The Descent of Man and Selection in Relation to Sex_ Vol. II (1st Edition) Charles Darwin 1424 downloads.pdf

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Selection in Relation to Sex, Vol. II (1st Edition)

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Title: The Descent of Man and Selection in Relation to Sex, Vol. II (1st
Edition)

Author: Charles Darwin

Release date: June 25, 2011 [eBook #36520]

Language: English

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*** START OF THE PROJECT GUTENBERG EBOOK THE DESCENT
OF MAN AND SELECTION IN RELATION TO SEX, VOL. II (1ST
EDITION) ***

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E-text prepared by Steven Gibbs, Turgut Dincer,
and the Online Distributed Proofreading Team
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Note: Project Gutenberg has Volume I of this book. See
http://www.gutenberg.org/ebooks/34967

THE

DESCENT OF MAN,
AND

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SELECTION IN RELATION TO SEX.

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

IN TWO VOLUMES.—Vol. II.
WITH ILLUSTRATIONS.

LONDON:
JOHN MURRAY, ALBEMARLE STREET.
1871.
[The right of Translation is reserved.]

ERRATA.

VOL. I.

Page line For read
27 13 kaolo koala.
31 6 prostratica prostatica.
59, note86 2 speech species.
74, note107 — Browne Brown.

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118, note167 — Vol. I. Vol. II.
insert ‘Proc. Royal
128, note184 4 Before vol. xiv.
Soc.
208 2 prostratica prostatica.
322 5 Actineæ Actiniæ.
324 30 land-shells land-snails.
330 16 figs. 4 and 5 figs. 4, 5, and 6.
334 17 Birgos Birgus.
339 8 attractions attentions.
341 3 dragon-flys dragon-flies.
378 17 Typhæus Typhœus.
384 31 tesselatum tessellatum.
397 9 Hypopira Hypopyra.
405 21 Acrœidæ Acræidæ.

VOL. II.

32 30 chamelion chameleon.
115 4 mail male.
178 23 Chloehaga Chloephaga.
227, note281 — Ramphaston Ramphastos.
240, note289 — Mr. H. Brown Mr. R. Brown.
240, note290 2 elephus elaphus.
242 14 walruses narwhals.
339 27 Durfur Darfur.

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CONTENTS.

PART II.

SEXUAL SELECTION—continued.

CHAPTER XII.
Secondary Sexual Characters of Fishes, Amphibians, and Reptiles.
Fishes: Courtship and battles of the males—Larger size of the females—Males, bright colours
and ornamental appendages; other strange characters—Colours and appendages acquired
by the males during the breeding-season alone—Fishes with both sexes brilliantly
coloured—Protective colours—The less conspicuous colours of the female cannot be
accounted for on the principle of protection—Male fishes building nests, and taking
charge of the ova and young. Amphibians: Differences in structure and colour between the
sexes—Vocal organs. Reptiles: Chelonians—Crocodiles—Snakes, colours in some cases
protective—Lizards, battles of—Ornamental appendages—Strange differences in
structure between the sexes—Colours—Sexual differences almost as great as with birds 1-37

CHAPTER XIII.
Secondary Sexual Characters of Birds.
Sexual differences—Law of battle—Special weapons—Vocal organs—Instrumental music—
Love-antics and dances—Decorations, permanent and seasonal—Double and single 38-
annual moults—Display of ornaments by the males 98

CHAPTER XIV.
Birds—continued
Choice exerted by the female—Length of courtship—Unpaired birds—Mental qualities and
taste for the beautiful—Preference or antipathy shewn by the female for particular males
—Variability of birds—Variations sometimes abrupt—Laws of variation—Formation of 99-
ocelli—Gradations of character—Case of Peacock, Argus pheasant, and Urosticte 153

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CHAPTER XV.
Birds—continued.
Discussion why the males alone of some species, and both sexes of other species, are brightly
coloured—On sexually-limited inheritance, as applied to various structures and to
brightly-coloured plumage—Nidification in relation to colour—Loss of nuptial plumage 154-
during the winter 182

CHAPTER XVI.
Birds—continued.
The immature plumage in relation to the character of the plumage in both sexes when adult—
Six classes of cases—Sexual differences between the males of closely-allied or
representative species—The female assuming the characters of the male—Plumage of the
young in relation to the summer and winter plumage of the adults—On the increase of
beauty in the Birds of the World—Protective colouring—Conspicuously-coloured birds— 183-
Novelty appreciated—Summary of the four chapters on birds 238

CHAPTER XVII.
Secondary Sexual Characters of Mammals.
The law of battle—Special weapons, confined to the males—Cause of absence of weapons in
the female—Weapons common to both sexes, yet primarily acquired by the male—Other
uses of such weapons—Their high importance—Greater size of the male—Means of 239-
defence—On the preference shewn by either sex in the pairing of quadrupeds 273

CHAPTER XVIII.
Secondary Sexual Characters of Mammals.—continued.
Voice—Remarkable sexual peculiarities in seals—Odour—Development of the hair—Colour
of the hair and skin—Anomalous case of the female being more ornamented than the male
—Colour and ornaments due to sexual selection—Colour acquired for the sake of
protection—Colour, though common to both sexes, often due to sexual selection—On the
disappearance of spots and stripes in adult quadrupeds—On the colours and ornaments of 274-
the Quadrumana—Summary 315

CHAPTER XIX.
Secondary Sexual Characters of Mammals.—continued.
Differences between man and woman—Causes of such differences and of certain characters
common to both sexes—Law of battle—Differences in mental powers—and voice—On
the influence of beauty in determining the marriages of mankind—Attention paid by
savages to ornaments—Their ideas of beauty in woman—The tendency to exaggerate 316-
each natural peculiarity 354

CHAPTER XX.
Secondary Sexual Characters of Man—continued.

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SEXUAL SELECTION.

CHAPTER XII.

Secondary Sexual Characters of Fishes, Amphibians, and Reptiles.
Fishes: Courtship and battles of the males—Larger size of the females—Males, bright colours and
ornamental appendages; other strange characters—Colours and appendages acquired by the
males during the breeding-season alone—Fishes with both sexes brilliantly coloured—
Protective colours—The less conspicuous colours of the female cannot be accounted for on the
principle of protection—Male fishes building nests, and taking charge of the ova and young.
Amphibians: Differences in structure and colour between the sexes—Vocal organs. Reptiles:
Chelonians—Crocodiles—Snakes, colours in some cases protective—Lizards, battles of—
Ornamental appendages— Strange differences in structure between the sexes—Colours—
Sexual differences almost as great as with birds.

We have now arrived at the great sub-kingdom of the Vertebrata, and will
commence with the lowest class, namely Fishes. The males of
Plagiostomous fishes (sharks, rays) and of Chimæroid fishes are provided
with claspers which serve to retain the female, like the various structures
possessed by so many of the lower animals. Besides the claspers, the males
of many rays have clusters of strong sharp spines on their heads, and
several rows along “the upper outer surface of their pectoral fins.” These
are present in the males of some species, which have the other parts of their
bodies smooth. They are only temporarily developed during the breeding-
season; and Dr. Günther suspects that they are brought into action as
prehensile organs by the doubling inwards and downwards of the two sides
of the body. It is a remarkable fact that the females and not the males of
some species, as of Raia clavata, have their backs studded with large hook-
formed spines.1
Owing to the element which fishes inhabit, little is known about their
courtship, and not much about their battles. The male stickleback
(Gasterosteus leiurus) has been described as “mad with delight” when the

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female comes out of her hiding-place and surveys the nest which he has
made for her. “He darts round her in every direction, then to his
accumulated materials for the nest, then back again in an instant; and as she
does not advance he endeavours to push her with his snout, and then tries to
pull her by the tail and side-spine to the nest.”2 The males are said to be
polygamists;3 they are extraordinarily bold and pugnacious, whilst “the
females are quite pacific.” Their battles are at times desperate; “for these
puny combatants fasten tight on each other for several seconds, tumbling
over and over again, until their strength appears completely exhausted.”
With the rough-tailed stickleback (G. trachurus) the males whilst fighting
swim round and round each other, biting and endeavouring to pierce each
other with their raised lateral spines. The same writer adds,4 “the bite of
these little furies is very severe. They also use their lateral spines with such
fatal effect, that I have seen one during a battle absolutely rip his opponent
quite open, so that he sank to the bottom and died.” When a fish is
conquered, “his gallant bearing forsakes him; his gay colours fade away;
and he hides his disgrace among his peaceable companions, but is for some
time the constant object of his conqueror’s persecution.”
The male salmon is as pugnacious as the little stickleback; and so is the
male trout, as I hear from Dr. Günther. Mr. Shaw saw a violent contest
between two male salmons which lasted the whole day; and Mr. R. Buist,
Superintendent of Fisheries, informs me that he has often watched from the
bridge at Perth the males driving away their rivals whilst the females were
spawning. The males “are constantly fighting and tearing each other on the
spawning-beds, and many so injure each other as to cause the death of
numbers, many being seen swimming near the banks of the river in a state
of exhaustion, and apparently in a dying state.”5 The keeper of the
Stormontfield breeding-ponds visited, as Mr. Buist informs me, in June,
1868, the northern Tyne, and found about 300 dead salmon, all of which
with one exception were males; and he was convinced that they had lost
their lives by fighting.

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Fig. 26. Head of male of common salmon (Salmo salar) during the
breeding-season.
[This drawing, as well as all the others in the present chapter, have been
executed by the well-known artist, Mr. G. Ford, under the kind
superintendence of Dr. Günther, from specimens in the British Museum.]

The most curious point about the male salmon is that during the breeding-
season, besides a slight change in colour, “the lower jaw elongates, and a
cartilaginous projection turns upwards from the point, which, when the jaws
are closed, occupies a deep cavity between the intermaxillary bones of the
upper jaw.”6 (Figs. 26 and 27.) In our salmon this change of structure lasts
only during the breeding-season; but in the Salmo lycaodon of N.W.
America the change, as Mr. J. K. Lord7 believes, is permanent and best
marked in the older males which have previously ascended the rivers. In
these old males the jaws become developed into immense hook-like
projections, and the teeth grow into regular fangs, often more than half an
inch in length. With the European salmon, according to Mr. Lloyd,8 the
temporary hook-like structure serves to strengthen and protect the jaws,
when one male charges another with wonderful violence; but the greatly
developed teeth of the male American salmon may be compared with the
tusks of many male mammals, and they indicate an offensive rather than a
protective purpose.

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Fig. 27. Head of female salmon.

The salmon is not the only fish in which the teeth differ in the two sexes.
This is the case with many rays. In the thornback (Raia clavata) the adult
male has sharp, pointed teeth, directed backwards, whilst those of the
female are broad and flat, forming a pavement; so that these teeth differ in
the two sexes of the same species more than is usual in distinct genera of
the same family. The teeth of the male become sharp only when he is adult:
whilst young they are broad and flat like those of the female. As so
frequently occurs with secondary sexual characters, both sexes of some
species of rays, for instance R. batis, possess, when adult, sharp, pointed
teeth; and here a character, proper to and primarily gained by the male,
appears to have been transmitted to the offspring of both sexes. The teeth
are likewise pointed in both sexes of R. maculata, but only when
completely adult; the males acquiring them at an earlier age than the
females. We shall hereafter meet with analogous cases with certain birds, in
which the male acquires the plumage common to both adult sexes, at a
somewhat earlier age than the female. With other species of rays the males
even when old never possess sharp teeth, and consequently both sexes when
adult are provided with broad, flat teeth like those of the young, and of the
mature females of the above-mentioned species.9 As the rays are bold,
strong and voracious fishes, we may suspect that the males require their
sharp teeth for fighting with their rivals; but as they possess many parts
modified and adapted for the prehension of the female, it is possible that
their teeth may be used for this purpose.

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In regard to size, M. Carbonnier10 maintains that with almost all fishes the
female is larger than the male; and Dr. Günther does not know of a single
instance in which the male is actually larger than the female. With some
Cyprinodonts the male is not even half as large as the female. As with many
kinds of fishes the males habitually fight together; it is surprising that they
have not generally become through the effects of sexual selection larger and
stronger than the females. The males suffer from their small size, for
according to M. Carbonnier they are liable to be devoured by the females of
their own species when carnivorous, and no doubt by other species.
Increased size must be in some manner of more importance to the females,
than strength and size are to the males for fighting with other males; and
this perhaps is to allow of the production of a vast number of ova.

Fig. 28. Callionymus lyra. Upper figure, male; lower figure, female.

In many species the male alone is ornamented with bright colours; or these
are much brighter in the male than the female. The male, also, is sometimes
provided with appendages which appear to be of no more use to him for the
ordinary purposes of life than are the tail-feathers to the peacock. I am
indebted for most of the following facts to the great kindness of Dr.

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Günther. There is reason to suspect that many tropical fishes differ sexually
in colour and structure; and there are some striking cases with our British
fishes. The male Callionymus lyra has been called the gemmeous dragonet
“from its brilliant gem-like colours.” When freshly taken from the sea the
body is yellow of various shades, striped and spotted with vivid blue on the
head; the dorsal fins are pale brown with dark longitudinal bands; the
ventral, caudal and anal fins being bluish-black. The female, or sordid
dragonet, was considered by Linnæus and by many subsequent naturalists
as a distinct species; it is of a dingy reddish-brown, with the dorsal fin
brown and the other fins white. The sexes differ also in the proportional size
of the head and mouth, and in the position of the eyes;11 but the most
striking difference is the extraordinary elongation in the male (fig. 28) of
the dorsal fin. The young males resemble in structure and colour the adult
females. Throughout the genus Callionymus,12 the male is generally much
more brightly spotted than the female, and in several species, not only the
dorsal, but the anal fin of the male is much elongated.
The male of the Cottus scorpius, or sea-scorpion, is more slender and
smaller than the female. There is also a great difference in colour between
them. It is difficult, as Mr. Lloyd13 remarks, “for any one, who has not seen
this fish during the spawning-season, when its hues are brightest, to
conceive the admixture of brilliant colours with which it, in other respects
so ill-favoured, is at that time adorned.” Both sexes of the Labrus mixtus,
although very different in colour, are beautiful; the male being orange with
bright-blue stripes, and the female bright-red with some black spots on the
back.
In the very distinct family of the Cyprinodontidæ—inhabitants of the fresh
waters of foreign lands—the sexes sometimes differ much in various
characters. In the male of the Mollienesia petenensis,14 the dorsal fin is
greatly developed and is marked with a row of large, round, ocellated,
bright-coloured spots; whilst the same fin in the female is smaller, of a
different shape, and marked only with irregularly-curved brown spots. In
the male the basal margin of the anal fin is also a little produced and dark-
coloured. In the male of an allied form, the Xiphophorus Hellerii (fig. 29),
the inferior margin of the anal fin is developed into a long filament, which
is striped, as I hear from Dr. Günther, with bright colours. This filament
does not contain any muscles, and apparently cannot be of any direct use to

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the fish. As in the case of the Callionymus, the males whilst young resemble
in colour and structure the adult females. Sexual differences such as these
may be strictly compared with those which are so frequent with
gallinaceous birds.15

Fig. 29. Xiphophorus Hellerii. Upper figure, male; lower figure, female.

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Fig. 30. Plecostomus barbatus. Upper figure, head of male; lower figure,
female.

In a siluroid fish, inhabiting the fresh waters of South America, namely the
Plecostomus barbatus16 (fig. 30), the male has its mouth and interoperculum
fringed with a beard of stiff hairs, of which the female shews hardly a trace.
These hairs are of the nature of scales. In another species of the same genus,
soft flexible tentacles project from the front part of the head of the male,
which are absent in the female. These tentacles are prolongations of the true
skin, and therefore are not homologous with the stiff hairs of the former
species; but it can hardly be doubted that both serve the same purpose.
What this purpose may be it is difficult to conjecture; ornament does not
here seem probable, but we can hardly suppose that stiff hairs and flexible
filaments can be useful in any ordinary way to the males alone. The
Monacanthus scopas, which was shewn to me in the British Museum by Dr.
Günther, presents a nearly analogous case. The male has a cluster of stiff,
straight spines, like those of a comb, on the sides of the tail; and these in a
specimen six inches long were nearly an inch and a half in length; the
female has on the same place a cluster of bristles, which may be compared

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with those of a tooth-brush. In another species, the M. peronii, the male has
a brush like that possessed by the female of the last species, whilst the sides
of the tail in the female are smooth. In some other species the same part of
the tail can be perceived to be a little roughened in the male and perfectly
smooth in the female; and lastly in others, both sexes have smooth sides. In
that strange monster, the Chimæra monstrosa, the male has a hook-shaped
bone on the top of the head, directed forwards, with its rounded end covered
with sharp spines; in the female “this crown is altogether absent,” but what
its use may be is utterly unknown.17
The structures as yet referred to are permanent in the male after he has
arrived at maturity; but with some Blennies and in another allied genus18 a
crest is developed on the head of the male only during the breeding-season,
and their bodies at the same time become more brightly-coloured. There can
be little doubt that this crest serves as a temporary sexual ornament, for the
female does not exhibit a trace of it. In other species of the same genus both
sexes possess a crest, and in at least one species neither sex is thus
provided. In this case and in that of the Monacanthus, we have good
instances to how great an extent the sexual characters of closely-allied
forms may differ. In many of the Chromidæ, for instance in Geophagus and
especially in Cichla, the males, as I hear from Professor Agassiz,19 have a
conspicuous protuberance on the forehead, which is wholly wanting in the
females and in the young males. Professor Agassiz adds, “I have often
observed these fishes at the time of spawning when the protuberance is
largest, and at other seasons when it is totally wanting and the two sexes
shew no difference whatever in the outline of the profile of the head. I never
could ascertain that it subserves any special function, and the Indians on the
Amazon know nothing about its use.” These protuberances in their
periodical appearance resemble the fleshy caruncles on the heads of certain
birds; but whether they serve as ornaments must remain at present doubtful.
The males of those fishes, which differ permanently in colour from the
females, often become more brilliant, as I hear from Professor Agassiz and
Dr. Günther, during the breeding-season. This is likewise the case with a
multitude of fishes, the sexes of which at all other seasons of the year are
identical in colour. The tench, roach, and perch may be given as instances.
The male salmon at this season is “marked on the cheeks with orange-
coloured stripes, which give it the appearance of a Labrus, and the body

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partakes of a golden-orange tinge. The females are dark in colour, and are
commonly called black-fish.”20 An analogous and even greater change takes
place with the Salmo eriox or bull-trout; the males of the char (S. umbla) are
likewise at this season rather brighter in colour than the females.21 The
colours of the pike (Esox reticulatus) of the United States, especially of the
male, become, during the breeding-season, exceedingly intense, brilliant,
and iridescent.22 Another striking instance out of many is afforded by the
male stickleback (Gasterosteus leiurus), which is described by Mr.
Warington,23 as being then “beautiful beyond description.” The back and
eyes of the female are simply brown, and the belly white. The eyes of the
male, on the other hand, are “of the most splendid green, having a metallic
lustre like the green feathers of some humming-birds. The throat and belly
are of a bright crimson, the back of an ashy-green, and the whole fish
appears as though it were somewhat translucent and glowed with an internal
incandescence.” After the breeding-season these colours all change, the
throat and belly become of a paler red, the back more green, and the
glowing tints subside.
That with fishes there exists some close relation between their colours and
their sexual functions we can clearly see;—firstly, from the adult males of
certain species being differently coloured from the females, and often much
more brilliantly;—secondly, from these same males, whilst immature,
resembling the mature females;—and, lastly, from the males, even of those
species which at all other times of the year are identical in colour with the
females, often acquiring brilliant tints during the spawning-season. We
know that the males are ardent in their courtship and sometimes fight
desperately together. If we may assume that the females have the power of
exerting a choice and of selecting the more highly-ornamented males, all
the above facts become intelligible through the principle of sexual selection.
On the other hand, if the females habitually deposited and left their ova to
be fertilised by the first male which chanced to approach, this fact would be
fatal to the efficiency of sexual selection; for there could be no choice of a
partner. But, as far as is known, the female never willingly spawns except in
the close presence of a male, and the male never fertilises the ova except in
the close presence of a female. It is obviously difficult to obtain direct
evidence with respect to female fishes selecting their partners. An excellent
observer,24 who carefully watched the spawning of minnows (Cyprinus

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phoxinus), remarks that owing to the males, which were ten times as
numerous as the females, crowding closely round them, he could “speak
only doubtfully on their operations. When a female came among a number
of males they immediately pursued her; if she was not ready for shedding
her spawn, she made a precipitate retreat; but if she was ready, she came
boldly in among them, and was immediately pressed closely by a male on
each side; and when they had been in that situation a short time, were
superseded by other two, who wedged themselves in between them and the
female, who appeared to treat all her lovers with the same kindness.”
Notwithstanding this last statement, I cannot, from the several previous
considerations, give up the belief that the males which are the most
attractive to the females, from their brighter colours or other ornaments, are
commonly preferred by them; and that the males have thus been rendered
more beautiful in the course of ages.
We have next to inquire whether this view can be extended, through the law
of the equal transmission of characters to both sexes, to those groups in
which the males and females are brilliant in the same or nearly the same
degree and manner. In such a genus as Labrus, which includes some of the
most splendid fishes in the world, for instance, the Peacock Labrus (L.
pavo), described,25 with pardonable exaggeration, as formed of polished
scales of gold encrusting lapis-lazuli, rubies, sapphires, emeralds and
amethysts, we may, with much probability, accept this belief; for we have
seen that the sexes in at least one species differ greatly in colour. With some
fishes, as with many of the lowest animals, splendid colours may be the
direct result of the nature of their tissues and of the surrounding conditions,
without any aid from selection. The goldfish (Cyprinus auratus), judging
from the analogy of the golden variety of the common carp, is, perhaps, a
case in point, as it may owe its splendid colours to a single abrupt variation,
due to the conditions to which this fish has been subjected under
confinement. It is, however, more probable that these colours have been
intensified through artificial selection, as this species has been carefully
bred in China from a remote period.26 Under natural conditions it does not
seem probable that beings so highly organised as fishes, and which live
under such complex relations, should become brilliantly coloured without
suffering some evil or receiving some benefit from so great a change, and
consequently without the intervention of natural selection.

