the
great lamellicorn group, placed by some authors at
the head of the Order, and in which we sometimes
see a mutual attachment between the sexes, that we
find the males of some species possessing weapons for
sexual strife, others furnished with wonderful horns,
many with stridulating organs, and others ornamented
with splendid metallic tints. Hence it seems probable
that all these characters have been gained through
the same means, namely sexual selection.

When we treat of Birds, we shall see that they pre-
sent in their secondary sexual characters the closest
analogy with insects. Thus, many male birds are
highly pugnacious, and some are furnished with special
weapons for fighting with their rivals. They possess
organs which are used during the breeding-season for
producing vocal and instrumental music. They are
frequently ornamented with combs, horns, wattles and
plumes of the most diversified kinds, and are decorated
with beautiful colours, all evidentlv for the sake of dis-
play. We shall find that, as with insects, both sexes,
in certain groups, are equally beautiful, and are equally
provided with ornaments which are usually confined to
the male sex. In other groups both sexes are equally
plain-coloured and unornamented. Lastly, in some few

Chap. XI.

SUMMARY ON INSECTS.

423

anomalous cases, the females are more beautiful than
the males. We shall often find, in the same group of
birds, every gradation from no difference between the
sexes, to an extreme difference. In the latter case we
shall see that the females, like female insects, often
possess more or less plain traces of the characters which
properly belong to the males. The analogy, indeed, in
all these respects between birds and insects, is curiously
close. Whatever explanation applies to the one class
probably applies to the other ; and this explanation, as
we shall hereafter attempt to shew, is almost certainly
sexual selection.

END OF VOL. I.

&\C4

LONDON :

PRINTED BY WILLIAM CLOWES AND SONS, DUKE STREET, STAMFORD STUEF.T,

AND CHARING CROSS.

50a, Albemarle Street, London,
January, 1871.

MR. MURRAY'S

LIST OF POPULAR WORKS.

One Shilling.

THE PRINCIPAL SPEECHES AND ADDRESSES OF

H.R.H. THE PRINCE CONSORT, with an Introduction giving some Out-
lines of his Character. With Portrait.

MUSIC AND THE ART OF DRESS. Two Essays. By A

LADY.

THE FALL OF JERUSALEM. A Dramatic Poem. By DEAN
MILMAN.

THEODORE HOOK. A Sketch. By J. G. LOCKHARt!

HISTORY OF THE GUILLOTINE. By Rt. Hon. J. W.

CROKER. Woodcuts.

THE CHACE. A Descriptive Essay. By C. J. APPERLEY

(Nimrod). With Woodcuts.

REJECTED ADDRESSES; or, The New Theatrum Poetarum.
By HORACE and JAMES SMITH. With Woodcuts.

THE ROAD. A Descriptive Essay. By C. J. APPERLEY

(Nimrod). With Woodcuts.

MAXIMS AND HINTS ON ANGLING, CHESS, SHOOTING,

AND OTHER MATTERS; also, the Miseries of Fishing. By RICHARD
PENN. With Woodcuts.

THE STORY OF JOAN OF ARC. By LORD MAHON.

THE PROGRESS OF LITERATURE AND SCIENCE—
THE STUDY OF HISTORY— AND ANTIQUITIES AND ART IN
ROME. By EARL STANHOPE.

CHILDE HAROLD'S PILGRIMAGE. A Romaunt. By LORD
BYRON. With Woodcuts.

HISTORY AND ANTIQUITIES OF NORTHAMPTONSHIRE.
By REV. THOMAS JAMES.

THE HONEY BEE. By REV. THOMAS JAMES.

THE FLOWER GARDEN, with an Essay on the Poetry of
Gardening. By REV. THOMAS JAMES.

B

LIST OF POPULAR WORKS

One Shilling and Sixpence.
THE AKT OF DINING ; or, Gastronomy and Gastronomers.

By A. HAYWARD.

WELLINGTON ; — His Character, — Actions, — and Writings.

By JULES MAUREL. With Preface by Loed Ellesmeke.

WORKMEN'S EARNINGS -STRIKES— AND SAVINGS. By

SAMUEL SMILES.

THE TURF. A Descriptive Essay. By C. J. APPERLEY

(Nimrod). "With Woodcuts.

THE STORY OF PUSS IN BOOTS. WTith 12 Illustrations.
By OT 10 SPECKTER.

PROGRESSIVE GEOGRAPHY FOR CHILDREN. By Rt .

Hox. J. W. CROKER.

HYMNS, written and adapted to the Weekly Church Service of the
Year. By BISHOP HEBER.

Tivo Shillings.

THE AMBER WITCH : the most interesting Trial for Witchcraft.
From the German. By LADY DUFF GORDON.

OLIVER CROMWELL AND JOHN BUNYAN: Biographies.
By ROBERT SOUTHEY.

LIFE OF SIR FRANCIS DRAKE. WTith his Voyages and Ex-
ploits by Sea and Land. By JOHN BARROW.

CAMPAIGNS OF THE BRITISH ARMY AT WASHINGTON

AND NEW ORLEANS. By REV. G. R. GLE1G.

THE FRENCH IN ALGIERS ; the Soldier of the Foreign Legion
and Prisoners of Abd-el-Kadir. Translated by LADY DUFF GORDON.

HISTORY OF THE FALL OF THE JESUITS IN THE 18th
CENTURY. By COUNT ALEXIS DE ST. PRIEST.

LIVONIAN TALES ; the Disponent— the Wolves— the Jewess

By A LADY.

NOTES FROM LIFE. Money — Humility — Independence —
Wisdom — Choice in Marriage — Children — Lite Poetic. By HENRY
TAYLOR.

THE SIEGE OF GIBRALTAR, 1779-83. With a Descrip-
tion and Account of that Garrison from the Earliest Periods. By JOHN
DRINKWATER.

SIR ROBERT SALE'S BRIGADE IN AFFGHANISTAN AND

THE DEFENCE OF JELLALABAD. By REV. G. R. GLE1G.

PUBLISHED BY MR. MURRAY.

THE TWO SIEGES OF VIENNA BY THE TURKS. Trans-
lated from the German. By LORD ELLESMERE.

THE WAYSIDE CROSS ; or, The Raid of Gomez : a Tale of the
Carlist War. By CAPT. MILMAN.

ADVENTURES ON THE ROAD TO PARIS DURING THE

CAMPAIGNS OF 1S13-14. From the Autobiography of HENRY
STEFFENS.

STOKERS AND POKERS ; or, The North1- Western Railway.
— The Electric Telegraph— and Railway Clearing House. By SIR FRANCIS
HEAD.

TRAVELS IN EGYPT, NUBIA, SYRIA, AND THE HOLY

LAND, with a Journey round the Dead Sea, and through the Country East of
the Jordan. By IRBY and MANGLES.

WESTERN BARBARY : An Account of the Wild Tribes and
Savage Animals. By JOHN H. DRUMMOND HAY.

LETTERS FROM THE SHORES OF THE BALTIC. By A

LADY.

NOTES AND SKETCHES OF NEW SOUTH WALES

DURING A RESIDENCE OF MANY YEARS IN THAT COLONY.
By MRS. CHARLES MEREDITH.

RECOLLECTIONS OF BUSH LIFE IN AUSTRALIA, during
a Residence of Eight Years in the Interior. By REV. H. W. HAYGARTH.

JOURNAL OF A RESIDENCE AMONG THE NEGROES

IN THE WEST INDIES. By M. G.LEWIS.

MEMOIRS OF FATHER RIPA DURING THIRTEEN YEARS'

RESIDENCE AT THE COURT OF PEK1N, in the Service of the Emperor
of China, with an Account of the Foundation of the College for the Education
of Young Chinese in Naples. From the Italian. By FORTUNATO PRAND1.

PHILIP MUSGRAVE; or, Memoirs of a Church of England
Missionary in the North American Colonies. By REV. J. ABBOTT.

A MONTH IN NORWAY. By JOHN G. HOLLWAY.

LETTERS FROM MADRAS ; or, First Impressions of Life and
Manners in India. By A LADY.

ROUGH NOTES TAKEN DURING SOME RAPID RIDES

ACR< SS THE PAMPAS AND AMONG THE ANDES. By SIR FRANCIS
HEAD.

A VOYAGE UP THE RIVER AMAZON, including a Visit to

PARA. ByAYILLIAM H. EDWARDS.

e %

4 LIST OF POPULAR WORKS

A POPULAE ACCOUNT of the MANNERS and CUSTOMS

OF INDIA. By REV. CHARLES ACLAND.

ADVENTURES IN THE LIBYAN DESERT AND THE
OASIS OF JUPITER AMMON. By BAYLE ST. JOHN.

LIFE OF SIR FOWELL BUXTON. By his Son, CHARLES
BUXTON. With an Inquiry into the Results of Emancipation.

Two Shillings and Sixpence.

THE CONTINUITY OF SCRIPTURE, as declared by the
Testimony of our Lord and of the Evangelists and Apostles. By LORD
HATHERLEY.

LIFE OF LORD BACON. By LORD CAMPBELL.
CHILDE HAROLD'S PILGRIMAGE. A Romaunt. By LORD

BYRON.

TALES AND POEMS. By LORD BYRON.

LITTLE ARTHUR'S HISTORY OF ENGLAND. By LADY

CALLCOTT. With 28 Woodcuts.

THE EMIGRANT. By SIR FRANCIS HEAD.

THE FABLES OF ^ESOP ; A New Veksion, chiefly from
Original Sources. By REV. THOMAS JAMES. With 100 Woodcuts.

STORIES FOR CHILDREN SELECTED FROM THE HIS-
TORY OF ENGLAND. By Rt. Hon. J. W. CROKER. With 24 Woodcuts.

THE COMPLETE POETICAL WORKS OF LORD BYRON.

THE PHILOSOPHY OF THE MORAL FEELINGS. By

JOHN ABERCROMBIE.

POEMS AND SONGS. By ALLAN CUNNINGHAM. With

Biographical Notice.

THE FRANCHISE ; a Privilege and not a Right. Proved by the

Political Experience of the Ancients. By H. S. TREMENHEERE.

Three Shillings.

" THE FORTY-FIVE ; " A Narrative of the Insurrection of
1745 in Scotland. To which are added Letters of Prince Charles Stuart.
By LORD MAHON.

PUBLISHED BY MR. MURRAY.

Three Shillings and Sixpence.

THE BIBLE IN SPAIN; or, The Adventures and Imprisonment

ot an Englishman in attempting to Circulate the Scriptures in the Peninsula.
By GEORGE BORROW.