Page 21

What, then, must we conclude in regard to the many fishes, both sexes of
which are splendidly coloured? Mr. Wallace27 believes that the species
which frequent reefs, where corals and other brightly-coloured organisms
abound, are brightly coloured in order to escape detection by their enemies;
but according to my recollection they were thus rendered highly
conspicuous. In the freshwaters of the Tropics there are no brilliantly-
coloured corals or other organisms for the fishes to resemble; yet many
species in the Amazons are beautifully coloured, and many of the
carnivorous Cyprinidæ in India are ornamented with “bright longitudinal
lines of various tints.”28 Mr. M’Clelland, in describing these fishes goes so
far as to suppose that “the peculiar brilliancy of their colours” serves as “a
better mark for kingfishers, terns, and other birds which are destined to
keep the number of these fishes in check;” but at the present day few
naturalists will admit that any animal has been made conspicuous as an aid
to its own destruction. It is possible that certain fishes may have been
rendered conspicuous in order to warn birds and beasts of prey (as
explained when treating of caterpillars) that they were unpalatable; but it is
not, I believe, known that any fish, at least any freshwater fish, is rejected
from being distasteful to fish-devouring animals. On the whole, the most
probable view in regard to the fishes, of which both sexes are brilliantly
coloured, is that their colours have been acquired by the males as a sexual
ornament, and have been transferred in an equal or nearly equal degree to
the other sex.
We have now to consider whether, when the male differs in a marked
manner from the female in colour or in other ornaments, he alone has been
modified, with the variations inherited only by his male offspring; or
whether the female has been specially modified and rendered inconspicuous
for the sake of protection, such modifications being inherited only by the
females. It is impossible to doubt that colour has been acquired by many
fishes as a protection: no one can behold the speckled upper surface of a
flounder, and overlook its resemblance to the sandy bed of the sea on which
it lives. One of the most striking instances ever recorded of an animal
gaining protection by its colour (as far as can be judged in preserved
specimens) and by its form, is that given by Dr. Günther29 of a pipefish,
which, with its reddish streaming filaments, is hardly distinguishable from
the sea-weed to which it clings with its prehensile tail. But the question now

Page 22

under consideration is whether the females alone have been modified for
this object. Fishes offer valuable evidence on this head. We can see that one
sex will not be modified through natural selection for the sake of protection
more than the other, supposing both to vary, unless one sex is exposed for a
longer period to danger, or has less power of escaping from such danger
than the other sex; and it does not appear that with fishes the sexes differ in
these respects. As far as there is any difference, the males, from being
generally of smaller size, and from wandering more about, are exposed to
greater danger than the females; and yet, when the sexes differ, the males
are almost always the most conspicuously coloured. The ova are fertilised
immediately after being deposited, and when this process lasts for several
days, as in the case of the salmon,30 the female, during the whole time, is
attended by the male. After the ova are fertilised they are, in most cases, left
unprotected by both parents, so that the males and females, as far as
oviposition is concerned, are equally exposed to danger, and both are
equally important for the production of fertile ova; consequently the more
or less brightly-coloured individuals of either sex would be equally liable to
be destroyed or preserved, and both would have an equal influence on the
colours of their offspring or the race.
Certain fishes, belonging to several families, make nests; and some of these
fishes take care of their young when hatched. Both sexes of the brightly-
coloured Crenilabrus massa and melops work together in building their
nests with sea-weed, shells, &c.31 But the males of certain fishes do all the
work, and afterwards take exclusive charge of the young. This is the case
with the dull-coloured gobies,32 in which the sexes are not known to differ
in colour, and likewise with the sticklebacks (Gasterosteus), in which the
males become brilliantly coloured during the spawning-season. The male of
the smooth-tailed stickleback (G. leiurus) performs during a long time the
duties of a nurse with exemplary care and vigilance, and is continually
employed in gently leading back the young to the nest when they stray too
far. He courageously drives away all enemies, including the females of his
own species. It would indeed be no small relief to the male if the female,
after depositing her eggs, were immediately devoured by some enemy, for
he is forced incessantly to drive her from the nest.33
The males of certain other fishes inhabiting South America and Ceylon, and
belonging to two distinct orders, have the extraordinary habit of hatching

Page 23

the eggs laid by the females within their mouths or branchial cavities.34
With the Amazonian species which follow this habit, the males, as I am
informed by the kindness of Professor Agassiz, “not only are generally
brighter than the females, but the difference is greater at the spawning-
season than at any other time.” The species of Geophagus act in the same
manner; and in this genus, a conspicuous protuberance becomes developed
on the forehead of the males during the breeding-season. With the various
species of Chromids, as Professor Agassiz likewise informs me, sexual
differences in colour may be observed, “whether they lay their eggs in the
water among aquatic plants, or deposit them in holes, leaving them to come
out without further care, or build shallow nests in the river-mud, over which
they sit, as our Promotis does. It ought also to be observed that these sitters
are among the brightest species in their respective families; for instance,
Hygrogonus is bright green, with large black ocelli, encircled with the most
brilliant red.” Whether with all the species of Chromids it is the male alone
which sits on the eggs is not known. It is, however, manifest that the fact of
the eggs being protected or unprotected, has had little or no influence on the
differences in colour between the sexes. It is further manifest, in all the
cases in which the males take exclusive charge of the nests and young, that
the destruction of the brighter-coloured males would be far more influential
on the character of the race, than the destruction of the brighter-coloured
females; for the death of the male during the period of incubation or nursing
would entail the death of the young, so that these could not inherit his
peculiarities; yet, in many of these very cases the males are more
conspicuously coloured than the females.
In most of the Lophobranchii (Pipe-fish, Hippocampi, &c.) the males have
either marsupial sacks or hemispherical depressions on the abdomen, in
which the ova laid by the female are hatched. The males also shew great
attachment to their young.35 The sexes do not commonly differ much in
colour; but Dr. Günther believes that the male Hippocampi are rather
brighter than the females. The genus Solenostoma, however, offers a very
curious exceptional case,36 for the female is much more vividly coloured
and spotted than the male, and she alone has a marsupial sack and hatches
the eggs; so that the female of Solenostoma differs from all the other
Lophobranchii in this latter respect, and from almost all other fishes, in
being more brightly coloured than the male. It is improbable that this

Page 24

remarkable double inversion of character in the female should be an
accidental coincidence. As the males of several fishes which take exclusive
charge of the eggs and young are more brightly coloured than the females,
and as here the female Solenostoma takes the same charge and is brighter
than the male, it might be argued that the conspicuous colours of the sex
which is the most important of the two for the welfare of the offspring must
serve, in some manner, as a protection. But from the multitude of fishes, the
males of which are either permanently or periodically brighter than the
females, but whose life is not at all more important than that of the female
for the welfare of the species, this view can hardly be maintained. When we
treat of birds we shall meet with analogous cases in which there has been a
complete inversion of the usual attributes of the two sexes, and we shall
then give what appears to be the probable explanation, namely, that the
males have selected the more attractive females, instead of the latter having
selected, in accordance with the usual rule throughout the animal kingdom,
the more attractive males.
On the whole we may conclude, that with most fishes, in which the sexes
differ in colour or in other ornamental characters, the males originally
varied, with their variations transmitted to the same sex, and accumulated
through sexual selection by attracting or exciting the females. In many
cases, however, such characters have been transferred, either partially or
completely, to the females. In other cases, again, both sexes have been
coloured alike for the sake of protection; but in no instance does it appear
that the female alone has had her colours or other characters specially
modified for this purpose.
The last point which need be noticed is that in many parts of the world
fishes are known to make peculiar noises, which are described in some
cases as being musical. Very little has been ascertained with respect to the
means by which such sounds are produced, and even less about their
purpose. The drumming of the Umbrinas in the European seas is said to be
audible from a depth of twenty fathoms. The fishermen of Rochelle assert
“that the males alone make the noise during the spawning-time; and that it
is possible by imitating it, to take them without bait.”37 If this statement is
trustworthy, we have an instance in this, the lowest class of the Vertebrata,
of what we shall find prevailing throughout the other vertebrate classes, and
which prevails, as we have already seen, with insects and spiders; namely,

Page 25

that vocal and instrumental sounds so commonly serve as a love-call or as a
love-charm, that the power of producing them was probably first developed
in connection with the propagation of the species.

Fig. 31. Triton cristatus (half natural size, from Bell’s ‘British Reptiles’). Upper figure,
male during the breeding-season; lower figure, female.

Amphibians.

Urodela.—First for the tailed amphibians. The sexes of salamanders or
newts often differ much both in colour and structure. In some species
prehensile claws are developed on the forelegs of the males during the
breeding-season; and at this season in the male Triton palmipes the hind-
feet are provided with a swimming web, which is almost completely
absorbed during the winter; so that their feet then resemble those of the
female.38 This structure no doubt aids the male in his eager search and
pursuit of the female. With our common newts (Triton punctatus and
cristatus) a deep, much-indented crest is developed along the back and tail
of the male during the breeding-season, being absorbed during the winter. It
is not furnished, as Mr. St. George Mivart informs me, with muscles, and
therefore cannot be used for locomotion. As during the season of courtship
it becomes edged with bright colours, it serves, there can hardly be a doubt,
as a masculine ornament. In many species the body presents strongly

Page 26

contrasted, though lurid tints; and these become more vivid during the
breeding-season. The male, for instance, of our common little newt (Triton
punctatus) is “brownish-grey above, passing into yellow beneath, which in
the spring becomes a rich bright orange, marked everywhere with round
dark spots.” The edge of the crest also is then tipped with bright red or
violet. The female is usually of a yellowish-brown colour with scattered
brown dots; and the lower surface is often quite plain.39 The young are
obscurely tinted. The ova are fertilised during the act of deposition and are
not subsequently tended by either parent. We may therefore conclude that
the males acquired their strongly-marked colours and ornamental
appendages through sexual selection; these being transmitted either to the
male offspring alone or to both sexes.

Anura or Batrachia.—With many frogs and toads the colours evidently
serve as a protection, such as the bright green tints of tree-frogs and the
obscure mottled shades of many terrestrial species. The most conspicuously
coloured toad which I ever saw, namely the Phryniscus nigricans40 had the
whole upper surface of the body as black as ink, with the soles of the feet
and parts of the abdomen spotted with the brightest vermilion. It crawled
about the bare sandy or open grassy plains of La Plata under a scorching
sun, and could not fail to catch the eye of every passing creature. These
colours may be beneficial by making this toad known to all birds of prey as
a nauseous mouthful; for it is familiar to every one that these animals emit a
poisonous secretion, which causes the mouth of a dog to froth, as if attacked
by hydrophobia. I was the more struck with the conspicuous colours of this
toad, as close by I found a lizard (Proctotretus multimaculatus) which,
when frightened, flattened its body, closed its eyes, and then from its
mottled tints could hardly be distinguishable from the surrounding sand.

With respect to sexual differences of colour, Dr. Günther knows of no
striking instance with frogs or toads; yet he can often distinguish the male
from the female, by the tints of the former being a little more intense. Nor
does Dr. Günther know of any striking difference in external structure
between the sexes, excepting the prominences which become developed
during the breeding-season on the front-legs of the male, by which he is
enabled to hold the female. The Megalophrys montana41 (fig. 32) offers the

Page 27

best case of a certain amount of structural difference between the sexes; for
in the male the tip of the nose and the eyelids are produced into triangular
flaps of skin, and there is a little black tubercle on the back—characters
which are absent, or only feebly developed, in the females. It is surprising
that frogs and toads should not have acquired more strongly-marked sexual
differences; for though cold-blooded, their passions are strong. Dr. Günther
informs me that he has several times found an unfortunate female toad dead
and smothered from having been so closely embraced by three or four
males.

Fig. 32. Megalophrys montana. The two left-hand figures, the
male; the two right-hand figures, the female.

These animals, however, offer one interesting sexual difference, namely in
the musical powers possessed by the males; but to speak of music, when
applied to the discordant and overwhelming sounds emitted by male bull-
frogs and some other species, seems, according to our taste, a singularly
inappropriate expression. Nevertheless certain frogs sing in a decidedly
pleasing manner. Near Rio de Janeiro I used often to sit in the evening to
listen to a number of little Hylæ, which, perched on blades of grass close to
the water, sent forth sweet chirping notes in harmony. The various sounds
are emitted chiefly by the males during the breeding-season, as in the case
of the croaking of our common frog.42 In accordance with this fact the vocal
organs of the males are more highly developed than those of the females. In

Page 28

some genera the males alone are provided with sacs which open into the
larynx.43 For instance, in the edible frog (Rana esculenta) “the sacs are
peculiar to the males, and become, when filled with air in the act of
croaking, large globular bladders, standing out one on each side of the head,
near the corners of the mouth.” The croak of the male is thus rendered
exceedingly powerful; whilst that of the female is only a slight groaning
noise.44 The vocal organs differ considerably in structure in the several
genera of the family; and their development in all cases may be attributed to
sexual selection.

Reptiles.

Chelonia.—Tortoises and turtles do not offer well-marked sexual
differences. In some species, the tail of the male is longer than that of the
female. In some, the plastron or lower surface of the shell of the male is
slightly concave in relation to the back of the female. The male of the mud-
turtle of the United States (Chrysemys picta) has claws on its front-feet
twice as long as those of the female; and these are used when the sexes
unite.45 With the huge tortoise of the Galapagos Islands (Testudo nigra) the
males are said to grow to a larger size than the females: during the pairing-
season, and at no other time, the male utters a hoarse, bellowing noise,
which can be heard at the distance of more than a hundred yards; the
female, on the other hand, never uses her voice.46

Crocodilia.—The sexes apparently do not differ in colour; nor do I know
that the males fight together, though this is probable, for some kinds make a
prodigious display before the females. Bartram47 describes the male
alligator as striving to win the female by splashing and roaring in the midst
of a lagoon, “swollen to an extent ready to burst, with his head and tail
lifted up, he spins or twirls round on the surface of the water, like an Indian
chief rehearsing his feats of war.” During the season of love, a musky odour
is emitted by the submaxillary glands of the crocodile, and pervades their
haunts.48

Ophidia.—I have little to say about Snakes. Dr. Günther informs me that
the males are always smaller than the females, and generally have longer

Page 29

and slenderer tails; but he knows of no other difference in external
structure. In regard to colour, Dr. Günther can almost always distinguish the
male from the female by his more strongly-pronounced tints; thus the black
zigzag band on the back of the male English viper is more distinctly defined
than in the female. The difference is much plainer in the Rattle-snakes of N.
America, the male of which, as the keeper in the Zoological Gardens
shewed me, can instantly be distinguished from the female by having more
lurid yellow about its whole body. In S. Africa the Bucephalus capensis
presents an analogous difference, for the female “is never so fully
variegated with yellow on the sides, as the male.”49 The male of the Indian
Dipsas cynodon, on the other hand, is blackish-brown, with the belly partly
black, whilst the female is reddish or yellowish-olive with the belly either
uniform yellowish or marbled with black.

In the Tragops dispar of the same country, the male is bright green, and the
female bronze-coloured.50 No doubt the colours of some snakes serve as a
protection, as the green tints of tree-snakes and the various mottled shades
of the species which live in sandy places; but it is doubtful whether the
colours of many kinds, for instance of the common English snake or viper,
serve to conceal them; and this is still more doubtful with the many foreign
species which are coloured with extreme elegance.
During the breeding-season their anal scent-glands are in active function;51
and so it is with the same glands in lizards, and as we have seen with the
submaxillary glands of crocodiles. As the males of most animals search for
the females, these odoriferous glands probably serve to excite or charm the
female, rather than to guide her to the spot where the male may be found.52
Male snakes, though appearing so sluggish, are amorous; for many have
been observed crowding round the same female, and even round the dead
body of a female. They are not known to fight together from rivalry. Their
intellectual powers are higher than might have been anticipated. An
excellent observer in Ceylon, Mr. E. Layard,53 saw a Cobra thrust its head
through a narrow hole and swallow a toad. “With this incumbrance he could
not withdraw himself; finding this, he reluctantly disgorged the precious
morsel, which began to move off; this was too much for snake philosophy
to bear, and the toad was again seized, and again was the snake, after
violent efforts to escape, compelled to part with its prey. This time,

Page 30

however, a lesson had been learnt, and the toad was seized by one leg,
withdrawn, and then swallowed in triumph.”
It does not, however, follow because snakes have some reasoning power
and strong passions, that they should likewise be endowed with sufficient
taste to admire brilliant colours in their partners, so as to lead to the
adornment of the species through sexual selection. Nevertheless it is
difficult to account in any other manner for the extreme beauty of certain
species; for instance, of the coral-snakes of S. America, which are of a rich
red with black and yellow transverse bands. I well remember how much
surprise I felt at the beauty of the first coral-snake which I saw gliding
across a path in Brazil. Snakes coloured in this peculiar manner, as Mr.
Wallace states on the authority of Dr. Günther,54 are found nowhere else in
the world except in S. America, and here no less than four genera occur.
One of these, Elaps, is venomous; a second and widely-distinct genus is
doubtfully venomous, and the two others are quite harmless. The species
belonging to these distinct genera inhabit the same districts, and are so like
each other, that no one “but a naturalist would distinguish the harmless
from the poisonous kinds.” Hence, as Mr. Wallace believes, the innocuous
kinds have probably acquired their colours as a protection, on the principle
of imitation; for they would naturally be thought dangerous by their
enemies. The cause, however, of the bright colours of the venomous Elaps
remains to be explained, and this may perhaps be sexual selection.

Lacertilia.—The males of some, probably of many kinds of lizards fight
together from rivalry. Thus the arboreal Anolis cristatellus of S. America is
extremely pugnacious: “During the spring and early part of the summer,
two adult males rarely meet without a contest. On first seeing one another,
they nod their heads up and down three or four times, at the same time
expanding the frill or pouch beneath the throat; their eyes glisten with rage,
and after waving their tails from side to side for a few seconds, as if to
gather energy, they dart at each other furiously, rolling over and over, and
holding firmly with their teeth. The conflict generally ends in one of the
combatants losing his tail, which is often devoured by the victor.” The male
of this species is considerably larger than the female;55 and this, as far as Dr.
Günther has been able to ascertain, is the general rule with lizards of all
kinds.

Page 31

The sexes often differ greatly in various external characters. The male of the
above-mentioned Anolis is furnished with a crest, which runs along the
back and tail, and can be erected at pleasure; but of this crest the female
does not exhibit a trace. In the Indian Cophotis ceylanica, the female
possesses a dorsal crest, though much less developed than in the male; and
so it is, as Dr. Günther informs me, with the females of many Iguanas,
Chameleons and other lizards. In some species, however, the crest is
equally developed in both sexes, as in the Iguana tuberculata. In the genus
Sitana, the males alone are furnished with a large throat-pouch (fig. 33),
which can be folded up like a fan, and is coloured blue, black, and red; but
these splendid colours are exhibited only during the pairing-season. The
female does not possess even a rudiment of this appendage. In the Anolis
cristatellus, according to Mr. Austen, the throat-pouch, which is bright red
marbled with yellow, is present, though in a rudimental condition, in the
female. Again, in certain other lizards, both sexes are equally well provided
with throat-pouches. Here, as in
so many previous cases, we see
with species belonging to the
same group, the same character
confined to the males, or more
largely developed in the males
than in the females, or equally
developed in both sexes. The
little lizards of the genus Draco,
which glide through the air on
their rib-supported parachutes,
and which in the beauty of their Fig. 33. Sitana minor. Male, with the gular pouch
expanded (from Günther’s ‘Reptiles of India’).
colours baffle description, are
furnished with skinny appendages to the throat, “like the wattles of
gallinaceous birds.” These become erected when the animal is excited. They
occur in both sexes, but are best developed in the male when arrived at
maturity, at which age the middle appendage is sometimes twice as long as
the head. Most of the species likewise have a low crest running along the
neck; and this is much more developed in the full-grown males, than in the
females or young males.56

Page 32

There are other and much more remarkable differences between the sexes
of certain lizards. The male of Ceratophora aspera bears on the extremity
of his snout an appendage half as long as
the head. It is cylindrical, covered with
scales, flexible, and apparently capable of
erection: in the female it is quite
rudimental. In a second species of the same
genus a terminal scale forms a minute horn
on the summit of the flexible appendage;
and in a third species (C. Stoddartii, fig.
34) the whole appendage is converted into
a horn, which is usually of a white colour,
but assumes a purplish tint when the animal
Fig. 34. Ceratophora Stoddartii. is excited. In the adult male of this latter
Upper figure, male; lower figure, species the horn is half an inch in length,
female. but is of quite minute size in the female and
in the young. These appendages, as Dr.
Günther has remarked to me, may be compared with the combs of
gallinaceous birds, and apparently serve as ornaments.

Page 33

Fig. 35. Chamæleon bifurcus. Upper figure, male; lower
figure, female.

In the genus Chamæleon we come to the climax of difference between the
sexes. The upper part of the skull of the male C. bifurcus (fig. 35), an
inhabitant of Madagascar, is produced into two great, solid, bony
projections, covered with scales like the rest of the head; and of this
wonderful modification of structure the female exhibits only a rudiment.
Again, in Chamæleon Owenii (fig. 36), from the West Coast of Africa, the
male bears on his snout and forehead three curious horns, of which the
female has not a trace. These horns consist of an excrescence of bone
covered with a smooth sheath, forming part of the general integuments of
the body, so that they are identical in structure with those of a bull, goat, or
other sheath-horned ruminant. Although the three horns differ so much in
appearance from the two great prolongations of the skull in C. bifurcus, we
can hardly doubt that they serve the same general purpose in the economy
of these two animals. The first conjecture which will occur to every one is

Page 34

that they are used by the males for fighting together; but Dr. Günther, to
whom I am indebted for the foregoing details, does not believe that such
peaceable creatures would ever become pugnacious. Hence we are driven to
infer that these almost monstrous
deviations of structure serve as
masculine ornaments.
With many kinds of lizards, the
sexes differ slightly in colour, the
tints and stripes of the males being
brighter and more distinctly defined
than in the females. This, for
instance, is the case with the
previously-mentioned Cophotis and
with the Acanthodactylus capensis
of S. Africa. In a Cordylus of the
latter country, the male is either
much redder or greener than the
Fig. 36. Chamæleon Owenii. Upper figure,
female. In the Indian Calotes
male; lower figure, female.
nigrilabris there is a greater
difference in colour between the
sexes; the lips also of the male are black, whilst those of the female are
green. In our common little viviparous lizard (Zootoca vivipara) “the under
side of the body and base of the tail in the male are bright orange, spotted
with black; in the female these parts are pale greyish-green without spots.”57
We have seen that the males alone of Sitana possess a throat-pouch; and
this is splendidly tinted with blue, black, and red. In the Proctotretus tenuis
of Chile the male alone is marked with spots of blue, green, and coppery-
red.58 I collected in S. America fourteen species of this genus, and though I
neglected to record the sexes, I observed that certain individuals alone were
marked with emerald-like green spots, whilst others had orange-coloured
gorges; and these in both cases no doubt were the males.
In the foregoing species, the males are more brightly coloured than the
females, but with many lizards both sexes are coloured in the same elegant
or even magnificent manner; and there is no reason to suppose that such
conspicuous colours are protective. With some lizards, however, the green
tints no doubt serve for concealment; and an instance has already been

Page 35

incidently given of one species of Proctotretus which closely resembles the
sand on which it lives. On the whole we may conclude with tolerable safety
that the beautiful colours of many lizards, as well as various appendages
and other strange modifications of structure, have been gained by the males
through sexual selection for the sake of ornament, and have been
transmitted either to their male offspring alone or to both sexes. Sexual
selection, indeed, seems to have played almost as important a part with
reptiles as with birds. But the less conspicuous colours of the females in
comparison with those of the males cannot be accounted for, as Mr. Wallace
believes to be the case with birds, by the exposure of the females to danger
during incubation.

CHAPTER XIII.

Secondary Sexual Characters of Birds.
Sexual differences—Law of battle—Special weapons—Vocal organs—Instrumental music—Love-
antics and dances—Decorations, permanent and seasonal—Double and single annual moults—
Display of ornaments by the males.