THE GYPSIES OF SPAIN; their Manners, Customs, Religion,
and Language. By GEORGE BORROW'.

THE BEAUTIES OF LORD BYRON'S WRITINGS ; Poetry
and Prose. With Portrait.

LIFE OF LOUIS PRINCE OF CONDE, SURNAMED THE

GREAT. By LORD MAHON.

SKETCHES OF GERMAN LIFE AND SCENES FROM THE
WAR OF LIBERATION IN GERMANY. By VARNHAGEN VON
ERESE. Translated by SIR ALEXANDER DUFF GORDON.

THE TRUE STORY OF THE BxiTTLE OF WATERLOO. By

REV. G. R. GLEIG.

DEEDS OF NAVAL DARING; or, Anecdotes op the
British Navy. By EDWARD GltFARD.

TYPEE ; or, The Marquesas Islanders. By HERMAN MELVILLE.

OMOO : A Narrative of Adventures in the South Seas ; a Sequel
to 'Typee.' By HERMAN MELVILLE.

THE BIBLE IN THE HOLY LAND; being Extracts from

Dean Stanley's 'Sinai and Palestine.' With Woodcuts.

AN ESSAY ON ENGLISH POETRY. With Short Notices of the
British Poets. By THOMAS CAMPBELL.

HISTORICAL AND CRITICAL ESSAYS. By LORD MAHON.

LIFE OF LORD CLIVE. By REV. G. R. GLEIG.

LIFE OF SIR THOMAS MUNRO. With Selections from his
Correspondence. By REV. G. R. GLEiG.

THE RISE OF OUR INDIAN EMPIRE : being a History of

British India from its Origin till the Peace of 17o3. By LORD MAHON.

6 LIST OF POPULAR WORKS

A NARRATIVE OF THE SIEGE OF KAES, and of the Six

Months' Resistance by the Turkish Garrison under General Williams. With
Travels and Adventures in Armenia, &c. By HUMPHREY SANDW1TH,
M.D.

SKETCHES OF PERSIA. By SIR JOHN MALCOLM.

THE WILD SPORTS AND NATURAL HISTORY OF THE

HIGHLANDS. By CHARLES ST. JOHN.

GATHERINGS FROM SPAIN. By RICHARD FORD.
TRAVELS IN MEXICO AND THE ROCKY MOUNTAINS.

By G. F. RUXTON.

PORTUGAL AND GALLICIA ; with an Account of the
Social and Political state of the Basque Provinces. By LORD
CARNARVON.

A RESIDENCE AT SIERRA LEONE. Described from a
Journal kept on the Spot, and Letters written to Friends at Home. By
A LADY.

THE REMAINS IN VERSE AND PROSE OF ARTHUR

HENRY HALLAM. With Preface, Memoir, and Portrait.

THE POETICAL WORKS OF BISHOP HEBER ; containing

Palestine, Europe, the Red Sea, Hymns, &c. With Portrait.

GLEANINGS IN NATURAL HISTORY. By EDWARD

JESSE. With Woodcuts.

THE REJECTED ADDRESSES: or, The New Theatrura
Poetarum. By HORACE and JAMES SMITH. With Portrait and
Woodcuts.

CONSOLATIONS IN TRAVEL ; or, Tbe Last Days of a Philoso-
pher. By SIR HUMPHRY DAVY. With Woodcuts.

SALMONIA; or, Days of Fly-fishing. By SIR HUMPHRY

DAVY. With Woodcuts.

THE INTELLECTUAL POWERS AND THE INVESTIGA-
TION OF TKUTH. By JOHN ABERCROMBIE.

SPECIMENS OF THE TABLE-TALK OF THE LATE

SAMUEL TAYLOR COLERIDGE. With Portrait.

PRACTICAL INSTRUCTIONS IN GARDENING, for every
Month in the Year. By MRS. LOUDON. With Woodcuts.

PUBLISHED BY MR. MURRAY.

A SMALLER HISTORY OF ENGLAND. Edited by DE. WM.
SMITH. With Woodcuts.

A SMALLER HISTORY OF GREECE. Edited by DR. WM.
SMITH. With Woodcuts.

A SMALLER HISTORY OF ROME. Edited by DR. WM.
SMITH. With Woodcuts.

A SMALLER CLASSICAL MYTHOLOGY. With Translations
from the Ancient Poets. Edited by DR. WM. SMITH. With Woodcuts.

A SMALLER ANCIENT HISTORY, from the Earliest Times
to the Conquest of Alexander the Great. Edited by DR. WM. SMITH.
With Woodcuts.

A SMALLER HISTORY OF ENGLISH LITERATURE, from

the earliest period to the Georgian Era. Pxlited by DR. WM. SMITH.

SPECIMENS OF THE CHIEF WRITERS IN ENGLISH

LITERATURE. Chronologically arranged. Edited by DR. AVM. SMITH.

A SMALLER SCRIPTURE HISTORY.— I. Old Testament His-
tory ; II. Connection of Old and New Testaments; III. New Testament
History to a.d. 70. Edited by DR. WM. SMITH. With Woodcuts.

Four Shillings.

HISTORY OF ENGLAND, from the First Invasion by the
Romans, continued down to 1865. With Conversations at the end of each
Chapter. By MRS. MARKHAM. With 100 Woodcuts.