Secondary sexual characters are more diversified and conspicuous in birds,
though not perhaps entailing more important changes of structure, than in
any other class of animals. I shall, therefore, treat the subject at
considerable length. Male birds sometimes, though rarely, possess special
weapons for fighting with each other. They charm the females by vocal or
instrumental music of the most varied kinds. They are ornamented by all
sorts of combs, wattles, protuberances, horns, air-distended sacs, top-knots,
naked shafts, plumes and lengthened feathers gracefully springing from all
parts of the body. The beak and naked skin about the head, and the feathers
are often gorgeously coloured. The males sometimes pay their court by
dancing, or by fantastic antics performed either on the ground or in the air.
In one instance, at least, the male emits a musky odour which we may
suppose serves to charm or excite the female; for that excellent observer,
Mr. Ramsay,59 says of the Australian musk-duck (Biziura lobata) that “the
smell which the male emits during the summer months is confined to that
sex, and in some individuals is retained throughout the year; I have never

Page 36

even in the breeding-season, shot a female which had any smell of musk.”
So powerful is this odour during the pairing-season, that it can be detected
long before the bird can be seen.60 On the whole, birds appear to be the
most æsthetic of all animals, excepting of course man, and they have nearly
the same taste for the beautiful as we have. This is shewn by our enjoyment
of the singing of birds, and by our women, both civilised and savage,
decking their heads with borrowed plumes, and using gems which are
hardly more brilliantly coloured than the naked skin and wattles of certain
birds.
Before treating of the characters with which we are here more particularly
concerned, I may just allude to certain differences between the sexes which
apparently depend on differences in their habits of life; for such cases,
though common in the lower, are rare in the higher classes. Two humming-
birds belonging to the genus Eustephanus, which inhabit the island of Juan
Fernandez, were long thought to be specifically distinct, but are now
known, as Mr. Gould informs me, to be the sexes of the same species, and
they differ slightly in the form of the beak. In another genus of humming-
birds (Grypus), the beak of the male is serrated along the margin and
hooked at the extremity, thus differing much from that of the female. In the
curious Neomorpha of New Zealand, there is a still wider difference in the
form of the beak; and Mr. Gould has been informed that the male with his
“straight and stout beak” tears off the bark of trees, in order that the female
may feed on the uncovered larvæ with her weaker and more curved beak.
Something of the same kind may be observed with our goldfinch (Carduelis
elegans), for I am assured by Mr. J. Jenner Weir that the bird-catchers can
distinguish the males by their slightly longer beaks. The flocks of males, as
an old and trustworthy bird-catcher asserted, are commonly found feeding
on the seeds of the teazle (Dipsacus) which they can reach with their
elongated beaks, whilst the females more commonly feed on the seeds of
the betony or Scrophularia. With a slight difference of this nature as a
foundation, we can see how the beaks of the two sexes might be made to
differ greatly through natural selection. In all these cases, however,
especially in that of the quarrelsome humming-birds, it is possible that the
differences in the beaks may have been first acquired by the males in
relation to their battles, and afterwards led to slightly changed habits of life.

Page 37

Law of Battle.—Almost all male birds are extremely pugnacious, using
their beaks, wings, and legs for fighting together. We see this every spring
with our robins and sparrows. The smallest of all birds, namely the
humming-bird, is one of the most quarrelsome. Mr. Gosse61 describes a
battle, in which a pair of humming-birds seized hold of each other’s beaks,
and whirled round and round, till they almost fell to the ground; and M.
Montes de Oca, in speaking of another genus, says that two males rarely
meet without a fierce aerial encounter: when kept in cages “their fighting
has mostly ended in the splitting of the tongue of one of the two, which then
surely dies from being unable to feed.”62 With Waders, the males of the
common water-hen (Gallinula chloropus) “when pairing, fight violently for
the females: they stand nearly upright in the water and strike with their
feet.” Two were seen to be thus engaged for half an hour, until one got hold
of the head of the other which would have been killed, had not the observer
interfered; the female all the time looking on as a quiet spectator.63 The
males of an allied bird (Gallicrex cristatus), as Mr. Blyth informs me, are
one third larger than the females, and are so pugnacious during the
breeding-season, that they are kept by the natives of Eastern Bengal for the
sake of fighting. Various other birds are kept in India for the same purpose,
for instance the Bulbuls (Pycnonotus hæmorrhous) which “fight with great
spirit.”64
The polygamous Ruff (Machetes pugnax, fig. 37) is notorious for his
extreme pugnacity; and in the spring, the males, which are considerably
larger than the females, congregate day after day at a particular spot, where
the females propose to lay their eggs. The fowlers discover these spots by
the turf being trampled somewhat bare. Here they fight very much like
game-cocks, seizing each other with their beaks and striking with their
wings. The great ruff of feathers round the neck is then erected, and
according to Col. Montagu “sweeps the ground as a shield to defend the
more tender parts;” and this is the only instance known to me in the case of
birds, of any structure serving as a shield. The ruff of feathers, however,
from its varied and rich colours probably serves in chief part as an
ornament. Like most pugnacious birds, they seem always ready to fight, and
when closely confined often kill each other; but Montagu observed that
their pugnacity becomes greater during the spring, when the long feathers
on their necks are fully developed; and at this period the least movement by

Page 38

any one bird provokes a general battle.65 Of the pugnacity of web-footed
birds, two instances will suffice: in Guiana “bloody fights occur during the
breeding-season between the males of the wild musk-duck (Cairina
moschata); and where these fights have occurred the river is covered for
some distance with feathers.”66 Birds which seem ill-adapted for fighting
engage in fierce conflicts; thus with the pelican the stronger males drive
away the weaker ones, snapping with their huge beaks and giving heavy
blows with their wings. Male snipes fight together, “tugging and pushing
each other with their bills in the most curious manner imaginable.” Some
few species are believed never to fight; this is the case, according to
Audubon, with one of the woodpeckers of the United States (Picus
auratus), although “the hens are followed by even half a dozen of their gay
suitors.”67

Fig. 37. The Ruff or Machetes pugnax (from Brehm’s ‘Thierteben’).

The males of many birds are larger than the females, and this no doubt is an
advantage to them in their battles with their rivals, and has been gained
through sexual selection. The difference in size between the two sexes is
carried to an extreme point in several Australian species; thus the male
musk-duck (Biziura) and the male Cincloramphus cruralis (allied to our
pipits) are by measurement actually twice as large as their respective
females.68 With many other birds the females are larger than the males; and
as formerly remarked, the explanation often given, namely that the females
have most of the work in feeding their young, will not suffice. In some few
cases, as we shall hereafter see, the females apparently have acquired their

Page 39

greater size and strength for the sake of conquering other females and
obtaining possession of the males.
The males of many gallinaceous birds, especially of the polygamous kinds,
are furnished with special weapons for fighting with their rivals, namely
spurs, which can be used with fearful effect. It has been recorded by a
trustworthy writer69 that in Derbyshire a kite struck at a game-hen
accompanied by her chickens, when the cock rushed to the rescue and drove
his spur right through the eye and skull of the aggressor. The spur was with
difficulty drawn from the skull, and as the kite though dead retained his
grasp, the two birds were firmly locked together; but the cock when
disentangled was very little injured. The invincible courage of the game-
cock is notorious: a gentleman who long ago witnessed the following brutal
scene, told me that a bird had both its legs broken by some accident in the
cock-pit, and the owner laid a wager that if the legs could be spliced so that
the bird could stand upright, he would continue fighting. This was effected
on the spot, and the bird fought with undaunted courage until he received
his death-stroke. In Ceylon a closely-allied and wild species, the Gallus
Stanleyi, is known to fight desperately “in defence of his seraglio,” so that
one of the combatants is frequently found dead.70 An Indian partridge
(Ortygornis gularis), the male of which is furnished with strong and sharp
spurs, is so quarrelsome, “that the scars of former fights disfigure the breast
of almost every bird you kill.”71
The males of almost all gallinaceous birds, even those which are not
furnished with spurs, engage during the breeding-season in fierce conflicts.
The Capercailzie and Black-cock (Tetrao urogallus and T. tetrix), which are
both polygamists, have regular appointed places, where during many weeks
they congregate in numbers to fight together and to display their charms
before the females. M. W. Kowalevsky informs me that in Russia he has
seen the snow all bloody on the arenas where the Capercailzie have fought;
and the Black-cocks “make the feathers fly in every direction,” when
several “engage in a battle royal.” The elder Brehm gives a curious account
of the Balz, as the love-dance and love-song of the Black-cock is called in
Germany. The bird utters almost continuously the most strange noises: “he
holds his tail up and spreads it out like a fan, he lifts up his head and neck
with all the feathers erect, and stretches his wings from the body. Then he
takes a few jumps in different directions, sometimes in a circle, and presses

Page 40

the under part of his beak so hard against the ground that the chin-feathers
are rubbed off. During these movements he beats his wings and turns round
and round. The more ardent he grows the more lively he becomes, until at
last the bird appears like a frantic creature.” At such times the black-cocks
are so absorbed that they become almost blind and deaf, but less so than the
capercailzie: hence bird after bird may be shot on the same spot, or even
caught by the hand. After performing these antics the males begin to fight:
and the same black-cock, in order to prove his strength over several
antagonists, will visit in the course of one morning several Balz-places,
which remain the same during successive years.72
The peacock with his long train appears more like a dandy than a warrior,
but he sometimes engages in fierce contests: the Rev. W. Darwin Fox
informs me that two peacocks became so excited whilst fighting at some
little distance from Chester that they flew over the whole city, still fighting,
until they alighted on the top of St. John’s tower.
The spur, in those gallinaceous birds which are thus provided, is generally
single; but Polyplectron (see fig. 51, p. 90) has two or more on each leg;
and one of the Blood-pheasants (Ithaginis cruentus) has been seen with five
spurs. The spurs are generally confined to the male, being represented by
mere knobs or rudiments in the female; but the females of the Java peacock
(Pavo muticus) and, as I am informed by Mr. Blyth, of the small fire-backed
pheasant (Euplocamus erythropthalmus) possess spurs. In Galloperdix it is
usual for the males to have two spurs, and for the females to have only one
on each leg.73 Hence spurs may safely be considered as a masculine
character, though occasionally transferred in a greater or less degree to the
females. Like most other secondary sexual characters, the spurs are highly
variable both in number and development in the same species.

Page 41

Fig. 38. Palamedea cornuta (from Brehm), shewing the double-wing-
spurs, and the filament on the head.

Various birds have spurs on their wings. But the Egyptian goose
(Chenalopex ægyptiacus) has only “bare obtuse knobs,” and these probably
shew us the first steps by which true spurs have been developed in other
allied birds. In the spur-winged goose, Plectropterus gambensis, the males
have much larger spurs than the females; and they use them, as I am
informed by Mr. Bartlett, in fighting together, so that, in this case, the wing-
spurs serve as sexual weapons; but according to Livingstone, they are
chiefly used in the defence of the young. The Palamedea (fig. 38) is armed
with a pair of spurs on each wing; and these are such formidable weapons
that a single blow has driven a dog howling away. But it does not appear
that the spurs in this case, or in that of some of the spur-winged rails, are
larger in the male than in the female.74 In certain plovers, however, the
wing-spurs must be considered as a sexual character. Thus in the male of
our common peewit (Vanellus cristatus) the tubercle on the shoulder of the
wing becomes more prominent during the breeding-season, and the males
are known to fight together. In some species of Lobivanellus a similar
tubercle becomes developed during the breeding-season “into a short horny
spur.” In the Australian L. lobatus both sexes have spurs, but these are much
larger in the males than in the females. In an allied bird, the Hoplopterus
armatus, the spurs do not increase in size during the breeding-season; but

Page 42

these birds have been seen in Egypt to fight together, in the same manner as
our peewits, by turning suddenly in the air and striking sideways at each
other, sometimes with a fatal result. Thus also they drive away other
enemies.75
The season of love is that of battle; but the males of some birds, as of the
game-fowl and ruff, and even the young males of the wild turkey and
grouse,76 are ready to fight whenever they meet. The presence of the female
is the teterrima belli causa. The Bengali baboos make the pretty little males
of the amadavat (Estrelda amandava) fight together by placing three small
cages in a row, with a female in the middle; after a little time the two males
are turned loose, and immediately a desperate battle ensues.77 When many
males congregate at the same appointed spot and fight together, as in the
case of grouse and various other birds, they are generally attended by the
females,78 which afterwards pair with the victorious combatants. But in
some cases the pairing precedes instead of succeeding the combat: thus,
according to Audubon,79 several males of the Virginian goat-sucker
(Caprimulgus Virginianus) “court, in a highly entertaining manner, the
female, and no sooner has she made her choice, than her approved gives
chase to all intruders, and drives them beyond his dominions.” Generally
the males try with all their power to drive away or kill their rivals before
they pair. It does not, however, appear that the females invariably prefer the
victorious males. I have indeed been assured by M. W. Kowalevsky that the
female capercailzie sometimes steals away with a young male who has not
dared to enter the arena with the older cocks; in the same manner as
occasionally happens with the does of the red-deer in Scotland. When two
males contend in presence of a single female, the victor, no doubt,
commonly gains his desire; but some of these battles are caused by
wandering males trying to distract the peace of an already mated pair.80
Even with the most pugnacious species it is probable that the pairing does
not depend exclusively on the mere strength and courage of the male: for
such males are generally decorated with various ornaments, which often
become more brilliant during the breeding-season, and which are
sedulously displayed before the females. The males also endeavour to
charm or excite their mates by love-notes, songs, and antics; and the
courtship is, in many instances, a prolonged affair. Hence it is not probable
that the females are indifferent to the charms of the opposite sex, or that

Page 43

they are invariably compelled to yield to the victorious males. It is more
probable that the females are excited, either before or after the conflict, by
certain males, and thus unconsciously prefer them. In the case of Tetrao
umbellus, a good observer81 goes so far as to believe that the battles of the
males “are all a sham, performed to show themselves to the greatest
advantage before the admiring females who assemble around; for I have
never been able to find a maimed hero, and seldom more than a broken
feather.” I shall have to recur to this subject, but I may here add that with
the Tetrao cupido of the United States, about a score of males assemble at a
particular spot, and strutting about make the whole air resound with their
extraordinary noises. At the first answer from a female the males begin to
fight furiously, and the weaker give way; but then, according to Audubon,
both the victors and vanquished search for the female, so that the females
must either then exert a choice, or the battle must be renewed. So, again,
with one of the Field-starlings of the United States (Sturnella ludoviciana)
the males engage in fierce conflicts, “but at the sight of a female they all fly
after her, as if mad.”82

Vocal and instrumental Music.—With birds the voice serves to express
various emotions, such as distress, fear, anger, triumph, or mere happiness.
It is apparently sometimes used to excite terror, as with the hissing noise
made by some nestling-birds. Audubon83 relates that a night-heron (Ardea
nycticorax, Linn.) which he kept tame, used to hide itself when a cat
approached, and then “suddenly start up uttering one of the most frightful
cries, apparently enjoying the cat’s alarm and flight.” The common
domestic cock clucks to the hen, and the hen to her chickens, when a dainty
morsel is found. The hen, when she has laid an egg, “repeats the same note
very often, and concludes with the sixth above, which she holds for a longer
time;”84 and thus she expresses her joy. Some social birds apparently call to
each other for aid; and as they flit from tree to tree, the flock is kept
together by chirp answering chirp. During the nocturnal migrations of geese
and other water-fowl, sonorous clangs from the van may be heard in the
darkness overhead, answered by clangs in the rear. Certain cries serve as
danger-signals, which, as the sportsman knows to his cost, are well
understood by the same species and by others. The domestic cock crows,
and the humming-bird chirps, in triumph over a defeated rival. The true
song, however, of most birds and various strange cries are chiefly uttered

Page 44

during the breeding-season, and serve as a charm, or merely as a call-note,
to the other sex.
Naturalists are much divided with respect to the object of the singing of
birds. Few more careful observers ever lived than Montagu, and he
maintained that the “males of song-birds and of many others do not in
general search for the female, but, on the contrary, their business in the
spring is to perch on some conspicuous spot breathing out their full and
amorous notes, which, by instinct, the female knows, and repairs to the spot
to choose her mate.”85 Mr. Jenner Weir informs me that this is certainly the
case with the nightingale. Bechstein, who kept birds during his whole life,
asserts, “that the female canary always chooses the best singer, and that in a
state of nature the female finch selects that male out of a hundred whose
notes please her most.”86 There can be no doubt that birds closely attend to
each other’s song. Mr. Weir has told me of the case of a bullfinch which had
been taught to pipe a German waltz, and who was so good a performer that
he cost ten guineas; when this bird was first introduced into a room where
other birds were kept and he began to sing, all the others, consisting of
about twenty linnets and canaries, ranged themselves on the nearest side of
their cages, and listened with the greatest interest to the new performer.
Many naturalists believe that the singing of birds is almost exclusively “the
effect of rivalry and emulation,” and not for the sake of charming their
mates. This was the opinion of Daines Barrington and White of Selborne,
who both especially attended to this subject.87 Barrington, however, admits
that “superiority in song gives to birds an amazing ascendancy over others,
as is well known to bird-catchers.”
It is certain that there is an intense degree of rivalry between the males in
their singing. Bird-fanciers match their birds to see which will sing longest;
and I was told by Mr. Yarrell that a first-rate bird will sometimes sing till he
drops down almost dead, or, according to Bechstein,88 quite dead from
rupturing a vessel in the lungs. Whatever the cause may be, male birds, as I
hear from Mr. Weir, often die suddenly during the season of song. That the
habit of singing is sometimes quite independent of love is clear, for a sterile
hybrid canary-bird has been described89 as singing whilst viewing itself in a
mirror, and then dashing at its own image; it likewise attacked with fury a
female canary when put into the same cage. The jealousy excited by the act
of singing is constantly taken advantage of by bird-catchers; a male, in good

Page 45

song, is hidden and protected, whilst a stuffed bird, surrounded by limed
twigs, is exposed to view. In this manner a man, as Mr. Weir informs me,
has caught, in the course of a single day, fifty, and in one instance seventy,
male chaffinches. The power and inclination to sing differ so greatly with
birds that although the price of an ordinary male chaffinch is only sixpence,
Mr. Weir saw one bird for which the bird-catcher asked three pounds; the
test of a really good singer being that it will continue to sing whilst the cage
is swung round the owner’s head.
That birds should sing from emulation as well as for the sake of charming
the female, is not at all incompatible; and, indeed, might have been
expected to go together, like decoration and pugnacity. Some authors,
however, argue that the song of the male cannot serve to charm the female,
because the females of some few species, such as the canary, robin, lark,
and bullfinch, especially, as Bechstein remarks, when in a state of
widowhood, pour forth fairly melodious strains. In some of these cases the
habit of singing may be in part attributed to the females having been highly
fed and confined,90 for this disturbs all the usual functions connected with
the reproduction of the species. Many instances have already been given of
the partial transference of secondary masculine characters to the female, so
that it is not at all surprising that the females of some species should
possess the power of song. It has also been argued, that the song of the male
cannot serve as a charm, because the males of certain species, for instance,
of the robin, sing during the autumn.91 But nothing is more common than
for animals to take pleasure in practising whatever instinct they follow at
other times for some real good. How often do we see birds which fly easily,
gliding and sailing through the air obviously for pleasure. The cat plays
with the captured mouse, and the cormorant with the captured fish. The
weaver-bird (Ploceus), when confined in a cage, amuses itself by neatly
weaving blades of grass between the wires of its cage. Birds which
habitually fight during the breeding-season are generally ready to fight at all
times; and the males of the capercailzie sometimes hold their balzens or
leks at the usual place of assemblage during the autumn.92 Hence it is not at
all surprising that male birds should continue singing for their own
amusement after the season for courtship is over.
Singing is to a certain extent, as shewn in a previous chapter, an art, and is
much improved by practice. Birds can be taught various tunes, and even the

Page 46

unmelodious sparrow has learnt to sing like a linnet. They acquire the song
of their foster-parents,93 and sometimes that of their neighbours.94 All the
common songsters belong to the Order of Insessores, and their vocal organs
are much more complex than those of most other birds; yet it is a singular
fact that some of the Insessores, such as ravens, crows, and magpies,
possess the proper apparatus,95 though they never sing, and do not naturally
modulate their voices to any great extent. Hunter asserts96 that with the true
songsters the muscles of the larynx are stronger in the males than in the
females; but with this slight exception there is no difference in the vocal
organs of the two sexes, although the males of most species sing so much
better and more continuously than the females.
It is remarkable that only small birds properly sing. The Australian genus
Menura, however, must be excepted; for the Menura Alberti, which is about
the size of a half-grown turkey, not only mocks other birds, but “its own
whistle is exceedingly beautiful and varied.” The males congregate and
form “corroborying places,” where they sing, raising and spreading their
tails like peacocks and drooping their wings.97 It is also remarkable that the
birds which sing are rarely decorated with brilliant colours or other
ornaments. Of our British birds, excepting the bullfinch and goldfinch, the
best songsters are plain-coloured. The kingfisher, bee-eater, roller, hoopoe,
woodpeckers, &c., utter harsh cries; and the brilliant birds of the tropics are
hardly ever songsters.98 Hence bright colours and the power of song seem to
replace each other. We can perceive that if the plumage did not vary in
brightness, or if bright colours were dangerous to the species, other means
would have to be employed to charm the females; and the voice being
rendered melodious would offer one such means.

Page 47

Fig. 39. Tetrao cupido; male. (From Brehm.)

In some birds the vocal organs differ greatly in the two sexes. In the Tetrao
cupido (fig. 39) the male has two bare, orange-coloured sacks, one on each
side of the neck; and these are largely inflated when the male, during the
breeding-season, makes a curious hollow sound, audible at a great distance.
Audubon proved that the sound was intimately connected with this
apparatus, which reminds us of the air-sacks on each side of the mouth of
certain male frogs, for he found that the sound was much diminished when
one of the sacks of a tame bird was pricked, and when both were pricked it
was altogether stopped. The female has “a somewhat similar, though
smaller, naked space of skin on the neck; but this is not capable of
inflation.”99 The male of another kind of grouse (Tetrao urophasianus),
whilst courting the female, has his “bare yellow œsophagus inflated to a
prodigious size, fully half as large as the body;” and he then utters various
grating, deep hollow tones. With his neck-feathers erect, his wings lowered
and buzzing on the ground, and his long pointed tail spread out like a fan,
he displays a variety of grotesque attitudes. The œsophagus of the female is
not in any way remarkable.100
It seems now well made out that the great throat-pouch of the European
male bustard (Otis tarda), and of at least four other species, does not serve,
as was formerly supposed, to hold water, but is connected with the utterance
during the breeding-season of a peculiar sound resembling “ock.” The bird
whilst uttering this sound throws himself into the most extraordinary
attitudes. It is a singular fact that with the males of the same species the
sack is not developed in all the individuals.101 A crow-like bird inhabiting
South America (Cephalopterus ornatus, fig. 40) is called the umbrella-bird,
from its immense top-knot, formed of bare white quills surmounted by
dark-blue plumes, which it can elevate into a great dome no less than five
inches in diameter, covering the whole head. This bird has on its neck a
long, thin, cylindrical, fleshy appendage, which is thickly clothed with
scale-like blue feathers. It probably serves in part as an ornament, but
likewise as a resounding apparatus, for Mr. Bates found that it is connected
“with an unusual development of the trachea and vocal organs.” It is dilated
when the bird utters its singularly deep, loud, and long-sustained fluty note.
The head-crest and neck-appendage are rudimentary in the female.102

Page 48

Fig. 40. The Umbrella-bird or Cephalopterus ornatus (male, from Brehm).

The vocal organs of various web-footed and wading birds are
extraordinarily complex, and differ to a certain extent in the two sexes. In
some cases the trachea is convoluted, like a French horn, and is deeply
embedded in the sternum. In the wild swan (Cygnus ferus) it is more deeply
embedded in the adult male than in the female or young male. In the male
Merganser the enlarged portion of the trachea is furnished with an
additional pair of muscles.103 But the meaning of these differences between
the sexes of many Anatidæ is not at all understood; for the male is not
always the more vociferous; thus with the common duck, the male hisses,
whilst the female utters a loud quack.104 In both sexes of one of the cranes
(Grus virgo) the trachea penetrates the sternum, but presents “certain sexual
modifications.” In the male of the black stork there is also a well-marked
sexual difference in the length and curvature of the bronchi.105 So that
highly important structures have in these cases been modified according to
sex.
It is often difficult to conjecture whether the many strange cries and notes,
uttered by male birds during the breeding-season, serve as a charm or
merely as a call to the female. The soft cooing of the turtle-dove and of
many pigeons, it may be presumed, pleases the female. When the female of
the wild turkey utters her call in the morning, the male answers by a
different note from the gobbling noise which he makes, when with erected
feathers, rustling wings and distended wattles, he puffs and struts before
her.106 The spel of the black-cock certainly serves as a call to the female, for
it has been known to bring four or five females from a distance to a male

Page 49

under confinement; but as the black-cock continues his spel for hours
during successive days, and in the case of the capercailzie “with an agony
of passion,” we are led to suppose that the females which are already
present are thus charmed.107 The voice of the common rook is known to
alter during the breeding-season, and is therefore in some way sexual.108 But
what shall we say about the harsh screams of, for instance, some kinds of
macaws; have these birds as bad taste for musical sounds as they apparently
have for colour, judging by the inharmonious contrast of their bright yellow
and blue plumage? It is indeed possible that the loud voices of many male
birds may be the result, without any advantage being thus gained, of the
inherited effects of the continued use of their vocal organs, when they are
excited by the strong passions of love, jealousy, and rage; but to this point
we shall recur when we treat of quadrupeds.
We have as yet spoken only of the voice, but the males of various birds
practise, during their courtship, what may be called instrumental music.
Peacocks and Birds of Paradise rattle their quills together, and the vibratory
movement apparently serves merely to make a noise, for it can hardly add
to the beauty of their plumage. Turkey-cocks scrape their wings against the
ground, and some kinds of grouse thus produce a buzzing sound. Another
North American grouse, the Tetrao umbellus, when with his tail erect, his
ruffs displayed, “he shows off his finery to the females, who lie hid in the
neighbourhood,” drums rapidly with his “lowered wings on the trunk of a
fallen tree,” or, according to Audubon, against his own body; the sound thus
produced is compared by some to distant thunder, and by others to the quick
roll of a drum. The female never drums, “but flies directly to the place
where the male is thus engaged.” In the Himalayas the male of the Kalij
pheasant “often makes a singular drumming noise with his wings, not
unlike the sound produced by shaking a stiff piece of cloth.” On the west
coast of Africa the little black-weavers (Ploceus?) congregate in a small
party on the bushes round a small open space, and sing and glide through
the air with quivering wings, “which make a rapid whirring sound like a
child’s rattle.” One bird after another thus performs for hours together, but
only during the courting-season. At this same season the males of certain
nightjars (Caprimulgus) make a most strange noise with their wings. The
various species of woodpeckers strike a sonorous branch with their beaks,
with so rapid a vibratory movement that “the head appears to be in two

Page 50

places at once.” The sound thus produced is audible at a considerable
distance, but cannot be described; and I feel sure that its cause would never
be conjectured by any one who heard it for the first time. As this jarring
sound is made chiefly during the breeding-season, it has been considered as
a love-song; but it is perhaps more strictly a love-call. The female, when
driven from her nest, has been observed thus to call her mate, who
answered in the same manner and soon appeared. Lastly the male Hoopoe
(Upupa epops) combines vocal and instrumental music; for during the
breeding-season this bird, as Mr. Swinhoe saw, first draws in air and then
taps the end of its beak perpendicularly down against a stone or the trunk of
a tree, “when the breath being forced down the tubular bill produces the
correct sound.” When the male utters its cry without striking his beak the
sound is quite different.109

Fig. 41. Outer tail-feather of Scolopax gallinago (from
Proc. Zool. Soc. 1858).

In the foregoing cases sounds are made by the aid of structures already
present and otherwise necessary; but in the following cases certain feathers
have been specially modified for the express purpose of producing the
sounds. The drumming, or bleating, or neighing, or thundering noise, as
expressed by different observers, which is made by the common snipe
(Scolopax gallinago) must have surprised every one who has ever heard it.
This bird, during the pairing-season, flies to “perhaps a thousand feet in
height,” and after zig-zagging about for a time descends in a curved line,
with outspread tail and quivering pinions, with surprising velocity to the
earth. The sound is emitted only during this rapid descent. No one was able
to explain the cause, until M. Meves observed that on each side of the tail
the outer feathers are peculiarly formed (fig. 41), having a stiff sabre-
shaped shaft, with the oblique barbs of unusual length, the outer webs being
strongly bound together.

Page 51

He found that by blowing on these feathers, or by fastening them to a long
thin stick and waving them rapidly through the air, he could exactly
reproduce the drumming noise made by the living bird. Both sexes are
furnished with these feathers, but they are generally larger in the male than
in the female, and emit a deeper
note. In some species, as in S.
frenata (fig. 42), four feathers, and
in S. javensis (fig. 43), no less than
Fig. 42. Outer tail-feather of Scolopax
eight on each side of the tail are
frenata.
greatly modified. Different tones
are emitted by the feathers of the
different species when waved
through the air; and the Scolopax
Fig. 43. Outer tail-feather of Scolopax
Wilsonii of the United States makes
javensis.
a switching noise whilst descending
rapidly to the earth.110
In the male of the Chamæpetes unicolor (a large gallinaceous bird of
America) the first primary wing-feather is arched towards the tip and is
much more attenuated than in the female. In an allied bird, the Penelope
nigra, Mr. Salvin observed a male, which, whilst it flew downwards “with
outstretched wings, gave forth a kind of crashing, rushing noise,” like the
falling of a tree.111 The male alone of one of the Indian bustards
(Sypheotides auritus) has its primary wing-feathers greatly acuminated; and
the male of an allied species is known to make a humming noise whilst
courting the female.112 In a widely different group of birds, namely the
Humming-birds, the males alone of
certain kinds have either the shafts of
their primary wing-feathers broadly
dilated, or the webs abruptly excised
towards the extremity. The male, for
instance, of Selasphorus platycercus,
when adult, has the first primary wing- Fig. 44. Primary wing-feather of a
feather (fig. 44), excised in this manner.
Humming-bird, the Selasphorus
Whilst flying from flower to flower he platycercus (from a sketch by Mr.
makes “a shrill, almost whistling,
Salvin). Upper figure, that of male; lower
noise;”113 but it did not appear to Mr. figure, corresponding feather of female.

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Salvin that the noise was intentionally made.
Lastly, in several species of a sub-genus of Pipra or Manakin, the males
have their secondary wing-feathers modified, as described by Mr. Sclater, in
a still more remarkable manner. In the brilliantly-coloured P. deliciosa the
first three secondaries are thick-stemmed and curved towards the body; in
the fourth and fifth (fig. 45, a) the change is greater; and in the sixth and
seventh (b, c) the shaft “is thickened to an extraordinary degree, forming a
solid horny lump.” The barbs also are greatly changed in shape, in
comparison with the corresponding feathers (d, e, f) in the female. Even the
bones of the wing which support these singular feathers in the male are said
by Mr. Fraser to be much thickened. These little birds make an
extraordinary noise, the first “sharp note being not unlike the crack of a
whip.”114

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Fig. 45. Secondary wing-feathers of Pipra deliciosa(from Mr. Sclater, in
Proc, Zool. Soc. 1860). The three upper feathers, a, b, c, from the male;
the three lower corresponding feathers, d, e, f, from the female.

a. and d. Fifth secondary wing-feather of male and female, upper surface.
b and e. Sixth secondary, upper surface. c and f. Seventh secondary,
lower surface.

The diversity of the sounds, both vocal and instrumental, made by the males
of many species during the breeding-season, and the diversity of the means
for producing such sounds, are highly remarkable. We thus gain a high idea
of their importance for sexual purposes, and are reminded of the same
conclusion with respect to insects. It is not difficult to imagine the steps by
which the notes of a bird, primarily used as a mere call or for some other
purpose, might have been improved into a melodious love-song. This is
somewhat more difficult in the case of the modified feathers, by which the
drumming, whistling, or roaring noises are produced. But we have seen that
some birds during their courtship flutter, shake, or rattle their unmodified
feathers together; and if the females were led to select the best performers,
the males which possessed the strongest or thickest, or most attenuated
feathers, situated on any part of the body, would be the most successful; and
thus by slow degrees the feathers might be modified to almost any extent.
The females, of course, would not notice each slight successive alteration in
shape, but only the sounds thus produced. It is a curious fact that in the
same class of animals, sounds so different as the drumming of the snipe’s
tail, the tapping of the woodpecker’s beak, the harsh trumpet-like cry of
certain water-fowl, the cooing of the turtle-dove, and the song of the
nightingale, should all be pleasing to the females of the several species. But
we must not judge the tastes of distinct species by a uniform standard; nor
must we judge by the standard of man’s taste. Even with man, we should
remember what discordant noises, the beating of tom-toms and the shrill
notes of reeds, please the ears of savages. Sir S. Baker remarks,115 that “as
the stomach of the Arab prefers the raw meat and reeking liver taken hot
from the animal, so does his ear prefer his equally coarse and discordant
music to all other.”
Love-Antics and Dances.—The curious love-gestures of various birds,
especially of the Gallinaceæ, have already been incidentally noticed; so that
little need here be added. In Northern America, large numbers of a grouse,
the Tetrao phasianellus, meet every morning during the breeding-season on

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a selected level spot, and here they run round and round in a circle of about
fifteen or twenty feet in diameter, so that the ground is worn quite bare, like
a fairy-ring. In these Partridge-dances, as they are called by the hunters, the
birds assume the strangest attitudes, and run round, some to the left and
some to the right. Audubon describes the males of a heron (Ardea herodias)
as walking about on their long legs with great dignity before the females,
bidding defiance to their rivals. With one of the disgusting carrion-vultures
(Cathartes jota) the same naturalist states that “the gesticulations and
parade of the males at the beginning of the love-season are extremely
ludicrous.” Certain birds perform their love-antics on the wing, as we have
seen with the black African weaver, instead of on the ground. During the
spring our little whitethroat (Sylvia cinerea) often rises a few feet or yards
in the air above some bush, and “flutters with a fitful and fantastic motion,
singing all the while, and then drops to its perch.” The great English bustard
throws himself into indescribably odd attitudes whilst courting the female,
as has been figured by Wolf. An allied Indian bustard (Otis bengalensis) at
such times “rises perpendicularly into the air with a hurried flapping of his
wings, raising his crest and puffing out the feathers of his neck and breast,
and then drops to the ground;” he repeats this manœuvre several times
successively, at the same time humming in a peculiar tone. Such females as
happen to be near “obey this saltatory summons,” and when they approach
he trails his wings and spreads his tail like a turkey-cock.116

But the most curious case is afforded by three allied genera of Australian
birds, the famous Bower-birds,—no doubt the co-descendants of some
ancient species which first acquired the strange instinct of constructing
bowers for performing their love-antics. The bowers (fig. 46), which, as we
shall hereafter see, are highly decorated with feathers, shells, bones and
leaves, are built on the ground for the sole purpose of courtship, for their
nests are formed in trees. Both sexes assist in the erection of the bowers, but
the male is the principal workman. So strong is this instinct that it is
practised under confinement, and Mr. Strange has described117 the habits of
some Satin Bower-birds, which he kept in his aviary in New South Wales.
“At times the male will chase the female all over the aviary, then go to the
bower, pick up a gay feather or a large leaf, utter a curious kind of note, set
all his feathers erect, run round the bower and become so excited that his
eyes appear ready to start from his head; he continues opening first one

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wing, and then the other, uttering a low, whistling note, and, like the
domestic cock, seems to be picking up something from the ground, until at
last the female goes gently towards him.” Captain Stokes has described the
habits and “play-houses” of another species, the Great Bower-bird, which
was seen “amusing itself by flying backwards and forwards, taking a shell
alternately from each side, and carrying it through the archway in its
mouth.” These curious structures, formed solely as halls of assemblages,
where both sexes amuse themselves and pay their court, must cost the birds
much labour. The bower, for instance, of the fawn-breasted species, is
nearly four feet in length, eighteen inches in height, and is raised on a thick
platform of sticks.

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Fig. 46. Bower-bird, Chlamydera maculata, with bower (from Brehm).

Decoration.—I will first discuss the cases in which the males are
ornamented either exclusively or in a much higher degree than the females;
and in a succeeding chapter those in which both sexes are equally
ornamented, and finally the rare cases in which the female is somewhat
more brightly-coloured than the male. As with the artificial ornaments used
by savage and civilised men, so with the natural ornaments of birds, the
head is the chief seat of decoration.118 The ornaments, as mentioned at the
commencement of this chapter, are wonderfully diversified. The plumes on
the front or back of the head consist of variously-shaped feathers,
sometimes capable of erection or expansion, by which their beautiful
colours are fully displayed. Elegant ear-tufts (see fig. 39 ante) are
occasionally present. The head is sometimes covered with velvety down
like that of the pheasant; or is naked and vividly coloured; or supports
fleshy appendages, filaments, and solid protuberances. The throat, also, is
sometimes ornamented with a beard, or with wattles or caruncles. Such
appendages are generally brightly coloured, and no doubt serve as
ornaments, though not always ornamental in our eyes; for whilst the male is
in the act of courting the female, they often swell and assume more vivid
tints, as in the case of the male turkey. At such times the fleshy appendages
about the head of the male Tragopan pheasant (Ceriornis temminckii) swell
into a large lappet on the throat and into two horns, one on each side of the
splendid top-knot; and these are then coloured of the most intense blue
which I have ever beheld. The African hornbill (Bucorax abyssinicus)

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inflates the scarlet bladder-like wattle on its neck, and with its wings
drooping and tail expanded “makes quite a grand appearance.”119 Even the
iris of the eye is sometimes more brightly coloured in the male than in the
female; and this is frequently the case with the beak, for instance, in our
common blackbird. In Buceros corrugatus, the whole beak and immense
casque are coloured more conspicuously in the male than in the female; and
“the oblique grooves upon the sides of the lower mandible are peculiar to
the male sex.”120

The males are often ornamented with elongated feathers or plumes
springing from almost every part of the body. The feathers on the throat and
breast are sometimes developed into beautiful ruffs and collars. The tail-
feathers are frequently increased in length; as we see in the tail-coverts of
the peacock, and in the tail of the Argus pheasant. The body of this latter
bird is not larger than that of a fowl; yet the length from the end of the beak
to the extremity of the tail is no less than five feet three inches.121 The wing-
feathers are not elongated nearly so often as the tail-feathers; for their
elongation would impede the act of flight. Yet the beautifully ocellated
secondary wing-feathers of the male Argus pheasant are nearly three feet in
length; and in a small African nightjar (Cosmetornis vexillarius) one of the
primary wing-feathers, during the breeding-season, attains a length of
twenty-six inches, whilst the bird itself is only ten inches in length. In
another closely-allied genus of nightjars, the shafts of the elongated wing-
feathers are naked, except at the extremity, where there is a disc.122 Again,
in another genus of nightjars, the tail-feathers are even still more
prodigiously developed; so that we see the same kind of ornament gained
by the males of closely-allied birds, through the development of widely
different feathers.
It is a curious fact that the feathers of birds belonging to distinct groups
have been modified in almost exactly the same peculiar manner. Thus the
wing-feathers in one of the above-mentioned nightjars are bare along the
shaft and terminate in a disc; or are, as they are sometimes called, spoon or
racket-shaped. Feathers of this kind occur in the tail of a motmot
(Eumomota superciliaris), of a kingfisher, finch, humming-bird, parrot,
several Indian drongos (Dicrurus and Edolius, in one of which the disc
stands vertically), and in the tail of certain Birds of Paradise. In these latter

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birds, similar feathers, beautifully ocellated, ornament the head, as is
likewise the case with some gallinaceous birds. In an Indian bustard
(Sypheotides auritus) the feathers forming the ear-tufts, which are about
four inches in length, also terminate in discs.123 The barbs of the feathers in
various widely-distinct birds are filamentous or plumose, as with some
Herons, Ibises, Birds of Paradise and Gallinaceæ. In other cases the barbs
disappear, leaving the shafts bare; and these in the tail of the Paradisea
apoda attain a length of thirty-four inches.124 Smaller feathers when thus
denuded appear like bristles, as on the breast of the turkey-cock. As any
fleeting fashion in dress comes to be admired by man, so with birds a
change of almost any kind in the structure or colouring of the feathers in the
male appears to have been admired by the female. The fact of the feathers
in widely distinct groups, having been modified in an analogous manner, no
doubt depends primarily on all the feathers having nearly the same structure
and manner of development, and consequently tending to vary in the same
manner. We often see a tendency to analogous variability in the plumage of
our domestic breeds belonging to distinct species. Thus top-knots have
appeared in several species. In an extinct variety of the turkey, the top-knot
consisted of bare quills surmounted with plumes of down, so that they
resembled, to a certain extent, the racket-shaped feathers above described.
In certain breeds of the pigeon and fowl the feathers are plumose, with
some tendency in the shafts to be naked. In the Sebastopol goose the
scapular feathers are greatly elongated, curled, or even spirally twisted, with
the margins plumose.125

Fig. 47. Paradisea rubra, male (from Brehm).

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Fig. 48. Lophornis ornatus, male and female (from Brehm).

In regard to colour hardly anything need here be said; for every one knows
how splendid are the tints of birds, and how harmoniously they are
combined. The colours are often metallic and iridescent. Circular spots are
sometimes surrounded by one or more differently shaded zones, and are
thus converted into ocelli. Nor need much be said on the wonderful
differences between the sexes, or of the extreme beauty of the males of
many birds. The common peacock offers a striking instance. Female Birds
of Paradise are obscurely coloured and destitute of all ornaments, whilst the
males are probably the most highly decorated of all birds, and in so many
ways, that they must be seen to be appreciated. The elongated and golden-
orange plumes which spring from beneath the wings of the Paradisea
apoda (see fig. 47 of P. rubra, a much less beautiful species), when
vertically erected and made to vibrate, are described as forming a sort of
halo, in the centre of which the head “looks like a little emerald sun with its
rays formed by the two plumes.”126 In another most beautiful species the
head is bald, “and of a rich cobalt blue, crossed by several lines of black
velvety feathers.”127

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Fig. 49. Spathura underwoodi, male and female (from Brehm).

Male humming-birds (figs. 48 and 49) almost vie with Birds of Paradise in
their beauty, as every one will admit who has seen Mr. Gould’s splendid
volumes or his rich collection. It is very remarkable in how many different
ways these birds are ornamented. Almost every part of the plumage has
been taken advantage of and modified; and the modifications have been
carried, as Mr. Gould shewed me, to a wonderful extreme in some species
belonging to nearly every subgroup. Such cases are curiously like those
which we see in our fancy breeds, reared by man for the sake of ornament:
certain individuals originally varied in one character, and other individuals
belonging to the same species in other characters; and these have been
seized on by man and augmented to an extreme point—as the tail of the
fantail-pigeon, the hood of the jacobin, the beak and wattle of the carrier,
and so forth. The sole difference between these cases is that in the one the
result is due to man’s selection, whilst in the other, as with Humming-birds,
Birds of Paradise, &c., it is due to sexual selection,—that is to the selection
by the females of the more beautiful males.
I will mention only one other bird, remarkable from the extreme contrast in
colour between the sexes, namely the famous Bell-bird (Chasmorhynchus
niveus) of S. America, the note of which can be distinguished at the distance
of nearly three miles, and astonishes every one who first hears it. The male

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is pure white, whilst the female is dusky-green; and the former colour with
terrestrial species of moderate size and inoffensive habits is very rare. The
male, also, as described by Waterton, has a spiral tube, nearly three inches
in length, which rises from the base of the beak. It is jet-black, dotted over
with minute downy feathers. This tube can be inflated with air, through a
communication with the palate; and when not inflated hangs down on one
side. The genus consists of four species, the males of which are very
distinct, whilst the females, as described by Mr. Sclater in a most interesting
paper, closely resemble each other, thus offering an excellent instance of the
common rule that within the same group the males differ much more from
each other than do the females. In a second species (C. nudicollis) the male
is likewise snow-white, with the exception of a large space of naked skin on
the throat and round the eyes, which during the breeding-season is of a fine
green colour. In a third species (C. tricarunculatus) the head and neck alone
of the male are white, the rest of the body being chesnut-brown, and the
male of this species is provided with three filamentous projections half as
long as the body—one rising from the base of the beak and the two others
from the corners of the mouth.128
The coloured plumage and certain other ornaments of the males when adult
are either retained for life or are periodically renewed during the summer
and breeding-season. At this season the beak and naked skin about the head
frequently change colour, as with some herons, ibises, gulls, one of the bell-
birds just noticed, &c. In the white ibis, the cheeks, the inflatable skin of the
throat, and the basal portion of the beak, then become crimson.129 In one of
the rails, Gallicrex cristatus a large red caruncle is developed during this
same period on the head of the male. So it is with a thin horny crest on the
beak of one of the pelicans, P. erythrorhynchus; for after the breeding-
season, these horny crests are shed, like horns from the heads of stags, and
the shore of an island in a lake in Nevada was found covered with these
curious exuviæ.130
Changes of colour in the plumage according to the season depend firstly on
a double annual moult, secondly on an actual change of colour in the
feathers themselves, and thirdly on their dull-coloured margins being
periodically shed, or on these three processes more or less combined. The
shedding of the deciduary margins may be compared with the shedding by

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very young birds of their down; for the down in most cases arises from the
summits of the first true feathers.131
With respect to the birds which annually undergo a double moult, there are,
firstly, some kinds, for instance snipes, swallow-plovers (Glareolæ), and
curlews, in which the two sexes resemble each other and do not change
colour at any season. I do not know whether the winter-plumage is thicker
and warmer than the summer-plumage, which seems, when there is no
change of colour, the most probable cause of a double moult. Secondly,
there are birds, for instance certain species of Totanus and other grallatores,
the sexes of which resemble each other, but have a slightly different
summer and winter plumage. The difference, however, in colour in these
cases is so slight that it can hardly be an advantage to them; and it may,
perhaps, be attributed to the direct action of the different conditions to
which the birds are exposed during the two seasons. Thirdly, there are many
other birds the sexes of which are alike, but which are widely different in
their summer and winter plumage. Fourthly, there are birds, the sexes of
which differ from each other in colour; but the females, though moulting
twice, retain the same colours throughout the year, whilst the males undergo
a change, sometimes, as with certain bustards, a great change of colour.
Fifthly and lastly, there are birds the sexes of which differ from each other
in both their summer and winter plumage, but the male undergoes a greater
amount of change at each recurrent season than the female—of which the
Ruff (Machetes pugnax) offers a good instance.
With respect to the cause or purpose of the differences in colour between
the summer and winter plumage, this may in some instances, as with the
ptarmigan,132 serve during both seasons as a protection. When the difference
between the two plumages is slight it may perhaps be attributed, as already
remarked, to the direct action of the conditions of life. But with many birds
there can hardly be a doubt that the summer plumage is ornamental, even
when both sexes are alike. We may conclude that this is the case with many
herons, egrets, &c., for they acquire their beautiful plumes only during the
breeding-season. Moreover, such plumes, top-knots, &c., though possessed
by both sexes, are occasionally a little more highly developed in the male
than in the female; and they resemble the plumes and ornaments possessed
by the males alone of other birds. It is also known that confinement, by
affecting the reproductive system of male birds, frequently checks the