HISTORY OF FRANCE, from the Conquest by the Gauls, con-
tinued down to 1867. With Conversations at the end of each Chapter.
By MRS. MARKHAM. With 70 Woodcuts.

HISTORY OF GERMANY, from the Invasion of the Kingdom by
the Romans under Marids, continued down to 1867. On the Plan of MRS.
MARKHAM. With 50 Woodcuts.

SHALL AND WILL ; or, the Future Auxiliary Verb. By SIR
EDMUND HEAD.

Four Shillings and Sixpence.

CHILDREN OF THE LAKE. A Poem. By EDWARD
SALLESBURY.

A LADY'S DIARY OF THE SIEGE OF LUCKNOW.

HOUSEHOLD SURGERY ; or, Hints on Emergencies. By JOHN
F. SOUTH. With Woodcuts.

8 LIST OF POPULAR WORKS

Five Shillings.
ANCIENT SPANISH BALLADS ; Historical and Komantic.

Translated with Notes by J. G. LOCKHART. With Portrait and Illustra-
tions.

MISCELLANIES. By LOBD BYKON. 2 vols.

INTKODUCTIONS TO THE STUDY OF THE GREEK

CLASSIC POETS. By HENRY NELSON COLERIDGE.

HYMNS IN PROSE FOR CHILDREN. By MRS. BARBAULD.

With 112 Illustrations.

RECOLLECTIONS OF THE DRUSES, and some Notes on

their Religion. By LORD CARNARVON.

THE ORIGIN OF LANGUAGE. BASED ON MODERN RE-
SEARCHES. By REV. F. W. FARRAR.

MODERN DOMESTIC COOKERY. Founded on Principles of
Economy and Practical Knowledge, and adapted for Private Families.
With Woodcuts.

DRAMAS AND PLAYS. By LORD BYRON. 2 vols.

THE HORSE AND HIS RIDER. By SIR FRANCIS HEAD.

With Woodcuts.

HANDBOOK OF FAMILIAR QUOTATIONS, chiefly from

English Authors.

THE CHACE— THE TURF— AND THE ROAD. A Series of
Popular Essays. By C. J. AFPERLEY (Nijiuod). With Portrait and
Illustrations.

AUNT IDA'S WALKS AND TALKS. A Story Book for
Children. By A LADY.

STpRIES FOR DARLINGS. A Book for Boys and Girls. With

Illustrations.

THE CHARMED ROE. A Story Book for Young People. Illus-
trated by OTTO SPECKTER.

DON JUAN AND BEPPO. By LORD BYRON. 2 vols.

LIFE IN THE LIGHT OF GOD'S WORD. By ARCHBISHOP

THOMSON, D.D.

ATHENS AND ATTICA; Notes of a Tour. By BISHOP
WORDSWORTH, D.D. With Illustrations.

ANNALS OF THE WARS— XVIIIth CENTURY, 1700-1799.
Compiled from the most Authentic Sources. By SIR EDWARD OUST, D.C.L.
With Maps. 5 vols. Post 8vo. 5s. each.

ANNALS OF THE WARS — XIXth CENTURY, 1800-15.

Compiled from the most Authentic Sources. By SIR EDWARD CUST.
4 vols. Fcap. 8vo. 5s. each.

PUBLISHED BY MR. MURRAY.

Six Shillings.

BENEDICITE ; or, THE SONG of the THREE CHILDREN.

Being Illustrations of the Power, Beneficence, and Design inanilested by
the Creator in His Works. By DR. CHAPLIN CHILD.

OLD DECCAN DAYS; or, HINDOO FAIRY LEGENDS

current in Southern India. By M. FRERE. With Introduction by Sir
Bartle Frere. With Illustrations.

THE WILD GARDEN; or, Our Groves and Shrubberies

MADE BEAUTIFUL BY THE NATURALIZATION OF HaRDY EXOTIC PLANTS.

By WILLIAM ROBINSON. With Frontispiece.

MISSIONARY TRAVELS AND RESEARCHES IN SOUTH

AFRICA. By DaVID LIVINGSTONE, M.D. With Map and Illustrations.

FIVE YEARS OF A HUNTER'S LIFE IN SOUTH AFRICA ;

By GORDON CLIMM1NG. With Illustrations.

THOUGHTS ON ANIMALCULES ; or, The Invisible World, as
revealed by the Microscope. By GIDEON A. MANTELL. With Plates.

LIVES OF BRINDLEY AND THE EARLY ENGINEERS.

By SAMUEL SMILES. With Woodcuts.

LIFE OF TELFORD. With a History of Roads and Travelling

in England. By SAMUEL SMILES. With Woodcuts.

LIVES OF GEORGE AND ROBERT STEPHENSON. By

SAMUEL SMILES. With Woodcuts.

SELF-HELP. With Illustrations of Character and Conduct. By

SAMUEL SMILES.

INDUSTRIAL BIOGRAPHY : Iron-workers and Toolrnakers. A
Sequel to ' Self-help.' By SAMUEL SMILES.

THE HUGUENOTS IN ENGLAND AND IRELAND : their

Settlements, Churches, and Industries. By SAMUEL SMILES.

WILD WALES ; its People, Language, and Scenery. With In-
troductory Remarks. By GEORGE RORROW.

A MANUAL OF ETHNOLOGY; or, A Popular History of the
Races of the Old World. By CHARLES L. BRACE.

Seven Shillings.
JOURNALS OF A TOUR IN INDIA. By BISHOP HEBER.

2 vols.

ADVENTURES AMONG THE MARQUESAS AND SOUTH

SEA ISLANDERS. By HERMAN MELViLLE. 2 vols.

CONSTITUTIONAL PROGRESS. By MONTAGU BURROWS,

M.A.

LIFE AND POETICAL WORKS OF REV. GEORGE

CRABiiE. Edited by HIS SON. With Notes, Portrait, and Illustrations.

10 LIST OF POPULAR WORKS

Seven Shillings and Sixpence.

THE AET OF TRAVEL ; or. Hints on the Shifts and Con-
trivances available in Wild Countries. By FRANCIS GALLON. With
"Woodcuts.

VISITS TO THE MONASTERIES OF THE LEVANT. By

HON. R. CURZON. With Illustrations.

LETTERS FROM HIGH LATITUDES ; an Account of a Yacht
Voyage to Iceland, Jan Mayen, and Spitzbergen, &c. By LORD DUFFER1N.
With Illustrations.

BUBBLES FROM THE BRUNNEN OF NASSAU. By an

Old Man (SIR FRANCIS HEAD). With Illustrations.

NINEVEH AND ITS REMAINS ; a Narrative of an Expedition
to Assyria in 1845-47. By A. H. LAYARD. With Illustrations.

NINEVEH AND BABYLON ; a Narrative of a Second Expedi-
tion to Assyria in 1 849-51. By A. H. LAYARD. With Illustrations.

THREE YEAES' RESIDENCE IN ABYSSINIA, with Travels
in that Country. By MANSFIELD PARKYNS. With Illustrations.

FIVE YEAES IN DAMASCUS, with Teavels in Palmyra,

Lebanon, and among the Giant Cities of Bashan and The Haukan. By
REV. J. L. PORTER. With Illustrations.

THE VOYAGE OF THE ' FOX * IN THE AECTIC SEAS,

and the Discovery of the Fate of Sir John Franklin and his Companions.
By SIR LEOPOLD McCLINTOCK. With Illustrations.

EEMINISCENCES OF ATHENS AND THE MOEEA, during
Travels in Greece. By LORD CARNARVON. With Map.

PEN AND PENCIL SKETCHES IN INDIA. By GENEEAL

MUNDY. With illustrations.

PHILOSOPHY IN SPOET, MADE SCIENCE IN EAENEST :

or, The First Principles of Natural Philosophy explained by the Toys and
Sports of Youth. By DR. PARIS. With Woodcuts.

BLIND PEOPLE ; their Works and Ways. With Lives of some
famous Blind Men. By REV. B. G. JOHNS. With Illustrations.

HOEACE: A New Edition of the Text. Edited by DEAN
MILMAN. With 100 Woodcuts.

THE BOOK OF THE CHUECH. By EOBEET SOUTHEY.

A HANDBOOK FOE YOUNG PAINTEES. By C. E.
LESLIE, R.A. With 24 Illustrations.

A GEOGEAPHICAL HANDBOOK OF FEENS, with Tables
to show their Distribution. By K. M. LYELL. With a Frontispiece.

THE STOEY OF THE LIFE OF LOED BACON. By W.

HEPWORTH DIXON.

PUBLISHED BY MR. MURRAY. 11

A SMALLER DICTIONARY OF THE BIBLE : Its Antiquities,
Geography, Biography, and Natural History. By DR. WM. SMITH. With
Maps and Illustrations.

A SMALLER CLASSICAL DICTIONARY OF MYTHOLOGY,

BIOGRAPHY, AND GEOGRAPHY. By DR. WM. SMITH. With
200 Woodcuts.

A SMALLER DICTIONARY OF GREEK AND ROMAN

ANTIQUITIES. By DR. WM. SMITH. With 200 Woodcuts.

A SMALLER LATIN - ENGLISH DICTIONARY. With a

Dictionary of Proper Names, and Tables of the Roman Calendar, Measures,
Weights, and Moneys. By DR. WM. SMITH.

A SMALLER ENGLISH-LATIN DICTIONARY. By DR. WM.

SMITH.

THE STUDENT'S HUME; An Epitome of the History of

England. By DAVID HUME. Corrected and continued to 1868. With
Woodcuts.

THE STUDENTS CONSTITUTIONAL HISTORY OF ENG-
LAND. By HENRY HALLAM. With the Author's latest Additions.
Edited by Dr. Wm. Smith. \_In the Press.

THE STUDENT'S HISTORY OF THE MIDDLE AGES OF
EUROPE. By HENRY HALLAM. With the Author's Supplemental
Notes. Edited by Dr. Wm. Smith. [In the Press.

THE STUDENT'S HISTORY OF FRANCE. From the

Earliest Times to the Establishment of the Second Empire, 1852.
Wiih Woodcuts.

THE STUDENT'S HISTORY OF ROME. From the Earliest

Times to the Establishment of the Empire. With Chapters on the
History of Literature and Art. By DEAN LIDDELL. With Woodcuts.

THE STUDENT'S GIBBON; An Epitome of the History of
the Decline and Fall of the Roman Empire. By EDWARD GIBBON.
With Woodcuts.