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development of their secondary sexual characters, but has no immediate
influence on any other characters; and I am informed by Mr. Bartlett that
eight or nine specimens of the Knot (Tringa canutus) retained their
unadorned winter plumage in the Zoological Gardens throughout the year,
from which fact we may infer that the summer plumage though common to
both sexes partakes of the nature of the exclusively masculine plumage of
many other birds.133
From the foregoing facts, more especially from neither sex of certain birds
changing colour during either annual moult, or changing so slightly that the
change can hardly be of any service to them, and from the females of other
species moulting twice yet retaining the same colours throughout the year,
we may conclude that the habit of moulting twice in the year has not been
acquired in order that the male should assume during the breeding-season
an ornamental character; but that the double moult, having been originally
acquired for some distinct purpose, has subsequently been taken advantage
of in certain cases for gaining a nuptial plumage.
It appears at first sight a surprising circumstance that with closely-allied
birds, some species should regularly undergo a double annual moult, and
others only a single one. The ptarmigan, for instance, moults twice or even
thrice in the year, and the black-cock only once: some of the splendidly-
coloured honey-suckers (Nectariniæ) of India and some sub-genera of
obscurely-coloured pipits (Anthus) have a double, whilst others have only a
single annual moult.134 But the gradations in the manner of moulting, which
are known to occur with various birds, shew us how species, or whole
groups of species, might have originally acquired their double annual
moult, or having once gained the habit, have again lost it. With certain
bustards and plovers the vernal moult is far from complete, some feathers
being renewed, and some changed in colour. There is also reason to believe
that with certain bustards and rail-like birds, which properly undergo a
double moult, some of the older males retain their nuptial plumage
throughout the year. A few highly modified feathers may alone be added
during the spring to the plumage, as occurs with the disc-formed tail-
feathers of certain drongos (Bhringa) in India, and with the elongated
feathers on the back, neck, and crest of certain herons. By such steps as
these, the vernal moult might be rendered more and more complete, until a
perfect double moult was acquired. A gradation can also be shewn to exist

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in the length of time during which either annual plumage is retained; so that
the one might come to be retained for the whole year, the other being
completely lost. Thus the Machetes pugnax retains his ruff in the spring for
barely two months. The male widow-bird (Chera progne) acquires in Natal
his fine plumage and long tail-feathers in December or January and loses
them in March; so that they are retained during only about three months.
Most species which undergo a double moult keep their ornamental feathers
for about six months. The male, however, of the wild Gallus bankiva retains
his neck-hackles for nine or ten months; and when these are cast off, the
underlying black feathers on the neck are fully exposed to view. But with
the domesticated descendant of this species, the neck-hackles of the male
are immediately replaced by new ones; so that we here see, with respect to
part of the plumage, a double moult changed under domestication into a
single moult.135
The common drake (Anas boschas) is well known after the breeding-season
to lose his male plumage for a period of three months, during which time he
assumes that of the female. The male pintail-duck (Anas acuta) loses his
plumage for the shorter period of six weeks or two months; and Montagu
remarks that “this double moult within so short a time is a most
extraordinary circumstance, that seems to bid defiance to all human
reasoning.” But he who believes in the gradual modification of species will
be far from feeling surprise at finding gradations of all kinds. If the male
pintail were to acquire his new plumage within a still shorter period, the
new male feathers would almost necessarily be mingled with the old, and
both with some proper to the female; and this apparently is the case with the
male of a not distantly-allied bird, namely the Merganser serrator, for the
males are said to “undergo a change of plumage, which assimilates them in
some measure to the female.” By a little further acceleration in the process,
the double moult would be completely lost.136
Some male birds, as before stated, become more brightly coloured in the
spring, not by a vernal moult, but either by an actual change of colour in the
feathers, or by their obscurely-coloured deciduary margins being shed.
Changes of colour thus caused may last for a longer or shorter time. With
the Pelecanus onocrotalus a beautiful rosy tint, with lemon-coloured marks
on the breast, overspreads the whole plumage in the spring; but these tints,
as Mr. Sclater states, “do not last long, disappearing generally in about six

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weeks or two months after they have been attained.” Certain finches shed
the margins of their feathers in the spring, and then become brighter-
coloured, while other finches undergo no such change. Thus the Fringilla
tristis of the United States (as well as many other American species),
exhibits its bright colours only when the winter is past, whilst our
goldfinch, which exactly represents this bird in habits, and our siskin, which
represents it still more closely in structure, undergo no such annual change.
But a difference of this kind in the plumage of allied species is not
surprising, for with the common linnet, which belongs to the same family,
the crimson forehead and breast are displayed only during the summer in
England, whilst in Madeira these colours are retained throughout the year.137
Display by Male Birds of their Plumage.—Ornaments of all kinds, whether
permanently or temporarily gained, are sedulously displayed by the males,
and apparently serve to excite, or attract, or charm the females. But the
males will sometimes display their ornaments, when not in the presence of
the females, as occasionally occurs with grouse at their balz-places, and as
may be noticed with the peacock; this latter bird, however, evidently wishes
for a spectator of some kind, and will shew off his finery, as I have often
seen, before poultry or even pigs.138 All naturalists who have closely
attended to the habits of birds, whether in a state of nature or under
confinement, are unanimously of opinion that the males delight to display
their beauty. Audubon frequently speaks of the male as endeavouring in
various ways to charm the female. Mr. Gould, after describing some
peculiarities in a male humming-bird, says he has no doubt that it has the
power of displaying them to the greatest advantage before the female. Dr.
Jerdon139 insists that the beautiful plumage of the male serves “to fascinate
and attract the female.” Mr. Bartlett, at the Zoological Gardens, expressed
himself to me in the strongest terms to the same effect.

It must be a grand sight in the forests of India “to come suddenly on twenty
or thirty peafowl, the males displaying their gorgeous trains, and strutting
about in all the pomp of pride before the gratified females.” The wild
turkey-cock erects his glittering plumage, expands his finely-zoned tail and
barred wing-feathers, and altogether, with his gorged crimson and blue
wattles, makes a superb, though, to our eyes, grotesque appearance. Similar
facts have already been given with respect to grouse of various kinds.

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Turning to another Order. The male Rupicola crocea (fig. 50) is one of the
most beautiful birds in the world, being of a splendid orange, with some of
the feathers curiously truncated and plumose. The female is brownish-
green, shaded with red, and has a much smaller crest. Sir R. Schomburgk
has described their courtship; he found one of their meeting-places where
ten males and two females were present. The space was from four to five
feet in diameter, and appeared to have been cleared of every blade of grass
and smoothed as if by human hands. A male “was capering to the apparent
delight of several others. Now spreading its wings, throwing up its head, or
opening its tail like a fan; now strutting about with a hopping gait until
tired, when it gabbled some kind of note, and was relieved by another. Thus
three of them successively took the field, and then, with self-approbation,
withdrew to rest.” The Indians, in order to obtain their skins, wait at one of
the meeting-places till the birds are eagerly engaged in dancing, and then
are able to kill, with their poisoned arrows, four or five males, one after the
other.140 With Birds of Paradise a dozen or more full-plumaged males
congregate in a tree to hold a dancing-party, as it is called by the natives;
and here flying about, raising their wings, elevating their exquisite plumes,
and making them vibrate, the whole tree seems, as Mr. Wallace remarks, to
be filled with waving plumes. When thus engaged, they become so
absorbed that a skilful archer may shoot nearly the whole party. These birds,
when kept in confinement in the Malay Archipelago, are said to take much
care in keeping their feathers clean; often spreading them out, examining
them, and removing every speck of dirt. One observer, who kept several
pairs alive, did not doubt that the display of the male was intended to please
the female.141

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Fig. 50. Rupicola crocea, male (from Brehm).

The gold pheasant (Thaumalea picta) during his courtship not only expands
and raises his splendid frill, but turns it, as I have myself seen, obliquely
towards the female on whichever side she may be standing, obviously in
order that a large surface may be displayed before her.142 Mr. Bartlett has
observed a male Polyplectron (fig. 51) in the act of courtship, and has
shewn me a specimen stuffed in the attitude then assumed. The tail and
wing-feathers of this bird are ornamented with beautiful ocelli, like those on
the peacock’s train. Now when the peacock displays himself, he expands
and erects his tail transversely to his body, for he stands in front of the
female, and has to shew off, at the same time, his rich blue throat and
breast. But the breast of the Polyplectron is obscurely coloured, and the
ocelli are not confined to the tail-feathers. Consequently the Polyplectron
does not stand in front of the female; but he erects and expands his tail-
feathers a little obliquely, lowering the expanded wing on the same side,
and raising that on the opposite side. In this attitude the ocelli over the
whole body are exposed before the eyes of the admiring female in one
grand bespangled expanse. To whichever side she may turn, the expanded
wings and the obliquely-held tail are turned towards her. The male
Tragopan pheasant acts in nearly the same manner, for he raises the feathers
of the body, though not the wing itself, on the side which is opposite to the
female, and which would otherwise be concealed, so that nearly all the
beautifully-spotted feathers are exhibited at the same time.

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Fig. 51. Polyplectron chinquis. male (from Brehm.)

The case of the Argus pheasant is still more striking. The immensely
developed secondary wing-feathers, which are confined to the male, are
ornamented with a row of from twenty to twenty-three ocelli, each above an
inch in diameter. The feathers are also elegantly marked with oblique dark
stripes and rows of spots, like those on the skin of a tiger and leopard
combined. The ocelli are so beautifully shaded that, as the Duke of Argyll
remarks,143 they stand out like a ball lying loosely within a socket. But when
I looked at the specimen in the British Museum, which is mounted with the
wings expanded and trailing downwards, I was greatly disappointed, for the
ocelli appeared flat or even concave. Mr. Gould, however, soon made the
case clear to me, for he had made a drawing of a male whilst he was
displaying himself. At such times the long secondary feathers in both wings
are vertically erected and expanded; and these, together with the
enormously elongated tail-feathers, make a grand semicircular upright fan.
Now as soon as the wing-feathers are held in this position, and the light
shines on them from above, the full effect of the shading comes out, and
each ocellus at once resembles the ornament called a ball and socket. These
feathers have been shewn to several artists, and all have expressed their
admiration at the perfect shading.
It may well be asked, could such artistically-shaded ornaments have been
formed by means of sexual selection? But it will be convenient to defer
giving an answer to this question until we treat in the next chapter of the
principle of gradation.

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The primary wing-feathers, which in most gallinaceous birds are uniformly
coloured, are in the Argus pheasant not less wonderful objects than the
secondary wing-feathers. They are of a soft brown tint with numerous dark
spots, each of which consists of two or three black dots with a surrounding
dark zone. But the chief ornament is a space parallel to the dark-blue shaft,
which in outline forms a perfect second feather lying within the true feather.
This inner part is coloured of a lighter chesnut, and is thickly dotted with
minute white points. I have shewn this feather to several persons, and many
have admired it even more than the ball-and-socket feathers, and have
declared that it was more like a work of art than of nature. Now these
feathers are quite hidden on all ordinary occasions, but are fully displayed
when the long secondary feathers are erected, though in a widely different
manner; for they are expanded in front like two little fans or shields, one on
each side of the breast near the ground.
The case of the male Argus pheasant is eminently interesting, because it
affords good evidence that the most refined beauty may serve as a charm
for the female, and for no other purpose. We must conclude that this is the
case, as the primary wing-feathers are never displayed, and the ball-and-
socket ornaments are not exhibited in full perfection, except when the male
assumes the attitude of courtship. The Argus pheasant does not possess
brilliant colours, so that his success in courtship appears to have depended
on the great size of his plumes, and on the elaboration of the most elegant
patterns. Many will declare that it is utterly incredible that a female bird
should be able to appreciate fine shading and exquisite patterns. It is
undoubtedly a marvellous fact that she should possess this almost human
degree of taste, though perhaps she admires the general effect rather than
each separate detail. He who thinks that he can safely gauge the
discrimination and taste of the lower animals, may deny that the female
Argus pheasant can appreciate such refined beauty; but he will then be
compelled to admit that the extraordinary attitudes assumed by the male
during the act of courtship, by which the wonderful beauty of his plumage
is fully displayed, are purposeless; and this is a conclusion which I for one
will never admit.
Although so many pheasants and allied gallinaceous birds carefully display
their beautiful plumage before the females, it is remarkable, as Mr. Bartlett
informs me, that this is not the case with the dull-coloured Eared and Cheer

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pheasants (Crossoptilon auritum and Phasianus Wallichii); so that these
birds seem conscious that they have little beauty to display. Mr. Bartlett has
never seen the males of either of these species fighting together, though he
has not had such good opportunities for observing the Cheer as the Eared
pheasant. Mr. Jenner Weir, also, finds that all male birds with rich or
strongly-characterised plumage are more quarrelsome than the dull-
coloured species belonging to the same groups. The goldfinch, for instance,
is far more pugnacious than the linnet, and the blackbird than the thrush.
Those birds which undergo a seasonal change of plumage likewise become
much more pugnacious at the period when they are most gaily ornamented.
No doubt the males of some obscurely-coloured birds fight desperately
together, but it appears that when sexual selection has been highly
influential, and has given bright colours to the males of any species, it has
also very often given a strong tendency to pugnacity. We shall meet with
nearly analogous cases when we treat of mammals. On the other hand, with
birds the power of song and brilliant colours have rarely been both acquired
by the males of the same species; but in this case, the advantage gained
would have been identically the same, namely success in charming the
female. Nevertheless it must be owned that the males of several brilliantly-
coloured birds have had their feathers specially modified for the sake of
producing instrumental music, though the beauty of this cannot be
compared, at least according to our taste, with that of the vocal music of
many songsters.
We will now turn to male birds which are not ornamented in any very high
degree, but which nevertheless display, during their courtship, whatever
attractions they may possess. These cases are in some respects more curious
than the foregoing, and have been but little noticed. I owe the following
facts, selected from a large body of valuable notes, sent to me by Mr. Jenner
Weir, who has long kept birds of many kinds, including all the British
Fringillidæ and Emberizidæ. The bullfinch makes his advances in front of
the female, and then puffs out his breast, so that many more of the crimson
feathers are seen at once than otherwise would be the case. At the same time
he twists and bows his black tail from side to side in a ludicrous manner.
The male chaffinch also stands in front of the female, thus shewing his red
breast, and “blue bell,” as the fanciers call his head; the wings at the same
time being slightly expanded, with the pure white bands on the shoulders

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thus rendered conspicuous. The common linnet distends his rosy breast,
slightly expands his brown wings and tail, so as to make the best of them by
exhibiting their white edgings. We must, however, be cautious in
concluding that the wings are spread out solely for display, as some birds
act thus whose wings are not beautiful. This is the case with the domestic
cock, but it is always the wing on the side opposite to the female which is
expanded, and at the same time scraped on the ground. The male goldfinch
behaves differently from all other finches: his wings are beautiful, the
shoulders being black, with the dark-tipped wing-feathers spotted with
white and edged with golden yellow. When he courts the female, he sways
his body from side to side, and quickly turns his slightly expanded wings
first to one side then to the other, with a golden flashing effect. No other
British finch, as Mr. Weir informs me, turns during his courtship from side
to side in this manner; not even the closely-allied male siskin, for he would
not thus add to his beauty.
Most of the British Buntings are plain-coloured birds; but in the spring the
feathers on the head of the male reed-bunting (Emberiza schœniculus)
acquire a fine black colour by the abrasion of the dusky tips; and these are
erected during the act of courtship. Mr. Weir has kept two species of
Amadina from Australia: the A. castanotis is a very small and chastely-
coloured finch, with a dark tail, white rump, and jet-black upper tail-
coverts, each of the latter being marked with three large conspicuous oval
spots of white.144 This species, when courting the female, slightly spreads
out and vibrates these parti-coloured tail-coverts in a very peculiar manner.
The male Amadina Lathami behaves very differently, exhibiting before the
female his brilliantly-spotted breast and scarlet rump and scarlet upper tail-
coverts. I may here add from Dr. Jerdon, that the Indian Bulbul (Pycnonotus
hæmorrhous) has crimson under tail-coverts, and the beauty of these
feathers, it might be thought, could never be well exhibited; but the bird
“when excited often spreads them out laterally, so that they can be seen
even from above.”145 The common pigeon has iridescent feathers on the
breast, and every one must have seen how the male inflates his breast whilst
courting the female, thus showing off these feathers to the best advantage.
One of the beautiful bronze-winged pigeons of Australia (Ocyphaps
lophotes) behaves, as described to me by Mr. Weir, very differently: the
male, whilst standing before the female, lowers his head almost to the

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ground, spreads out and raises perpendicularly his tail, and half expands his
wings. He then alternately and slowly raises and depresses his body, so that
the iridescent metallic feathers are all seen at once, and glitter in the sun.
Sufficient facts have now been given to shew with what care male birds
display their various charms, and this they do with the utmost skill. Whilst
preening their feathers, they have frequent opportunities for admiring
themselves and of studying how best to exhibit their beauty. But as all the
males of the same species display themselves in exactly the same manner, it
appears that actions, at first perhaps intentional, have become instinctive. If
so, we ought not to accuse birds of conscious vanity; yet when we see a
peacock strutting about, with expanded and quivering tail-feathers, he
seems the very emblem of pride and vanity.
The various ornaments possessed by the males are certainly of the highest
importance to them, for they have been acquired in some cases at the
expense of greatly impeded powers of flight or of running. The African
nightjar (Cosmetornis), which during the pairing-season has one of its
primary wing-feathers developed into a streamer of extreme length, is thus
much retarded in its flight, although at other times remarkable for its
swiftness. The “unwieldy size” of the secondary wing-feathers of the male
Argus pheasant are said “almost entirely to deprive the bird of flight.” The
fine plumes of male Birds of Paradise trouble them during a high wind. The
extremely long tail-feathers of the male widow-birds (Vidua) of Southern
Africa render “their flight heavy;” but as soon as these are cast off they fly
as well as the females. As birds always breed when food is abundant, the
males probably do not suffer much inconvenience in searching for food
from their impeded powers of movement; but there can hardly be a doubt
that they must be much more liable to be struck down by birds of prey. Nor
can we doubt that the long train of the peacock and the long tail and wing-
feathers of the Argus pheasant must render them a more easy prey to any
prowling tiger-cat than would otherwise be the case. Even the bright colours
of many male birds cannot fail to make them conspicuous to their enemies
of all kinds. Hence it probably is, as Mr. Gould has remarked, that such
birds are generally of a shy disposition, as if conscious that their beauty was
a source of danger, and are much more difficult to discover or approach,
than the sombre-coloured and comparatively tame females, or than the
young and as yet unadorned males.146

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It is a more curious fact that the males of some birds which are provided
with special weapons for battle, and which in a state of nature are so
pugnacious that they often kill each other, suffer from possessing certain
ornaments. Cock-fighters trim the hackles and cut off the comb and gills of
their cocks; and the birds are then said to be dubbed. An undubbed bird, as
Mr. Tegetmeier insists, “is at a fearful disadvantage: the comb and gills
offer an easy hold to his adversary’s beak, and as a cock always strikes
where he holds, when once he has seized his foe, he has him entirely in his
power. Even supposing that the bird is not killed, the loss of blood suffered
by an undubbed cock is much greater than that sustained by one that has
been trimmed.”147 Young turkey-cocks in fighting always seize hold of each
other’s wattles; and I presume that the old birds fight in the same manner. It
may perhaps be objected that the comb and wattles are not ornamental, and
cannot be of service to the birds in this way; but even to our eyes, the
beauty of the glossy black Spanish cock is much enhanced by his white face
and crimson comb; and no one who has ever seen the splendid blue wattles
of the male Tragopan pheasant, when distended during the act of courtship,
can for a moment doubt that beauty is the object gained. From the foregoing
facts we clearly see that the plumes and other ornaments of the male must
be of the highest importance to him; and we further see that beauty in some
cases is even more important than success in battle.

CHAPTER XIV.

Birds—continued.
Choice exerted by the female—Length of courtship—Unpaired birds—Mental qualities and taste for
the beautiful—Preference or antipathy shewn by the female for particular males—Variability of
birds—Variations sometimes abrupt—Laws of variation—Formation of ocelli—Gradations of
character—Case of Peacock, Argus pheasant, and Urosticte.