THE STUDENTS HISTORY OF GREECE. From the

Earliest Times to the Roman Conquest. With Chapters on the
History of Literature and Art. By DR. WM. SMITH. With Woodcuts.

THE STUDENTS ANCIENT HISTORY OF THE EAST.

From the Earliest Times to the Conquests of Alexander the Great, including
Egypt, Assyria, Babylonia, Media, Persia, Asia Minor, and Phoenicia. By
PHILIP SMITH, B.A. With Woodcuts.

THE STUDENT'S OLD TESTAMENT HISTORY. From the

Creation to the Return of the Jews from Captivity. With Maps and
Woodcuts.

THE STUDENT'S NEW TESTAMENT HISTORY. With an

Introdui tion, containing the connection of the Old and New Testaments.
With Maps and Woodcuts.

THE STUDENT'S MANUAL OF THE ENGLISH LAN-
GUAGE. By GEORGE P. MARSH. Edited, with additional Chapters and

Notes.

12 LIST OF POPULAR WORKS.

THE STUDENT'S MANUAL OF ENGLISH LITERATURE.

By T. B. SHAW, M.A. Edited, with Notes and Illustrations.

THE STUDENT'S SPECIMENS OF ENGLISH LITERA-
TURE. Selected from the Best Writers. By THOS. B. SHAW, M.A.
Edited, with Additions.

THE STUDENT'S MANUAL OF ANCIENT GEOGRAPHY.

By REV. W. L. BE VAN. With Woodcuts.

THE STUDENT'S MANUAL OF MODERN GEOGRAPHY.

Mathematical, Physical, and Descriptive. By REV. W. L. BEVAN. With
Woodcuts.

THE STUDENTS MANUAL OF MORAL PHILOSOPHY.

With Quotations and References. By WILLIAM FLEMING, D.D.

THE STUDENT'S BLACKSTONE. A Systematic Abridgment

OF THE entire Commentaries. By R. MALCOLM KERR, LL.D.

A PRACTICAL HEBREW GRAMMAR. With the Hebrew

text of Genesis i.-vi. and Psalms i.-vi., Grammatical Analysis aud Vocabulary
By REV. STANLEY LEATHES.

Nine Shillings.

THE CONNECTION OF THE PHYSICAL SCIENCES. By

MARY SOMERVILLE. With Woodcuts.

PHYSICAL GEOGRAPHY. By MARY SOMERVILLE.

Revised by H. W. Bates. With Portrait.

THE STUDENT'S ELEMENTS OF GEOLOGY. By SIR

CHARLES LYELL. With 600 Woodcuts.

A MANUAL OF SCIENTIFIC INQUIRY. By Various Writers.
Edited by SIR J. F. HERSCHEL and REV. ROBERT MAIN.

POETICAL WORKS OF LORD BYRON. With Notes, Illus-

tratiuns, and Portrait.

LIFE OF LORD BYRON ; with his Letters and Journals. By
THOMAS MOORE. With Portraits.

ARCHBISHOP BECKET; A BIOGRAPHY. By CANON

ROBERTSON, M.A. With Illustrations.

PICTURES OF THE CHINESE, DRAWN BY THEMSELVES.

Described by REV. R. H. COBBOLD. With 34 Illustrations.

THE ENGLISH BATTLES AND SIEGES OF THE PENIN-
SULAR WAR. By SIR WILLIAM NAPIER. With Portrait.

THE YOUNG OFFICER'S COMPANION; or, Essays on

Military Duties and (Qualities : with Illustrations from History. By
LORD DE ROS.

DOG-BREAKING; the most Expeditious, Certain, and Easy
Method, whether great Excellence or only Mediocrity be required. With
a Few Hints for those who Love the Dog and the Gun. By GENERAL
HUTCHINSON. With Woodcuts.

LIST OF SCHOOL CLASSICS. 13

By DR. WILLIAM SMITH.
PKINCIPIA LATINA, Part I. A Fikst Latin Course. A

Grammar, Delectus, and Exercise Book with Vocabularies. 13th Edition.
3s. 6d.

%* This Edition contains the Accidence arranged for the ' Public School Latin
Primer.'

PRINCIPIA LATINA, Part II. Latin Eeading Book. An

Introduction to Ancient Mythology, Geography, Roman Antiquities, and
History. With Notes and a Dictionary. 3s. 6d.

PRINCIPIA LATINA, Part III. Latin Poetry. 1. Easy

Hexameters and Pentameters. 2. Eclogaj Ovidianaj. 3. Prosody and Metre.
4. First Latin Verse Book. 3s. 6d.

PRINCIPIA LATINA, Part IV. Latin Prose Composition.

Rules of Syntax, with Examples, Explanations of Synonyms, and Exercises
on the Syntax. 3s. 6d.

PRINCIPIA LATINA, Part V. Short Tales and Anecdotes
from Ancient History, for Translation into Latin Prose. 3s.

A LATIN-ENGLISH VOCABULARY, with a Latin-English
Dictionary to Phaedrus, Cornelius Nepos, and Caesar's 'Gallic War.' 3s. 6d.

THE STUDENTS LATIN GRAMMAR. By WM. SMITH,

D.C.L., and THE0PH1LUS D. HALL. 6s.

A SMALLER LATIN GRAMMAR. Abridged from the above
Work. 3s. Qd.

INITIA GR.ECA, Part I. A First Greek Course, containing
Grammar, Delectus, Exercise Book, and Vocabularies. By DR. WM. SMITH.
3s. 6(i.

INITIA GR^ECA, Part II. A Reading Book ; containing short
Tales, Anecdotes, Fables, Mythology, and Grecian History. With a Lexicon.
3s. 6d.

INITIA GRiECA, Part III. Greek Prose Composition; con-
taining the Rules of Syntax, with copious Examples and Exercises. 3s. 6d.

THE STUDENT'S GREEK GRAMMAR. By PROFESSOR

CUR ITUS, and WM. SMITH, D.C.L. 6s.

A SMALLER GREEK GRAMMAR. Abridged from the above

Work. 3s. 6cZ.

PRINCIPIA GR^ECA. A First Greek Course. A Grammar,

Delectus, and Exercise Book, with Vocabularies. By H. E. HUTTON, M.A
3s. 6d.

MATTHIAS GREEK GRAMMAR. Abridged by BLOM-
F1ELD. Revised and enlarged, by E. S. Crooke, B.A. 4s.

KING EDWARD VI.'S FIRST LATIN BOOK; including
a Short Syntax and Prosody with an English Translation. 2s. 6d.

KING EDWARD VI.'S LATIN GRAMMAR. 3s. 6d.

ENGLISH NOTES FOR LATIN ELEGIACS; designed for
Early Proficients in the Art of Latjn Versification. By REV. W. OXENHAM.
3s. 6d.

14 LIST OF HANDBOOKS FOR TRAVELLERS.

THE CONTINENT, &c.
HANDBOOK— TRAVEL TALK, — English, French, German,

and Italian. 3s. 6d.

HANDBOOK — NORTH GERMANY, Holland, Belgium,

Prussia, and the Rhine to Switzerland. With Map and Plans.

HANDBOOK — SOUTH GERMANY, The Tyrol, Bavaria,
Austria, Salzburg, Sttria, Hungary, and The Danube, from Ulm to the
Black Sea. With Map and Plans. 10s.

HANDBOOK — SWITZERLAND, The Alps of Savoy and

Piedmont. With Maps and Plans. 10s.

HANDBOOK — FRANCE, Normandy, Brittany, The French
Alps, Dauphlne, Provence, and the Pyrenees, With Maps. 12s.

HANDBOOK— PARIS AND ITS ENVIRONS. With Map and

Plans. 3s. 6d.

*** Murray's Plan of Paris. 3s. 6d.
HANDBOOK — CORSICA AND SARDINIA. With Map. 4s.
HANDBOOK — SPAIN, Madrid, The Castile.?, The Basque

Provinces, Leon, The Asturias, Galicia, Estremadura, Andalusia,
Ronda, Granada, Murcia, Valencia, Catalonia, Aragon, Navarre,
The Balearic Islands, &c, &c. With Maps. 2 vols. 24s.

HANDBOOK — PORTUGAL, Lisbon, Porto, Cintra, Mafra,

&c. With Map. 9s.

HANDBOOK — NORTH ITALY, Piedmont, Nice, Lombardy,

Venice, Parma, Modena, and Romagna. With Map and Plans. 12s.

HANDBOOK — CENTRAL ITALY, Tuscany, Florence, Lucca,
Umbria, The Marches, and the Patrimony of St. Peter. With Map.
10s.

HANDBOOK — ROME AND ITS ENVIRONS. With Map and

Plans. 9s.

HANDBOOK — SOUTH ITALY, Two Sicilies, Naples, Pompeh,

Herculaneum, Vesuvius, Abruzzi, &c. With Map. 10s.

HANDBOOK — SICILY, Palermo, Messina, Catania, Syracuse,
Etna, and the Ruins of the Greek Temples. With Map. 12s.

HANDBOOK — E G Y P T, The Nile, Alexandria, Cairo,
Thebes, and the Overland Route to India. With Map. 15s.

HANDBOOK — GREECE, The Ionian Islands, Athens,
Albania , Thessaly, and Macedonia. With Map.

HANDBOOK — Constantinople, The Bosphoris, Dardanelles,

Brousa, and Plain of Troy, with General Hints for Travellers in Turkey,
&c. With Map

LIST OF HANDBOOKS FOR TRAVELLERS. 15

HANDBOOK — DENMABK, Norway, Sweden, and Iceland.

With Map and Plans.

HANDBOOK — KUSSIA, St. Petersburg, Moscow, Finland, &c.

With Map. 15s.

HANDBOOK— INDIA, Bombay and Madras. With Map. 2 vols.

12s. each.

HANDBOOK — HOLY LAND, Syria, Palestine, Sinai, Edom,

and the Syrian Deserts. With Map. 2 vols. 24s.

KNAPSACK GUIDES FOR TRAVELLERS.
KNAPSACK GUIDE TO SWITZEKLAND. With Plans. 5s.
KNAPSACK GUIDE TO NOEWAY. With Map. 6s.
KNAPSACK GUIDE TO ITALY. With Plans. 6s.
KNAPSACK GUIDE TO THE TYROL. With Plans. 6s.

ENGLAND AND WALES.

HANDBOOK — LONDON AS IT IS. With Map and Plans.

3S. 6d.