When the sexes differ in beauty, in the power of singing, or in producing
what I have called instrumental music, it is almost invariably the male
which excels the female. These qualities, as we have just seen, are evidently
of high importance to the male. When they are gained for only a part of the
year, this is always shortly before the breeding-season. It is the male alone

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who elaborately displays his varied attractions, and often performs strange
antics on the ground or in the air, in the presence of the female. Each male
drives away or, if he can, kills all his rivals. Hence we may conclude, that it
is the object of the male to induce the female to pair with him, and for this
purpose he tries to excite or charm her in various ways; and this is the
opinion of all those who have carefully studied the habits of living birds.
But there remains a question which has an all important bearing on sexual
selection, namely, does every male of the same species equally excite and
attract the female? or does she exert a choice, and prefer certain males? This
question can be answered in the affirmative by much direct and indirect
evidence. It is much more difficult to decide what qualities determine the
choice of the females; but here again we have some direct and indirect
evidence that it is to a large extent the external attractions of the male,
though no doubt his vigour, courage, and other mental qualities come into
play. We will begin with the indirect evidence.
Length of Courtship.—The lengthened period during which both sexes of
certain birds meet day after day at an appointed place, probably depends
partly on the courtship being a prolonged affair, and partly on the reiteration
of the act of pairing. Thus in Germany and Scandinavia the balzens or leks
of the Black-cocks, last from the middle of March, all through April into
May. As many as forty or fifty, or even more birds congregate at the leks;
and the same place is often frequented during successive years. The lek of
the Capercailzie lasts from the end of March to the middle or even end of
May. In North America “the partridge dances” of the Tetrao phasianellus
“last for a month or more.” Other kinds of grouse both in North America
and Eastern Siberia148 follow nearly the same habits. The fowlers discover
the hillocks where the Ruffs congregate by the grass being trampled bare,
and this shews that the same spot is long frequented. The Indians of Guiana
are well acquainted with the cleared arenas, where they expect to find the
beautiful Cocks of the Rock; and the natives of New Guinea know the trees
where from ten to twenty full-plumaged male Birds of

Paradise congregate. In this latter case it is not expressly stated that the
females meet on the same trees, but the hunters, if not specially asked,
would not probably mention their presence, as their skins are valueless.
Small parties of an African weaver (Ploceus) congregate, during the

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breeding-season, and perform for hours their graceful evolutions. Large
numbers of the Solitary snipe (Scolopax major) assemble during the dusk in
a morass; and the same place is frequented for the same purpose during
successive years; here they may be seen running about “like so many large
rats,” puffing out their feathers, flapping their wings, and uttering the
strangest cries.149
Some of the above-mentioned birds, namely, the black-cock, capercailzie,
pheasant-grouse, the ruff, the Solitary snipe, and perhaps some others, are,
as it is believed, polygamists. With such birds it might have been thought
that the stronger males would simply have driven away the weaker, and
then at once have taken possession of as many females as possible; but if it
be indispensable for the male to excite or please the female, we can
understand the length of the courtship and the congregation of so many
individuals of both sexes at the same spot. Certain species which are strictly
monogamous likewise hold nuptial assemblages; this seems to be the case
in Scandinavia with one of the ptarmigans, and their leks last from the
middle of March to the middle of May. In Australia the lyre-bird or Menura
superba forms “small round hillocks,” and the M. Alberti scratches for itself
shallow holes, or, as they are called by the natives, corroborying places,
where it is believed both sexes assemble. The meetings of the M. superba
are sometimes very large; and an account has lately been published150 by a
traveller, who heard in a valley beneath him, thickly covered with scrub, “a
din which completely astonished” him; on crawling onwards he beheld to
his amazement about one hundred and fifty of the magnificent lyre-cocks,
“ranged in order of battle, and fighting with indescribable fury.” The
bowers of the Bower-birds are the resort of both sexes during the breeding-
season; and “here the males meet and contend with each other “for the
favours of the female, and here the latter assemble and coquet with the
males.” With two of the genera, the same bower is resorted to during many
years.151
The common magpie (Corvus pica, Linn.), as I have been informed by the
Rev. W. Darwin Fox, used to assemble from all parts of Delamere Forest, in
order to celebrate the “great magpie marriage.” Some years ago these birds
abounded in extraordinary numbers, so that a gamekeeper killed in one
morning nineteen males, and another killed by a single shot seven birds at
roost together. Whilst they were so numerous, they had the habit very early

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in the spring of assembling at particular spots, where they could be seen in
flocks, chattering, sometimes fighting, bustling and flying about the trees.
The whole affair was evidently considered by the birds as of the highest
importance. Shortly after the meeting they all separated, and were then
observed by Mr. Fox and others to be paired for the season. In any district
in which a species does not exist in large numbers, great assemblages
cannot, of course, be held, and the same species may have different habits
in different countries. For instance, I have never met with any account of
regular assemblages of black game in Scotland, yet these assemblages are
so well known in Germany and Scandinavia that they have special names.
Unpaired Birds.—From the facts now given, we may conclude that with
birds belonging to widely-different groups their courtship is often a
prolonged, delicate, and troublesome affair. There is even reason to suspect,
improbable as this will at first appear, that some males and females of the
same species, inhabiting the same district, do not always please each other
and in consequence do not pair. Many accounts have been published of
either the male or female of a pair having been shot, and quickly replaced
by another. This has been observed more frequently with the magpie than
with any other bird, owing perhaps to its conspicuous appearance and nest.
The illustrious Jenner states that in Wiltshire one of a pair was daily shot no
less than seven times successively, “but all to no purpose, for the remaining
magpie soon found another mate;” and the last pair reared their young. A
new partner is generally found on the succeeding day; but Mr. Thompson
gives the case of one being replaced on the evening of the same day. Even
after the eggs are hatched, if one of the old birds is destroyed a mate will
often be found; this occurred after an interval of two days, in a case recently
observed by one of Sir J. Lubbock’s keepers.152 The first and most obvious
conjecture is that male magpies must be much more numerous than the
females; and that in the above cases, as well in many others which could be
given, the males alone had been killed. This apparently holds good in some
instances, for the gamekeepers in Delamere Forest assured Mr. Fox that the
magpies and carrion-crows which they formerly killed in succession in
large numbers near their nests were all males; and they accounted for this
fact by the males being easily killed whilst bringing food to the sitting
females. Macgillivray, however, gives, on the authority of an excellent
observer, an instance of three magpies successively killed on the same nest

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which were all females; and another case of six magpies successively killed
whilst sitting on the same eggs, which renders it probable that most of them
were females, though the male will sit on the eggs, as I hear from Mr. Fox,
when the female is killed.
Sir J. Lubbock’s gamekeeper has repeatedly shot, but how many times he
could not say, one of a pair of jays (Garrulus glandarius), and has never
failed shortly afterwards to find the survivor rematched. The Rev. W. D.
Fox, Mr. F. Bond, and others, have shot one of a pair of carrion-crows
(Corvus corone), but the nest was soon again tenanted by a pair. These birds
are rather common; but the peregrine falcon (Falco peregrinus) is rare, yet
Mr. Thompson states that in Ireland “if either an old male or female be
killed in the breeding-season (not an uncommon circumstance), another
mate is found within a very few days, so that the eyries, notwithstanding
such casualties, are sure to turn out their complement of young.” Mr. Jenner
Weir has known the same thing to occur with the peregrine falcons at
Beachy Head. The same observer informs me that three kestrels, all males
(Falco tinnunculus), were killed one after the other whilst attending the
same nest; two of these were in mature plumage, and the third in the
plumage of the previous year. Even with the rare golden eagle (Aquila
chrysaëtos), Mr. Birkbeck was assured by a trustworthy gamekeeper in
Scotland, that if one is killed, another is soon found. So with the white owl
(Strix flammea), it has been observed that “the survivor readily found a
mate, and the mischief went on.”
White of Selborne, who gives the case of the owl, adds that he knew a man,
who from believing that partridges when paired were disturbed by the males
fighting, used to shoot them; and though he had widowed the same female
several times she was always soon provided with a fresh partner. This same
naturalist ordered the sparrows, which deprived the house-martins of their
nests, to be shot: but the one which was left, “be it cock or hen, presently
procured a mate, and so for several times following.” I could add analogous
cases relating to the chaffinch, nightingale, and redstart. With respect to the
latter bird (Phœnicura ruticilla), the writer remarks that it was by no means
common in the neighbourhood, and he expresses much surprise how the
sitting female could so soon give effectual notice that she was a widow. Mr.
Jenner Weir has mentioned to me a nearly similar case: at Blackheath he
never sees or hears the note of the wild bullfinch, yet when one of his caged

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males has died, a wild one in the course of a few days has generally come
and perched near the widowed female, whose call-note is far from loud. I
will give only one other fact, on the authority of this same observer; one of
a pair of starlings (Sturnus vulgaris) was shot in the morning; by noon a
new mate was found; this was again shot, but before night the pair was
complete; so that the disconsolate widow or widower was thrice consoled
during the same day. Mr. Engleheart also informs me that he used during
several years to shoot one of a pair of starlings which built in a hole in a
house at Blackheath; but the loss was always immediately repaired. During
one season he kept an account and found that he had shot thirty-five birds
from the same nest; these consisted of both males and females, but in what
proportion he could not say: nevertheless after all this destruction, a brood
was reared.153
These facts are certainly remarkable. How is it that so many birds are ready
immediately to replace a lost mate? Magpies, jays, carrion-crows,
partridges, and some other birds, are never seen during the spring by
themselves, and these offer at first sight the most perplexing case. But birds
of the same sex, although of course not truly paired, sometimes live in pairs
or in small parties, as is known to be the case with pigeons and partridges.
Birds also sometimes live in triplets, as has been observed with starlings,
carrion-crows, parrots, and partridges. With partridges two females have
been known to live with one male, and two males with one female. In all
such cases it is probable that the union would be easily broken. The males
of certain birds may occasionally be heard pouring forth their love-song
long after the proper time, shewing that they have either lost or never
gained a mate. Death from accident or disease of either one of a pair, would
leave the other bird free and single; and there is reason to believe that
female birds during the breeding-season are especially liable to premature
death. Again, birds which have had their nests destroyed, or barren pairs, or
retarded individuals, would easily be induced to desert their mates, and
would probably be glad to take what share they could of the pleasures and
duties of rearing offspring, although not their own.154 Such contingencies as
these probably explain most of the foregoing cases.155 Nevertheless it is a
strange fact that within the same district, during the height of the breeding-
season, there should be so many males and females always ready to repair
the loss of a mated bird. Why do not such spare birds immediately pair

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together? Have we not some reason to suspect, and the suspicion has
occurred to Mr. Jenner Weir, that inasmuch as the act of courtship appears
to be with many birds a prolonged and tedious affair, so it occasionally
happens that certain males and females do not succeed during the proper
season, in exciting each other’s love, and consequently do not pair? This
suspicion will appear somewhat less improbable after we have seen what
strong antipathies and preferences female birds occasionally evince towards
particular males.
Mental Qualities of Birds, and their taste for the beautiful.—Before we
discuss any further the question whether the females select the more
attractive males or accept the first whom they may encounter, it will be
advisable briefly to consider the mental powers of birds. Their reason is
generally, and perhaps justly, ranked as low; yet some facts could be
given156 leading to an opposite conclusion. Low powers of reasoning,
however, are compatible, as we see with mankind, with strong affections,
acute perception, and a taste for the beautiful; and it is with these latter
qualities that we are here concerned. It has often been said that parrots
become so deeply attached to each other that when one dies the other for a
long time pines; but Mr. Jenner Weir thinks that with most birds the strength
of their affection has been much exaggerated. Nevertheless when one of a
pair in a state of nature has been shot, the survivor has been heard for days
afterwards uttering a plaintive call; and Mr. St. John gives157 various facts
proving the attachment of mated birds. Starlings, however, as we have seen,
may be consoled thrice in the same day for the loss of their mates. In the
Zoological Gardens parrots have clearly recognised their former masters
after an interval of some months. Pigeons have such excellent local
memories that they have been known to return to their former homes after
an interval of nine months, yet, as I hear from Mr. Harrison Weir, if a pair
which would naturally remain mated for life be separated for a few weeks
during the winter and matched with other birds, the two, when brought
together again, rarely, if ever, recognise each other.
Birds sometimes exhibit benevolent feelings; they will feed the deserted
young even of distinct species, but this perhaps ought to be considered as a
mistaken instinct. They will also feed, as shewn in an earlier part of this
work, adult birds of their own species which have become blind. Mr.
Buxton gives a curious account of a parrot which took care of a frost-bitten

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and crippled bird of a distinct species, cleansed her feathers and defended
her from the attacks of the other parrots which roamed freely about his
garden. It is a still more curious fact that these birds apparently evince some
sympathy for the pleasures of their fellows. When a pair of cockatoos made
a nest in an acacia tree, “it was ridiculous to see the extravagant interest
taken in the matter by the others of the same species.” These parrots, also,
evinced unbounded curiosity, and clearly had “the idea of property and
possession.”158
Birds possess acute powers of observation. Every mated bird, of course,
recognises its fellow. Audubon states that with the mocking-thrushes of the
United States (Mimus polyglottus) a certain number remain all the year
round in Louisiana, whilst the others migrate to the Eastern States; these
latter, on their return, are instantly recognised, and always attacked, by their
Southern brethren. Birds under confinement distinguish different persons,
as is proved by the strong and permanent antipathy or affection which they
shew, without any apparent cause, towards certain individuals. I have heard
of numerous instances with jays, partridges, canaries, and especially
bullfinches. Mr. Hussey has described in how extraordinary a manner a
tamed partridge recognised everybody; and its likes and dislikes were very
strong. This bird seemed “fond of gay colours, and no new gown or cap
could be put on without catching his attention.”159 Mr. Hewitt has carefully
described the habits of some ducks (recently descended from wild birds),
which, at the approach of a strange dog or cat, would rush headlong into the
water, and exhaust themselves in their attempts to escape; but they knew so
well Mr. Hewitt’s own dogs and cats that they would lie down and bask in
the sun close to them. They always moved away from a strange man, and so
they would from the lady who attended them, if she made any great change
in her dress. Audubon relates that he reared and tamed a wild turkey which
always ran away from any strange dog; this bird escaped into the woods,
and some days afterwards Audubon saw, as he thought, a wild turkey, and
made his dog chase it; but to his astonishment, the bird did not run away,
and the dog, when he came up, did not attack the bird, for they mutually
recognised each other as old friends.160
Mr. Jenner Weir is convinced that birds pay particular attention to the
colours of other birds, sometimes out of jealousy, and sometimes as a sign
of kinship. Thus he turned a reed-bunting (Emberiza schœniculus), which

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had acquired its black head, into his aviary, and the new-comer was not
noticed by any bird, except by a bullfinch, which is likewise black-headed.
This bullfinch was a very quiet bird, and had never before quarrelled with
any of its comrades, including another reed-bunting, which had not as yet
become black-headed: but the reed-bunting with a black head was so
unmercifully treated, that it had to be removed. Mr. Weir was also obliged
to turn out a robin, as it fiercely attacked all birds with any red in their
plumage, but no other kinds; it actually killed a red-breasted crossbill, and
nearly killed a goldfinch. On the other hand, he has observed that some
birds, when first introduced into his aviary, fly towards the species which
resemble them most in colour, and settle by their sides.
As male birds display with so much care their fine plumage and other
ornaments in the presence of the females, it is obviously probable that these
appreciate the beauty of their suitors. It is, however, difficult to obtain direct
evidence of their capacity to appreciate beauty. When birds gaze at
themselves in a looking-glass (of which many instances have been
recorded) we cannot feel sure that it is not from jealousy at a supposed
rival, though this is not the conclusion of some observers. In other cases it
is difficult to distinguish between mere curiosity and admiration. It is
perhaps the former feeling which, as stated by Lord Lilford,161 attracts the
Ruff strongly towards any bright object, so that, in the Ionian Islands, it
“will dart down to a bright-coloured handkerchief, regardless of repeated
shots.” The common lark is drawn down from the sky, and is caught in large
numbers, by a small mirror made to move and glitter in the sun. Is it
admiration or curiosity which leads the magpie, raven, and some other birds
to steal and secrete bright objects, such as silver articles or jewels?
Mr. Gould states that certain humming-birds decorate the outside of their
nests, “with the utmost taste; they instinctively fasten thereon beautiful
pieces of flat lichen, the larger pieces in the middle, and the smaller on the
part attached to the branch. Now and then a pretty feather is intertwined or
fastened to the outer sides, the stem being always so placed, that the feather
stands out beyond the surface.” The best evidence, however, of a taste for
the beautiful is afforded by the three genera of Australian bower-birds
already mentioned. Their bowers (see fig. 46, p. 70), where the sexes
congregate and play strange antics, are differently constructed, but what
most concerns us is, that they are decorated in a different manner by the

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several species. The Satin bower-bird collects gaily-coloured articles, such
as the blue tail-feathers of parrakeets, bleached bones and shells, which it
sticks between the twigs, or arranges at the entrance. Mr. Gould found in
one bower a neatly-worked stone tomahawk and a slip of blue cotton,
evidently procured from a native encampment. These objects are
continually rearranged, and carried about by the birds whilst at play. The
bower of the Spotted bower-bird “is beautifully lined with tall grasses, so
disposed that the heads nearly meet, and the decorations are very profuse.”
Round stones are used to keep the grass-stems in their proper places, and to
make divergent paths leading to the bower. The stones and shells are often
brought from a great distance. The Regent bird, as described by Mr.
Ramsay, ornaments its short bower with bleached land-shells belonging to
five or six species, and with “berries of various colours, blue, red, and
black, which give it when fresh a very pretty appearance. Besides these
there were several newly-picked leaves and young shoots of a pinkish
colour, the whole shewing a decided taste for the beautiful.” Well may Mr.
Gould say “these highly decorated halls of assembly must be regarded as
the most wonderful instances of bird-architecture yet discovered;” and the
taste, as we see, of the several species certainly differs.162

Preference for particular Males by the Females.—Having made these
preliminary remarks on the discrimination and taste of birds, I will give all
the facts known to me, which bear on the preference shewn by the female
for particular males. It is certain that distinct species of birds occasionally
pair in a state of nature and produce hybrids. Many instances could be
given: thus Macgillivray relates how a male blackbird and female thrush
“fell in love with each other,” and produced offspring.163 Several years ago
eighteen cases had been recorded of the occurrence in Great Britain of
hybrids between the black grouse and pheasant;164 but most of these cases
may perhaps be accounted for by solitary birds not finding one of their own
species to pair with. With other birds, as Mr. Jenner Weir has reason to
believe, hybrids are sometimes the result of the casual intercourse of birds
building in close proximity. But these remarks do not apply to the many
recorded instances of tamed or domestic birds, belonging to distinct species,
which have become absolutely fascinated with each other, although living
with their own species. Thus Waterton165 states that out of a flock of twenty-
three Canada geese, a female paired with a solitary Bernicle gander,

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although so different in appearance and size; and they produced hybrid
offspring. A male Wigeon (Mareca penelope), living with females of the
same species, has been known to pair with a Pintail duck, Querquedula
acuta. Lloyd describes the remarkable attachment between a shield-drake
(Tadorna vulpanser) and a common duck. Many additional instances could
be given; and the Rev. E. S. Dixon remarks that “Those who have kept
many different species of geese together, well know what unaccountable
attachments they are frequently forming, and that they are quite as likely to
pair and rear young with individuals of a race (species) apparently the most
alien to themselves, as with their own stock.”
The Rev. W. D. Fox informs me that he possessed at the same time a pair of
Chinese geese (Anser cygnoides), and a common gander with three geese.
The two lots kept quite separate, until the Chinese gander seduced one of
the common geese to live with him. Moreover, of the young birds hatched
from the eggs of the common geese, only four were pure, the other eighteen
proving hybrids; so that the Chinese gander seems to have had prepotent
charms over the common gander. I will give only one other case; Mr.
Hewitt states that a wild duck, reared in captivity, “after breeding a couple
of seasons with her own mallard, at once shook him off on my placing a
male Pintail on the water. It was evidently a case of love at first sight, for
she swam about the new-comer caressingly, though he appeared evidently
alarmed and averse to her overtures of affection. From that hour she forgot
her old partner. Winter passed by, and the next spring the Pintail seemed to
have become a convert to her blandishments, for they nested and produced
seven or eight young ones.”
What the charm may have been in these several cases, beyond mere novelty,
we cannot even conjecture. Colour, however, sometimes comes into play;
for in order to raise hybrids from the siskin (Fringilla spinus) and the
canary, it is much the best plan, according to Bechstein, to place birds of the
same tint together. Mr. Jenner Weir turned a female canary into his aviary,
where there were male linnets, goldfinches, siskins, greenfinches,
chaffinches, and other birds, in order to see which she would choose; but
there never was any doubt, and the greenfinch carried the day. They paired
and produced hybrid offspring.

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With the members of the same species the fact of the female preferring to
pair with one male rather than with another is not so likely to excite
attention, as when this occurs between distinct species. Such cases can best
be observed with domesticated or confined birds; but these are often
pampered by high feeding, and sometimes have their instincts vitiated to an
extreme degree. Of this latter fact I could give sufficient proofs with
pigeons, and especially with fowls, but they cannot be here related. Vitiated
instincts may also account for some of the hybrid unions above referred to;
but in many of these cases the birds were allowed to range freely over large
ponds, and there is no reason to suppose that they were unnaturally
stimulated by high feeding.
With respect to birds in a state of nature, the first and most obvious
supposition which will occur to everyone is that the female at the proper
season accepts the first male whom she may encounter; but she has at least
the opportunity for exerting a choice, as she is almost invariably pursued by
many males. Audubon—and we must remember that he spent a long life in
prowling about the forests of the United States and observing the birds—
does not doubt that the female deliberately chooses her mate; thus, speaking
of a woodpecker, he says the hen is followed by half-a-dozen gay suitors,
who continue performing strange antics, “until a marked preference is
shewn for one.” The female of the red-winged starling (Agelæus phœniceus)
is likewise pursued by several males, “until, becoming fatigued, she alights,
receives their addresses, and soon makes a choice.” He describes also how
several male nightjars repeatedly plunge through the air with astonishing
rapidity, suddenly turning, and thus making a singular noise; “but no sooner
has the female made her choice, than the other males are driven away.”
With one of the vultures (Cathartes aura) of the United States, parties of
eight or ten or more males and females assemble on fallen logs, “exhibiting
the strongest desire to please mutually,” and after many caresses, each male
leads off his partner on the wing. Audubon likewise carefully observed the
wild flocks of Canada geese (Anser Canadensis), and gives a graphic
description of their love-antics; he says that the birds which had been
previously mated “renewed their courtship as early as the month of January,
while the others would be contending or coquetting for hours every day,
until all seemed satisfied with the choice they had made, after which,
although they remained together, any person could easily perceive that they

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were careful to keep in pairs. I have observed also that the older the birds,
the shorter were the preliminaries of their courtship. The bachelors and old
maids, whether in regret, or not caring to be disturbed by the bustle, quietly
moved aside and lay down at some distance from the rest.”166 Many similar
statements with respect to other birds could be cited from this same
observer.
Turning now to domesticated and confined birds, I will commence by
giving what little I have learnt respecting the courtship of fowls. I have
received long letters on this subject from Messrs. Hewitt and Tegetmeier,
and almost an essay from the late Mr. Brent. It will be admitted by every
one that these gentlemen, so well known from their published works, are
careful and experienced observers. They do not believe that the females
prefer certain males on account of the beauty of their plumage; but some
allowance must be made for the artificial state under which they have long
been kept. Mr. Tegetmeier is convinced that a game-cock, though disfigured
by being dubbed with his hackles trimmed, would be accepted as readily as
a male retaining all his natural ornaments. Mr. Brent, however, admits that
the beauty of the male probably aids in exciting the female; and her
acquiescence is necessary. Mr. Hewitt is convinced that the union is by no
means left to mere chance, for the female almost invariably prefers the most
vigorous, defiant, and mettlesome male; hence it is almost useless, as he
remarks, “to attempt true breeding if a game-cock in good health and
condition runs the locality, for almost every hen on leaving the roosting-
place will resort to the game-cock, even though that bird may not actually
drive away the male of her own variety.” Under ordinary circumstances the
males and females of the fowl seem to come to a mutual understanding by
means of certain gestures, described to me by Mr. Brent. But hens will often
avoid the officious attentions of young males. Old hens, and hens of a
pugnacious disposition, as the same writer informs me, dislike strange
males, and will not yield until well beaten into compliance. Ferguson,
however, describes how a quarrelsome hen was subdued by the gentle
courtship of a Shanghai cock.167
There is reason to believe that pigeons of both sexes prefer pairing with
birds of the same breed; and dovecot-pigeons dislike all the highly
improved breeds.168 Mr. Harrison Weir has lately heard from a trustworthy
observer, who keeps blue pigeons, that these drive away all other coloured