HANDBOOK — ESSEX, CAMBEIDGE, SUFFOLK, AND
NORFOLK — Chelmsford, Colchester, Maldon, Cambridge, Ely, New-
market, Bury, Ipswich, Woodbridge, Felixstowe, Lowestoft, Norwich
Yarmouth, Cromer, &c. With Maps and Plans. 12s.

HANDBOOK — KENT AND SUSSEX — Canterbury, Dover,
Ramsgate, Rochester, Chatham, Brighton, Chichester, Worthing, Has-
tings, Lewes, Arundel. With Map. lus.

HANDBOOK — SUEEEY AND HANTS— Kingston, Croydon,
Reigate, Guildford, Dorking, Boxhill, Winchester, Southampton, Ports-
mouth, and The Isle of Wight. With Map. IDs.

HANDBOOK — BE EKS, BUCKS, AND OXON — Windsor,
Eton, Reading, Aylesbury, Henley, Oxford, and the Thames. With
Map. 7a. 6d.

HANDBOOK — WILTS, DOKSET, AND SOMEESET —

Salisbury, Chippenham, Weymouth, Sherborne, Wells, Bath, Bristol,
Taunton, &c. With Map. 10s.

HANDBOOK — DEVON AND C 0 E N WALL — Exeter,

Ilfracombe, Linton, Sidmouth, Dawlish, Teignmouth, Plymouth, Devon-
port, Torquay, Launceston, Penzance, Falmouth, The Lizard, Land's
End, &c. With Map. 10s.

16 LIST OF HANDBOOKS FOR TRAVELLERS.

HANDBOOK — GLOUCESTER, HEREFORD, AND

WORCESTER — Cirencester, Cheltenham, Stroud, Tewkesbury, Leo-
minster, Ross, Malvern, Kidderminster, Dudley, Bromsgrove, Evesham.
With Map. 6s. 6d.

HANDBOOK — N 0 R T H W A L E S — Bangor, Carnarvon,

Beaumaris, Snowdon, Conway, &c. With Map. 6s. 6d.

HANDBOOK — SOUTH WALES — Monmouth, Carmarthen,

Tenby, Swansea, and the Wye, &c. With Map. 7s.

HANDBOOK — DERBY, NOTTS, LEICESTER, AN^»

STAFFORD — Matlock, Bakewell, Chatsworth, The Peak, Bu>
Hardwick, Dove Dale, Ashborne. Southwell, Mansfield, Retf»j -o,
Burton, Belvoir, Melton Mowbray, Wolverhampton, Litchfield,
Walsall, Tamworth. With Map. 7s. 6d.

HANDBOOK — SHROPSHIRE, CHESHIRE, AND LANCA-
SHIRE]— Shrewsbury, Ludlow, Bridgnorth, Oswestry, Chester, Crewe,
Alderley, Stockport, Birkenhead, Warrington, Bury, Manchester,
Liverpool, Burnlei, Clttheroe, Bolton, Blackburn, Wigan, Preston,
Rochdale, Lancaster, Southport, Blackpool, &c. With Map. 10s.

HANDBOOK — YORKSHIRE — Donoaster, Hull, Selby,

Beverley, Scarborough, Whitby, Harrogate, Ripon, Leeds, Wakefield,
Bradford, Halifax, Huddersfield, Sheffield. With Map. 12s.

HANDBOOK — DURHAM AND NORTHUMBERLAND —

Newcastle, Darllngton, Bishop Auckland, Stockton, Hartlepool, Sun-
derland, Shields, Berwick, Tynemouth, Alnwick. With Map. 9s.

HANDBOOK — WESTMORLAND AND CUMBERLAND —

Lancaster, Furness Abbey, Ambleside, Kendal, Windermere, Coniston,
Keswick, Grasmere, Carlisle, Cockermouth, Penrith, Appleby. With
Map. 6s.

*** Murray's Map op the Lake District, 3s. 6d.

HANDBOOK — SCOTLAND — Edinburgh, Melrose, Kelso,
Glasgow, Dumfries, Ayk, Stirling, Arrax, The Clyde. Oban, Inverary,
I/Och Lomond, Loch Katrine and Trosachs, Caledonian Canal, Inver-
ness, Perth, Dundee, Aberdeen, Braemar, Skye, Caithness, Ross, and
Sutherland. With Maps and Plans. 9s.

HANDBOOK — IRELAND— Dublin, Belfast, Donegal, Gal-
way, Wexford, Cork, Limerick, Waterford, Killarney, Munster.
With Map. 12s.

JOHN MURRAY, 50a, ALBEMARLE STREET.

LONDON: PRINTED BY WILLIAM CLOWES AND SONS, STAMFORD STREET,

AND CHARING CROSS.

>

y

Nratffnuf

SiiSlE

flilHi

■ ;:;' - m h

■.-■■'■''

■■-'■'''■■■"■■■'

* H; % M&l nSUVlMllllfat'

- ■■"::■; ■"";■■"■

■■■■--•:■.,.

sShHmSS

'.■.'.."■■'•'■'-:■-.■■".'">-■ 1; :;-'t ;"■'""

• '■'••'''-■*

SiSa lMfi§S ^^

'■'•::^.'::- ;.-.!^r-.':- :

■an

**■

MaMaaWKa

MHHh

■mmw

' '■■■■•-'.■■'■!',"'•

'-■■-'■:• .':■■::'-. :-i',:-..,:-:;

:i- '■•■■:.•'.'•■'• ::

SjmB