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varieties, such as white, red, and yellow; and from another observer, that a
female dun carrier could not be matched, after repeated trials, with a black
male, but immediately paired with a dun. Generally colour alone appears to
have little influence on the pairing of pigeons. Mr. Tegetmeier, at my
request, stained some of his birds with magenta, but they were not much
noticed by the others.
Female pigeons occasionally feel a strong antipathy towards certain males,
without any assignable cause. Thus MM. Boitard and Corbié, whose
experience extended over forty-five years, state: “Quand une femelle
éprouve de l’antipathie pour un mâle avec lequel on veut l’accoupler,
malgré tous les feux de l’amour, malgré l’alpiste et le chènevis dont on la
nourrit pour augmenter son ardeur, malgré un emprisonnement de six mois
et même d’un an, elle refuse constamment ses caresses; les avances
empressées, les agaceries, les tournoiemens, les tendres roucoulemens, rien
ne peut lui plaire ni l’émouvoir; gonflée, boudeuse, blottie dans un coin de
sa prison, elle n’en sort que pour boire et manger, ou pour repousser avec
une espèce de rage des caresses devenues trop pressantes.”169 On the other
hand, Mr. Harrison Weir has himself observed, and has heard from, several
breeders, that a female pigeon will occasionally take a strong fancy for a
particular male, and will desert her own mate for him. Some females,
according to another experienced observer, Riedel,170 are of a profligate
disposition, and prefer almost any stranger to their own mate. Some
amorous males, called by our English fanciers “gay birds,” are so successful
in their gallantries, that, as Mr. H. Weir informs me, they must be shut up,
on account of the mischief which they cause.
Wild turkeys in the United States, according to Audubon, “sometimes pay
their addresses to the domesticated females, and are generally received by
them with great pleasure.” So that these females apparently prefer the wild
to their own males.171
Here is a more curious case. Sir R. Heron during many years kept an
account of the habits of the peafowl, which he bred in large numbers. He
states that “the hens have frequently great preference to a particular
peacock. They were all so fond of an old pied cock, that one year, when he
was confined though still in view, they were constantly assembled close to
the trellice-walls of his prison, and would not suffer a japanned peacock to

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touch them. On his being let out in the autumn, the oldest of the hens
instantly courted him, and was successful in her courtship. The next year he
was shut up in a stable, and then the hens all courted his rival.”172 This rival
was a japanned or black-winged peacock, which to our eyes is a more
beautiful bird than the common kind.
Lichtenstein, who was a good observer and had excellent opportunities of
observation at the Cape of Good Hope, assured Rudolphi that the female
widow-bird (Chera progne) disowns the male, when robbed of the long tail-
feathers with which he is ornamented during the breeding-season. I presume
that this observation must have been made on birds under confinement.173
Here is another striking case; Dr. Jaeger,174 director of the Zoological
Gardens of Vienna, states that a male silver pheasant, who had been
triumphant over the other males and was the accepted lover of the females,
had his ornamental plumage spoiled. He was then immediately superseded
by a rival, who got the upper hand and afterwards led the flock.
Not only does the female exert a choice, but in some few cases she courts
the male, or even fights for his possession. Sir R. Heron states that with
peafowl, the first advances are always made by the female; something of
the same kind takes place, according to Audubon, with the older females of
the wild turkey. With the capercailzie, the females flit round the male,
whilst he is parading at one of the places of assemblage, and solicit his
attention.175 We have seen that a tame wild-duck seduced after a long
courtship an unwilling Pintail drake. Mr. Bartlett believes that the
Lophophorus, like many other gallinaceous birds, is naturally polygamous,
but two females cannot be placed in the same cage with a male, as they
fight so much together. The following instance of rivalry is more surprising
as it relates to bullfinches, which usually pair for life. Mr. Jenner Weir
introduced a dull-coloured and ugly female into his aviary, and she
immediately attacked another mated female so unmercifully that the latter
had to be separated. The new female did all the courtship, and was at last
successful, for she paired with the male; but after a time she met with a just
retribution, for, ceasing to be pugnacious, Mr. Weir replaced the old female,
and the male then deserted his new and returned to his old love.
In all ordinary cases the male is so eager that he will accept any female, and
does not, as far as we can judge, prefer one to the other; but exceptions to

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this rule, as we shall hereafter see, apparently occur in some few groups.
With domesticated birds, I have heard of only one case in which the males
shew any preference for particular females, namely, that of the domestic
cock, who, according to the high authority of Mr. Hewitt, prefers the
younger to the older hens. On the other hand, in effecting hybrid unions
between the male pheasant and common hens, Mr. Hewitt is convinced that
the pheasant invariably prefers the older birds. He does not appear to be in
the least influenced by their colour, but “is most capricious in his
attachments.”176 From some inexplicable cause he shews the most
determined aversion to certain hens, which no care on the part of the
breeder can overcome. Some hens, as Mr. Hewitt informs me, are quite
unattractive even to the males of their own species, so that they may be kept
with several cocks during a whole season, and not one egg out of forty or
fifty will prove fertile. On the other hand with the Long-tailed duck
(Harelda glacialis), “it has been remarked,” says M. Ekström, “that certain
females are much more courted than the rest. Frequently, indeed, one sees
an individual surrounded by six or eight amorous males.” Whether this
statement is credible, I know not; but the native sportsmen shoot these
females in order to stuff them as decoys.177
With respect to female birds feeling a preference for particular males, we
must bear in mind that we can judge of choice being exerted, only by
placing ourselves in imagination in the same position. If an inhabitant of
another planet were to behold a number of young rustics at a fair, courting
and quarrelling over a pretty girl, like birds at one of their places of
assemblage, he would be able to infer that she had the power of choice only
by observing the eagerness of the wooers to please her, and to display their
finery. Now with birds, the evidence stands thus; they have acute powers of
observation, and they seem to have some taste for the beautiful both in
colour and sound. It is certain that the females occasionally exhibit, from
unknown causes, the strongest antipathies and preferences for particular
males. When the sexes differ in colour or in other ornaments, the males
with rare exceptions are the most highly decorated, either permanently or
temporarily during the breeding-season. They sedulously display their
various ornaments, exert their voices, and perform strange antics in the
presence of the females. Even well-armed males, who, it might have been
thought, would have altogether depended for success on the law of battle,

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are in most cases highly ornamented; and their ornaments have been
acquired at the expense of some loss of power. In other cases ornaments
have been acquired, at the cost of increased risk from birds and beasts of
prey. With various species many individuals of both sexes congregate at the
same spot, and their courtship is a prolonged affair. There is even reason to
suspect that the males and females within the same district do not always
succeed in pleasing each other and pairing.
What then are we to conclude from these facts and considerations? Does the
male parade his charms with so much pomp and rivalry for no purpose? Are
we not justified in believing that the female exerts a choice, and that she
receives the addresses of the male who pleases her most? It is not probable
that she consciously deliberates; but she is most excited or attracted by the
most beautiful, or melodious, or gallant males. Nor need it be supposed that
the female studies each stripe or spot of colour; that the peahen, for
instance, admires each detail in the gorgeous train of the peacock—she is
probably struck only by the general effect. Nevertheless after hearing how
carefully the male Argus pheasant displays his elegant primary wing-
feathers, and erects his ocellated plumes in the right position for their full
effect; or again, how the male goldfinch alternately displays his gold-
bespangled wings, we ought not to feel too sure that the female does not
attend to each detail of beauty. We can judge, as already remarked, of
choice being exerted, only from the analogy of our own minds; and the
mental powers of birds, if reason be excluded, do not fundamentally differ
from ours. From these various considerations we may conclude that the
pairing of birds is not left to chance; but that those males, which are best
able by their various charms to please or excite the female, are under
ordinary circumstances accepted. If this be admitted, there is not much
difficulty in understanding how male birds have gradually acquired their
ornamental characters. All animals present individual differences, and as
man can modify his domesticated birds by selecting the individuals which
appear to him the most beautiful, so the habitual or even occasional
preference by the female of the more attractive males would almost
certainly lead to their modification; and such modifications might in the
course of time be augmented to almost any extent, compatible with the
existence of the species.

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Variability of Birds, and especially of their secondary Sexual Characters.—
Variability and inheritance are the foundations for the work of selection.
That domesticated birds have varied greatly, their variations being inherited,
is certain. That birds in a state of nature present individual differences is
admitted by every one; and that they have sometimes been modified into
distinct races, is generally admitted.178 Variations are of two kinds, which
insensibly graduate into each other, namely, slight differences between all
the members of the same species, and more strongly-marked deviations
which occur only occasionally. These latter are rare with birds in a state of
nature, and it is very doubtful whether they have often been preserved
through selection, and then transmitted to succeeding generations.179
Nevertheless, it may be worth while to give the few cases relating chiefly to
colour (simple albinism and melanism being excluded), which I have been
able to collect.
Mr. Gould is well known rarely to admit the existence of varieties, for he
esteems very slight differences as specific; now he states180 that near Bogota
certain humming-birds belonging to the genus Cynanthus are divided into
two or three races or varieties, which differ from each other in the colouring
of the tail,—“some having the whole of the feathers blue, while others have
the eight central ones tipped with beautiful green.” It does not appear that
intermediate gradations have been observed in this or the following cases.
In the males alone of one of the Australian parrakeets “the thighs in some
are scarlet, in others grass-green.” In another parrakeet of the same country
“some individuals have the band across the wing-coverts bright-yellow,
while in others the same part is tinged with red.”181 In the United States
some few of the males of the Scarlet Tanager (Tanagra rubra) have “a
beautiful transverse band of glowing red on the smaller wing-coverts;”182
but this variation seems to be somewhat rare, so that its preservation
through sexual selection would follow only under unusually favourable
circumstances. In Bengal the Honey buzzard (Pernis cristata) has either a
small rudimental crest on its head, or none at all; so slight a difference
however would not have been worth notice, had not this same species
possessed in Southern India “a well-marked occipital crest formed of
several graduated feathers.”183
The following case is in some respects more interesting. A pied variety of
the raven, with the head, breast, abdomen, and parts of the wings and tail-

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feathers white, is confined to the Feroe Islands. It is not very rare there, for
Graba saw during his visit from eight to ten living specimens. Although the
characters of this variety are not quite constant, yet it has been named by
several distinguished ornithologists as a distinct species. The fact of the
pied birds being pursued and persecuted with much clamour by the other
ravens of the island was the chief cause which led Brünnich to conclude
that it was specifically distinct; but this is now known to be an error.184
In various parts of the northern seas a remarkable variety of the common
Guillemot (Uria troile) is found; and in Feroe, one out of every five birds,
according to Graba’s estimation, consists of this variety. It is
characterised185 by a pure white ring round the eye, with a curved narrow
white line, an inch and a half in length, extending back from the ring. This
conspicuous character has caused the bird to be ranked by several
ornithologists as a distinct species under the name of U. lacrymans, but it is
now known to be merely a variety. It often pairs with the common kind, yet
intermediate gradations have never been seen; nor is this surprising, for
variations which appear suddenly are often, as I have elsewhere shewn,186
transmitted either unaltered or not at all. We thus see that two distinct forms
of the same species may co-exist in the same district, and we cannot doubt
that if the one had possessed any great advantage over the other, it would
soon have been multiplied to the exclusion of the latter. If, for instance, the
male pied ravens, instead of being persecuted and driven away by their
comrades, had been highly attractive, like the pied peacock before
mentioned, to the common black females, their numbers would have rapidly
increased. And this would have been a case of sexual selection.
With respect to the slight individual differences which are common, in a
greater or less degree, to all the members of the same species, we have
every reason to believe that they are by far the most important for the work
of selection. Secondary sexual characters are eminently liable to vary, both
with animals in a state of nature and under domestication.187 There is also
reason to believe, as we have seen in our eighth chapter, that variations are
more apt to occur in the male than in the female sex. All these contingencies
are highly favourable for sexual selection. Whether characters thus acquired
are transmitted to one sex or to both sexes, depends exclusively in most
cases, as I hope to shew in the following chapter, on the form of inheritance
which prevails in the groups in question.

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It is sometimes difficult to form any opinion whether certain slight
differences between the sexes of birds are simply the result of variability
with sexually-limited inheritance, without the aid of sexual selection, or
whether they have been augmented through this latter process. I do not here
refer to the innumerable instances in which the male displays splendid
colours or other ornaments, of which the female partakes only to a slight
degree; for these cases are almost certainly due to characters primarily
acquired by the male, having been transferred, in a greater or less degree, to
the female. But what are we to conclude with respect to certain birds in
which, for instance, the eyes differ slightly in colour in the two sexes?188 In
some cases the eyes differ conspicuously; thus with the storks of the genus
Xenorhynchus those of the male are blackish-hazel, whilst those of the
females are gamboge-yellow; with many hornbills (Buceros), as I hear from
Mr. Blyth,189 the males have intense crimson, and the females white eyes. In
the Buceros bicornis, the hind margin of the casque and a stripe on the crest
of the beak are black in the male, but not so in the female. Are we to
suppose that these black marks and the crimson colour of the eyes have
been preserved or augmented through sexual selection in the males? This is
very doubtful; for Mr. Bartlett shewed me in the Zoological Gardens that
the inside of the mouth of this Buceros is black in the male and flesh-
coloured in the female; and their external appearance or beauty would not
be thus affected. I observed in Chili190 that the iris in the condor, when about
a year old, is dark-brown, but changes at maturity into yellowish-brown in
the male, and into bright red in the female. The male has also a small,
longitudinal, leaden-coloured, fleshy crest or comb. With many
gallinaceous birds the comb is highly ornamental, and assumes vivid
colours during the act of courtship; but what are we to think of the dull-
coloured comb of the condor, which does not appear to us in the least
ornamental? The same question may be asked in regard to various other
characters, such as the knob on the base of the beak of the Chinese goose
(Anser cygnoides), which is much larger in the male than in the female. No
certain answer can be given to these questions; but we ought to be cautious
in assuming that knobs and various fleshy appendages cannot be attractive
to the female, when we remember that with savage races of man various
hideous deformities=-deep scars on the face with the flesh raised into
protuberances, the septum of the nose pierced by sticks or bones, holes in
the ears and lips stretched widely open—are all admired as ornamental.

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Whether or not unimportant differences between the sexes, such as those
just specified, have been preserved through sexual selection, these
differences, as well as all others, must primarily depend on the laws of
variation. On the principle of correlated development, the plumage often
varies on different parts of the body, or over the whole body, in the same
manner. We see this well illustrated in certain breeds of the fowl. In all the
breeds the feathers on the neck and loins of the males are elongated, and are
called hackles; now when both sexes acquire a top-knot, which is a new
character in the genus, the feathers on the head of the male become hackle-
shaped, evidently on the principle of correlation; whilst those on the head of
the female are of the ordinary shape. The colour also of the hackles forming
the top-knot of the male, is often correlated with that of the hackles on the
neck and loins, as may be seen by comparing these feathers in the Golden
and Silver-spangled Polish, the Houdans, and Crève-cœur breeds. In some
natural species we may observe exactly the same correlation in the colours
of these same feathers, as in the males of the splendid Golden and Amherst
pheasants.
The structure of each individual feather generally causes any change in its
colouring to be symmetrical; we see this in the various laced, spangled, and
pencilled breeds of the fowl; and on the principle of correlation the feathers
over the whole body are often modified in the same manner. We are thus
enabled without much trouble to rear breeds with their plumage marked and
coloured almost as symmetrically as in natural species. In laced and
spangled fowls the coloured margins of the feathers are abruptly defined;
but in a mongrel raised by me from a black Spanish cock glossed with
green and a white game hen, all the feathers were greenish-black, excepting
towards their extremities, which were yellowish-white; but between the
white extremities and the black bases, there was on each feather a
symmetrical, curved zone of dark-brown. In some instances the shaft of the
feather determines the distribution of the tints; thus with the body-feathers
of a mongrel from the same black Spanish cock and a silver-spangled Polish
hen, the shaft, together with a narrow space on each side, was greenish-
black, and this was surrounded by a regular zone of dark-brown, edged with
brownish-white. In these cases we see feathers becoming symmetrically
shaded, like those which give so much elegance to the plumage of many
natural species. I have also noticed a variety of the common pigeon with the

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wing-bars symmetrically zoned with three bright shades, instead of being
simply black on a slaty-blue ground, as in the parent-species.
In many large groups of birds it may be observed that the plumage is
differently coloured in each species, yet that certain spots, marks, or stripes,
though likewise differently coloured, are retained by all the species.
Analogous cases occur with the breeds of the pigeon, which usually retain
the two wing-bars, though they may be coloured red, yellow, white, black,
or blue, the rest of the plumage being of some wholly different tint. Here is
a more curious case, in which certain marks are retained, though coloured
in almost an exactly reversed manner to what is natural; the aboriginal
pigeon has a blue tail, with the terminal halves of the outer webs of the two
outer tail-feathers white; now there is a sub-variety having a white instead
of a blue tail, with precisely that small part black which is white in the
parent-species.191
Formation and variability of the Ocelli or eye-like Spots on the Plumage of
Birds.—As no ornaments are more beautiful than the ocelli on the feathers
of various birds, on the hairy coats of some mammals, on the scales of
reptiles and fishes, on the skin of amphibians, on the wings of many
Lepidoptera and other insects, they deserve to be especially noticed. An
ocellus consists of a spot within a ring of another colour, like the pupil
within the iris, but the central spot is often surrounded by additional
concentric zones. The ocelli on the tail-coverts of the peacock offer a
familiar example, as well as those on the wings of the peacock-butterfly
(Vanessa). Mr. Trimen has given me a description of a S. African moth
(Gynanisa Isis), allied to our Emperor moth, in which a magnificent ocellus
occupies nearly the whole surface of each hinder wing; it consists of a black
centre, including a semi-transparent crescent-shaped mark, surrounded by
successive ochre-yellow, black, ochre-yellow, pink, white, pink, brown, and
whitish zones. Although we do not know the steps by which these
wonderfully beautiful and complex ornaments have been developed, the
process at least with insects has probably been a simple one; for, as Mr.
Trimen writes to me, “no characters of mere marking or coloration are so
unstable in the Lepidoptera as the ocelli, both in number and size.” Mr.
Wallace, who first called my attention to this subject, shewed me a series of
specimens of our common meadow-brown butterfly (Hipparchia Janira)
exhibiting numerous gradations from a simple minute black spot to an

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elegantly-shaded ocellus. In a S. African butterfly (Cyllo Leda belonging to
the same family, the ocelli are even still more variable. In some specimens
(A, fig. 52) large spaces on the upper surface of the wings are coloured
black, and include irregular white marks; and from this state a complete
gradation can be traced into a tolerably perfect (A1) ocellus, and this results
from the contraction of the irregular blotches of colour. In another series of
specimens a gradation can be followed from excessively minute white dots,
surrounded by a scarcely visible black line (B), into perfectly symmetrical
and large ocelli (B1).192 In cases like these, the development of a perfect
ocellus does not require a long course of variation and selection.

Fig. 52. Cyllo leda, Linn., from a drawing by Mr. Trimen, shewing the extreme range of
variation in the ocelli.
A. Specimen, from Mauritius, upper surface of B. Specimen, from Java, upper surface of hind-
fore-wing wing.
1 1
A . Specimen, from Natal, ditto. B . Specimen, from Mauritius, ditto.

With birds and many other animals it seems, from the comparison of allied
species, to follow, that circular spots are often generated by the breaking up
and contraction of stripes. In the Tragopan pheasant faint white lines in the
female represent the beautiful white spots in the male;193 and something of
the same kind may be observed in the two sexes of the Argus pheasant.
However this may be, appearances strongly favour the belief that, on the

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one hand, a dark spot is often formed by the colouring-matter being drawn
towards a central point from a surrounding zone, which is thus rendered
lighter. And, on the other hand, that a white spot is often formed by the
colour being driven away from a central point, so that it accumulates in a
surrounding darker zone. In either case an ocellus is the result. The
colouring matter seems to be a nearly constant quantity, but is redistributed,
either centripetally or centrifugally. The feathers of the common guinea-
fowl offer a good instance of white spots surrounded by darker zones; and
wherever the white spots are large and stand near each other, the
surrounding dark zones become confluent. In the same wing-feather of the
Argus pheasant dark spots may be seen surrounded by a pale zone, and
white spots by a dark zone. Thus the formation of an ocellus in its simplest
state appears to be a simple affair. By what further steps the more complex
ocelli, which are surrounded by many successive zones of colour, have been
generated, I will not pretend to say. But bearing in mind the zoned feathers
of the mongrel offspring from differently-coloured fowls, and the
extraordinary variability of the ocelli in many Lepidoptera, the formation of
these beautiful ornaments can hardly be a highly complex process, and
probably depends on some slight and graduated change in the nature of the
tissues.

Gradation of Secondary Sexual Characters.—Cases of gradation are
important for us, as they shew that it is at least possible that highly complex
ornaments may have been acquired by small successive steps. In order to
discover the actual steps by which the male of any existing bird has
acquired his magnificent colours or other ornaments, we ought to behold
the long line of his ancient and extinct progenitors; but this is obviously
impossible. We may, however, generally gain a clue by comparing all the
species of a group, if it be a large one; for some of them will probably
retain, at least in a partial manner, traces of their former characters. Instead
of entering on tedious details respecting various groups, in which striking
instances of gradation could be given, it seems the best plan to take some
one or two strongly-characterised cases, for instance that of the peacock, in
order to discover if any light can thus be thrown on the steps by which this
bird has become so splendidly decorated. The peacock is chiefly remarkable
from the extraordinary length of his tail-coverts; the tail itself not being
much elongated. The barbs along nearly the whole length of these feathers

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stand separate or are decomposed; but this is the case with the feathers of
many species, and with some varieties of the domestic fowl and pigeon. The
barbs coalesce towards the extremity of the shaft to form the oval disc or
ocellus, which is certainly one of the most beautiful objects in the world.
This consists of an iridescent, intensely blue, indented centre, surrounded
by a rich green zone, and this by a broad coppery-brown zone, and this by
five other narrow zones of slightly-different iridescent shades. A trifling
character in the disc perhaps deserves notice; the barbs, for a space along
one of the concentric zones are destitute, to a greater or less degree, of their
barbules, so that a part of the disc is surrounded by an almost transparent
zone, which gives to it a highly-finished aspect. But I have elsewhere
described194 an exactly analogous variation in the hackles of a sub-variety of
the game-cock, in which the tips, having a metallic lustre, “are separated
from the lower part of the feather by a symmetrically-shaped transparent
zone, composed of the naked portions of the barbs.” The lower margin or
base of the dark-blue centre of the ocellus is deeply indented on the line of
the shaft. The surrounding zones likewise shew traces, as may be seen in
the drawing (fig. 53), of indentations, or rather breaks. These indentations
are common to the Indian and Javan peacocks (Pavo cristatus and P.
muticus); and they seemed to me to deserve particular attention, as probably
connected with the development of the ocellus; but for a long time I could
not conjecture their meaning.
If we admit the principle of gradual evolution, there must formerly have
existed many species which presented every successive step between the
wonderfully elongated tail-coverts of the peacock and the short tail-coverts
of all ordinary birds; and again between the magnificent ocelli of the
former, and the simpler ocelli or mere coloured spots of other birds; and so
with all the other characters of the peacock. Let us look to the allied
Gallinaceæ for any still-existing gradations. The species and sub-species of
Polyplectron inhabit countries adjacent to the native land of the peacock;
and they so far resemble this bird that they are sometimes called peacock-
pheasants. I am also informed by Mr. Bartlett that they resemble the
peacock in their voice and in some of their habits. During the spring the
males, as previously described, strut about before the comparatively plain-
coloured females, expanding and erecting their tail and wing-feathers,
which are ornamented with numerous ocelli. I request the reader to turn

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back to the drawing (fig. 51, p. 90) of a Polyplectron. In P. Napoleonis the
ocelli are confined to the tail, and the back is of a rich metallic blue, in
which respects this species approaches the Java peacock. P. Hardwickii
possesses a peculiar top-knot, somewhat like that of this same kind of
peacock. The ocelli on the wings and tail of the several species of
Polyplectron are either circular or oval, and consist of a beautiful,
iridescent, greenish-blue or greenish-purple disc, with a black border. This
border in P. chinquis shades into brown which is edged with cream-colour,
so that the ocellus is here surrounded with differently, though not brightly,
shaded concentric zones. The unusual length of the tail-coverts is another
highly remarkable character in Polyplectron; for in some of the species they
are half as long, and in others two-thirds of the length of the true tail-
feathers. The tail-coverts are ocellated, as in the peacock. Thus the several
species of Polyplectron manifestly make a graduated approach in the length
of their tail-coverts, in the zoning of the ocelli, and in some other
characters, to the peacock.

Fig. 53. Feather of Peacock, about two-thirds of natural size, carefully drawn by Mr. Ford. The
transparent zone is represented by the outermost white zone, confined to the upper end of the
disc.

Notwithstanding this approach, the first species of Polyplectron which I
happened to examine almost made me give up the search; for I found not
only that the true tail-feathers, which in the peacock are quite plain, were
ornamented with ocelli, but that the ocelli on all the feathers differed

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fundamentally from those of the peacock, in there being two on the same
feather, (fig. 54), one on each side of the shaft. Hence I concluded that the
early progenitors of the peacock could not
have resembled in any degree a Polyplectron.
But on continuing my search, I observed that
in some of the species the two ocelli stood
very near each other; that in the tail-feathers
of P. Hardwickii they touched each other; and,
finally, that in the tail-coverts of this same
species as well as of P. malaccense (fig. 55)
they were actually confluent. As the central
part alone is confluent, an indentation is left
at both the upper and lower ends; and the
surrounding coloured zones zones are
likewise indented.
A single ocellus is thus formed on each tail-
covert, though still plainly betraying its Fig. 54. Part of a tail-covert of
double origin. These confluent ocelli differ Polyplectron chinquis, with two
from the single ocelli of the peacock in oval ocelli of nat. size.
having an indentation at both ends, instead of
at the lower or basal end alone. The
explanation, however, of this difference is not
difficult; in some species of Polyplectron the
two oval ocelli on the same feather stand
parallel to each other; in other species (as in
P. chinquis) they converge towards one end;
now the partial confluence of two convergent
ocelli would manifestly leave a much deeper
indentation at the divergent than at the
convergent end. It is also manifest that if the
Fig. 55. Part of a tail-covert of
convergence were strongly pronounced and
Polyplectron malaccense, with
the confluence complete, the indentation at
the two oval ocelli, partially
the convergent end would tend to be quite
confluent, of nat. size.
obliterated.
The tail-feathers in both species of peacock are entirely destitute of ocelli,
and this apparently is related to their being covered up and concealed by the

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long tail-coverts. In this respect they differ remarkably from the tail-
feathers of Polyplectron, which in most of the species are ornamented with
larger ocelli than those on the tail-coverts. Hence I was led carefully to
examine the tail-feathers of the several species of Polyplectron in order to
discover whether the ocelli in any of them shewed any tendency to
disappear, and, to my great satisfaction, I was successful. The central tail-
feathers of P. Napoleonis have the two ocelli on each side of the shaft
perfectly developed; but the inner ocellus becomes less and less
conspicuous on the more exterior tail-feathers, until a mere shadow or
rudimentary vestige is left on the inner side of the outermost feather. Again,
in P. malaccense, the ocelli on the tail-coverts are, as we have seen,
confluent; and these feathers are of unusual length, being two-thirds of the
length of the tail-feathers, so that in both these respects they resemble the
tail-coverts of the peacock. Now in this species the two central tail-feathers
alone are ornamented, each with two brightly-coloured ocelli, the ocelli
having completely disappeared from the inner sides of all the other tail-
feathers. Consequently the tail-coverts and tail-feathers of this species of
Polyplectron make a near approach in structure and ornamentation to the
corresponding feathers of the peacock.
As far, then, as the principle of gradation throws light on the steps by which
the magnificent train of the peacock has been acquired, hardly anything
more is needed. We may picture to ourselves a progenitor of the peacock in
an almost exactly intermediate condition between the existing peacock,
with his enormously elongated tail-coverts, ornamented with single ocelli,
and an ordinary gallinaceous bird with short tail-coverts, merely spotted
with some colour; and we shall then see in our mind’s eye, a bird possessing
tail-coverts, capable of erection and expansion, ornamented with two
partially confluent ocelli, and long enough almost to conceal the tail-
feathers,—the latter having already partially lost their ocelli; we shall see in
short, a Polyplectron. The indentation of the central disc and surrounding
zones of the ocellus in both species of peacock, seems to me to speak
plainly in favour of this view; and this structure is otherwise inexplicable.
The males of Polyplectron are no doubt very beautiful birds, but their
beauty, when viewed from a little distance, cannot be compared, as I
formerly saw in the Zoological Gardens, with that of the peacock. Many
female progenitors of the peacock must, during a long line of descent, have

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appreciated this superiority; for they have unconsciously, by the continued
preference of the most beautiful males, rendered the peacock the most
splendid of living birds.
Argus pheasant.—Another excellent case for investigation is offered by the
ocelli on the wing-feathers of the Argus pheasant, which are shaded in so
wonderful a manner as to resemble balls lying within sockets, and which
consequently differ from ordinary ocelli. No one, I presume, will attribute
the shading, which has excited the admiration of many experienced artists,
to chance—to the fortuitous concourse of atoms of colouring matter. That
these ornaments should have been formed through the selection of many
successive variations, not one of which was originally intended to produce
the ball-and-socket effect, seems as incredible, as that one of Raphael’s
Madonnas should have been formed by the selection of chance daubs of
paint made by a long succession of young artists, not one of whom intended
at first to draw the human figure. In order to discover how the ocelli have
been developed, we cannot look to a long line of progenitors, nor to various
closely-allied forms, for such do not now exist. But fortunately the several
feathers on the wing suffice to give us a clue to the problem, and they prove
to demonstration that a gradation is at least possible from a mere spot to a
finished ball-and-socket ocellus.

The wing-feathers, bearing the ocelli, are covered with dark stripes or rows
of dark spots, each stripe or row running obliquely down the outer side of
the shaft to an ocellus. The spots are generally elongated in a transverse line
to the row in which they stand. They often become confluent, either in the
line of the row—and then they form a longitudinal stripe—or transversely,
that is, with the spots in the adjoining rows, and then they form transverse
stripes. A spot sometimes breaks up into smaller spots, which still stand in
their proper places.
It will be convenient first to describe a perfect ball-and-socket ocellus. This
consists of an intensely black circular ring, surrounding a space shaded so
as exactly to resemble a ball. The figure here given has been admirably
drawn by Mr. Ford, and engraved, but a woodcut cannot exhibit the
exquisite shading of the original. The ring is almost always slightly broken
or interrupted (see fig. 56) at a point in the upper half, a little to the right of

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and above the white shade on the enclosed ball; it is also sometimes broken
towards the base on the right hand. These little breaks have an important
meaning. The ring is always much thickened, with the edges ill-defined
towards the left-hand upper corner the feather being held erect, in the
position in which it is here
drawn. Beneath this thickened
part there is on the surface of the
ball an oblique almost pure-white
mark, which shades off
downwards into a pale-leaden
hue, and this into yellowish and
brown tints, which insensibly
become darker and darker
towards the lower part of the ball.
It is this shading which gives so
admirably the effect of light Fig. 56. Part of Secondary
shining on a convex surface. If wing-feather of Argus pheasant, shewing two, a
one of the balls be examined, it and b, perfect ocelli. A, B, C, &c., dark stripes
will be seen that the lower part is running obliquely down, each to an ocellus.
of a browner tint and is [Much of the web on both sides, especially to the
indistinctly separated by a curved left of the shaft, has been cut off.]
oblique line from the upper part,
which is yellower and more leaden; this oblique line runs at right angles to
the longer axis of the white patch of light, and indeed of all the shading; but
this difference in the tints, which cannot of course be shewn in the woodcut,
does not in the least interfere with the perfect shading of the ball.195 It
should be particularly observed that each ocellus stands in obvious
connection with a dark stripe, or row of dark spots, for both occur
indifferently on the same feather. Thus in fig. 56 stripe A runs to ocellus a;
B runs to ocellus b; stripe C is broken in the upper part, and runs down to
the next succeeding ocellus, not represented in the woodcut; D to the next
lower one, and so with the stripes E and F. Lastly, the several ocelli are
separated from each other by a pale surface bearing irregular black marks.
I will next describe the other extreme of the series, namely the first trace of
an ocellus. The short secondary wing-feather (fig. 57), nearest to the body,
is marked like the other feathers, with oblique, longitudinal, rather irregular,

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rows of spots. The lowest spot,
or that nearest the shaft, in the
five lower rows (excluding the
basal row) is a little larger than
the other spots in the same
row, and a little more
elongated in a transverse
direction. It differs also from
the other spots by being
bordered on its upper side with
some dull fulvous shading. But
this spot is not in any way
more remarkable than those on
the plumage of many birds,
and might easily be quite
overlooked. The next higher
spot in each row does not
differ at all from the upper
ones in the same row, although
in the following series it
becomes, as we shall see,
Fig. 57. Basal part of the Secondary wing-feather, greatly modified. The larger
nearest to the body. spots occupy exactly the same
relative position on this feather
as those occupied by the perfect ocelli on the longer wing-feathers.
By looking to the next two or three succeeding secondary wing-feathers, an
absolutely insensible gradation can be traced from one of the above-
described lower spots, together with the next higher one in the same row, to
a curious ornament, which cannot be called an ocellus, and which I will
name, from the want of a better term, an “elliptic ornament.” These are
shewn in the accompanying figure (fig. 58). We here see several oblique
rows, A, B, C, D (see the lettered diagram), &c., of dark spots of the usual
character. Each row of spots runs down to and is connected with one of the
elliptic ornaments, in exactly the same manner as each stripe in fig. 56 runs
down to, and is connected with, one of the ball-and-socket ocelli. Looking
to any one row, for instance, B, the lowest spot or mark (b) is thicker and

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considerably longer than the upper spots, and has its left extremity pointed
and curved upwards. This black mark is abruptly bordered on its upper side
by a rather broad space of richly-shaded tints, beginning with a narrow
brown zone, which passes into orange, and this into a pale leaden tint, with
the end towards the shaft much paler. This mark corresponds in every
respect with the larger, shaded spot, described in the last paragraph (fig.
57), but is more highly developed and more brightly coloured. To the right
and above this spot (b), with its bright shading, there is a long, narrow,
black mark (c), belonging to the same row, and which is arched a little
downwards so as to face (b). It is also narrowly edged on the lower side
with a fulvous tint. To the left of and above c, in the same oblique direction,
but always more or less distinct from it, there is another black mark (d).
This mark is generally sub-triangular and irregular in shape, but in the one
lettered in the diagram is unusually narrow, elongated, and regular. It
apparently consists of a lateral and broken prolongation of the mark (c), as I
infer from traces of similar prolongations from the succeeding upper spots;
but I do not feel sure of this. These three marks, b, c, and d, with the
intervening bright shades, form together the so-called elliptic ornament.
These ornaments stand in a line parallel to the shaft, and manifestly
correspond in position with the ball-and-socket ocelli. Their extremely
elegant appearance cannot be appreciated in the drawing, as the orange and
leaden tints, contrasting so well with the black marks, cannot be shewn.

Fig. 58. Portion of one of the Secondary wing-feathers near to the body; shewing the so-called
elliptic ornaments. The right-hand figure is given merely as a diagram for the sake of the letters of
reference.

A, B, C, &c. Rows of spots running down to and c. The next succeeding spot or mark in the same
forming the elliptic ornaments. row.
d. Apparently a broken prolongation of the spot c
b. Lowest spot or mark in row B.
in the same row B.

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Between one of the elliptic ornaments and a perfect ball-and-socket ocellus,
the gradation is so perfect that it is scarcely possible to decide when the
latter term ought to be used. I regret that I have not given an additional
drawing, besides fig. 58, which stands about half-way in the series between
one of the simple spots and a perfect ocellus. The passage from the elliptic
ornament into an ocellus is effected by the elongation and greater curvature
in opposed directions of the lower black mark (b), and more especially of
the upper one (c), together with the contraction of the irregular sub-
triangular or narrow mark (d), so that at last these three marks become
confluent, forming an irregular elliptic ring. This ring is gradually rendered
more and more circular and regular, at the same time increasing in diameter.
Traces of the junction of all three elongated spots or marks, especially of
the two upper ones, can still be observed in many of the most perfect ocelli.
The broken state of the black ring on the upper side of the ocellus in fig. 56
was pointed out. The irregular sub-triangular or narrow mark (d) manifestly
forms, by its contraction and equalisation, the thickened portion of the ring
on the left upper side of the perfect ball-and-socket ocellus. The lower part
of the ring is invariably a little thicker than the other parts (see fig. 56), and
this follows from the lower black mark of the elliptic ornament (b) having
been originally thicker than the upper mark (c). Every step can be followed
in the process of confluence and modification; and the black ring which
surrounds the ball of the ocellus is unquestionably formed by the union and
modification of the three black marks, b, c, d, of the elliptic ornament. The
irregular zigzag black marks between the successive ocelli (see again fig.
56) are plainly due to the breaking up of the somewhat more regular but
similar marks between the elliptic ornaments.
The successive steps in the shading of the ball-and-socket ocelli can be
followed out with equal clearness. The brown, orange, and pale-leaden
narrow zones which border the lower black mark of the elliptic ornament
can be seen gradually to become more and more softened and shaded into
each other, with the upper lighter part towards the left-hand corner rendered
still lighter, so as to become almost white. But even in the most perfect ball-
and-socket ocelli a slight difference in the tints, though not in the shading,
between the upper and lower parts of the ball can be perceived (as was
before especially noticed), the line of separation being oblique, in the same
direction with the bright coloured shades of the elliptic ornaments. Thus

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almost every minute detail in the shape and colouring of the ball-and-socket
ocelli can be shewn to follow from gradual changes in the elliptic
ornaments; and the development of the latter can be traced by equally small
steps from the union of two almost simple spots, the lower one (fig. 57)
having some dull fulvous shading on the upper side.
The extremities of the longer secondary feathers which bear the perfect
ball-and-socket ocelli are peculiarly ornamented. (Fig. 59.) The oblique
longitudinal stripes suddenly cease upwards and become confused, and
above this limit the whole upper end of the feather (a) is covered with white
dots, surrounded by little black rings, standing on a dark ground. Even the
oblique stripe belonging to the
uppermost ocellus (b) is represented
only by a very short irregular black
mark with the usual, curved, transverse
base. As this stripe is thus abruptly cut
off above, we can understand, from
what has gone before, how it is that the
upper thickened part of the ring is
absent in the uppermost ocellus; for, as
before stated, this thickened part is
apparently formed by a broken
prolongation of the next higher spot in
the same row. From the absence of the Fig. 59. Portion near
upper and thickened part of the ring, summit of one of the Secondary wing-
the uppermost ocellus, though perfect feathers, bearing perfect ball-and-socket
in all other respects, appears as if its top ocelli.
had been obliquely sliced off. It would, a. Ornamented upper part.
I think, perplex any one, who believes b. Uppermost, imperfect ball-and-socket
that the plumage of the Argus pheasant ocellus. (The shading above the white
was created as we now see it, to mark on the summit of the ocellus is here
account for the imperfect condition of a little too dark.)
the uppermost ocelli. I should add that c. Perfect ocellus.
in the secondary wing-feather farthest
from the body all the ocelli are smaller and less perfect than on the other
feathers, with the upper parts of the external black rings deficient, as in the
case just mentioned. The imperfection here seems to be connected with the

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fact that the spots on this feather shew less tendency than usual to become
confluent into stripes; on the contrary, they are often broken up into smaller
spots, so that two or three rows run down to each ocellus.
We have now seen that a perfect series can be followed, from two almost
simple spots, at first quite distinct from each other, to one of the wonderful
ball-and-socket ornaments. Mr. Gould, who kindly gave me some of these
feathers, fully agrees with me in the completeness of the gradation. It is
obvious that the stages in development exhibited by the feathers on the
same bird do not at all necessarily shew us the steps which have been
passed through by the extinct progenitors of the species; but they probably
give us the clue to the actual steps, and they at least prove to demonstration
that a gradation is possible. Bearing in mind how carefully the male Argus
pheasant displays his plumes before the female, as well as the many facts
rendering it probable that female birds prefer the more attractive males, no
one who admits the agency of sexual selection, will deny that a simple dark
spot with some fulvous shading might be converted, through the
approximation and modification of the adjoining spots, together with some
slight increase of colour, into one of the so-called elliptic ornaments. These
latter ornaments have been shewn to many persons, and all have admitted
that they are extremely pretty, some thinking them even more beautiful than
the ball-and-socket ocelli. As the secondary plumes became lengthened
through sexual selection, and as the elliptic ornaments increased in
diameter, their colours apparently became less bright; and then the
ornamentation of the plumes had to be gained by improvements in the
pattern and shading; and this process has been carried on until the
wonderful ball-and-socket ocelli have been finally developed. Thus we can
understand—and in no other way as it seems to me—the present condition
and origin of the ornaments on the wing-feathers of the Argus pheasant.
From the light reflected by the principle of gradation; from what we know
of the laws of variation; from the changes which have taken place in many
of our domesticated birds; and, lastly, from the character (as we shall
hereafter more clearly see) of the immature plumage of young birds—we
can sometimes indicate with a certain amount of confidence, the probable
steps by which the males have acquired their brilliant plumage and various
ornaments; yet in many cases we are involved in darkness. Mr. Gould
several years ago pointed out to me a humming-bird, the Urosticte

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benjamini, remarkable from the curious differences presented by the two
sexes. The male, besides a splendid gorget, has greenish-black tail-feathers,
with the four central ones tipped with white; in the female, as with most of
the allied species, the three outer tail-feathers on each side are tipped with
white, so that the male has the four central, whilst the female has the six
exterior feathers ornamented with white tips. What makes the case curious
is that, although the colouring of the tail differs remarkably in both sexes of
many kinds of humming-birds, Mr. Gould does not know a single species,
besides the Urosticte, in which the male has the four central feathers tipped
with white.

The Duke of Argyll, in commenting on this case,196 passes over sexual
selection, and asks, “What explanation does the law of natural selection
give of such specific varieties as these?” He answers “none whatever;” and
I quite agree with him. But can this be so confidently said of sexual
selection? Seeing in how many ways the tail-feathers of humming-birds
differ, why should not the four central feathers have varied in this one
species alone, so as to have acquired white tips? The variations may have
been gradual, or somewhat abrupt as in the case recently given of the
humming-birds near Bogota, in which certain individuals alone have the
“central tail-feathers tipped with beautiful green.” In the female of the
Urosticte I noticed extremely minute or rudimental white tips to the two
outer of the four central black tail-feathers; so that here we have an
indication of change of some kind in the plumage of this species. If we
grant the possibility of the central tail-feathers of the male varying in
whiteness, there is nothing strange in such variations having been sexually
selected. The white tips, together with the small white ear-tufts, certainly
add, as the Duke of Argyll admits, to the beauty of the male; and whiteness
is apparently appreciated by other birds, as may be inferred from such cases
as the snow-white male of the Bell-bird. The statement made by Sir E.
Heron should not be forgotten, namely that his peahens, when debarred
from access to the pied peacock, would not unite with any other male, and
during that season produced no offspring. Nor is it strange that variations in
the tail-feathers of the Urosticte should have been specially selected for the
sake of ornament, for the next succeeding genus in the family takes its
name of Metallura from the splendour of these feathers. Mr. Gould, after
describing the peculiar plumage of the Urosticte, adds, “that ornament and

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variety is the sole object, I have myself but little doubt.”197 If this be
admitted, we can perceive that the males which were decked in the most
elegant and novel manner would have gained an advantage, not in the
ordinary struggle for life, but in rivalry with other males, and would
consequently have left a larger number of offspring to inherit their newly-
acquired beauty.

CHAPTER XV.

Birds—continued.

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Discussion why the males alone of some species, and both sexes of other species, are brightly
coloured—On sexually-limited inheritance, as applied to various structures and to brightly-
coloured plumage—Nidification in relation to colour—Loss of nuptial plumage during the
winter.

We have in this chapter to consider, why with many kinds of birds the
female has not received the same ornaments as the male; and why with
many others, both sexes are equally, or almost equally, ornamented? In the
following chapter we shall consider why in some few rare cases the female
is more conspicuously coloured than the male.
In my ‘Origin of Species’198 I briefly suggested that the long tail of the
peacock would be inconvenient, and the conspicuous black colour of the
male capercailzie dangerous, to the female during the period of incubation;
and consequently that the transmission of these characters from the male to
the female offspring had been checked through natural selection. I still think
that this may have occurred in some few instances: but after mature
reflection on all the facts which I have been able to collect, I am now
inclined to believe that when the sexes differ, the successive variations have
generally been from the first limited in their transmission to the same sex in
which they first appeared. Since my remarks appeared, the subject of sexual
coloration has been discussed in some very interesting papers by Mr.
Wallace,199 who believes that in almost all cases the successive variations
tended at first to be transmitted equally to both sexes; but that the female
was saved, through natural selection, from acquiring the conspicuous
colours of the male, owing to the danger which she would thus have
incurred during incubation.
This view necessitates a tedious discussion on a difficult point, namely
whether the transmission of a character, which is at first inherited by both
sexes, can be subsequently limited in its transmission, by means of
selection, to one sex alone. We must bear in mind, as shewn in the
preliminary chapter on sexual selection, that characters which are limited in
their development to one sex are always latent in the other. An imaginary
illustration will best aid us in seeing the difficulty of the case: we may
suppose that a fancier wished to make a breed of pigeons, in which the
males alone should be coloured of a pale blue, whilst the females retained
their former slaty tint. As with pigeons characters of all kinds are usually
transmitted to both sexes equally, the fancier would have to try to convert

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this latter form of inheritance into sexually-limited transmission. All that he
could do would be to persevere in selecting every male pigeon which was in
the least degree of a paler blue; and the natural result of this process, if
steadily carried on for a long time, and if the pale variations were strongly
inherited or often recurred, would be to make his whole stock of a lighter
blue. But our fancier would be compelled to match, generation after
generation, his pale blue males with slaty females, for he wishes to keep the
latter of this colour. The result would generally be the production either of a
mongrel piebald lot, or more probably the speedy and complete loss of the
pale-blue colour, for the primordial slaty tint would be transmitted with
prepotent force. Supposing, however, that some pale-blue males and slaty
females were produced during each successive generation, and were always
crossed together; then the slaty females would have, if I may use the
expression, much blue blood in their veins, for their fathers, grandfathers,
etc., will all have been blue birds. Under these circumstances it is
conceivable (though I know of no distinct facts rendering it probable) that
the slaty females might acquire so strong a latent tendency to pale-blueness,
that they would not destroy this colour in their male offspring, their female
offspring still inheriting the slaty tint. If so, the desired end of making a
breed with the two sexes permanently different in colour might be gained.
The extreme importance, or rather necessity, of the desired character in the
above case, namely, pale-blueness, being present though in a latent state in
the female, so that the male offspring should not be deteriorated, will be
best appreciated as follows: the male of Sœmmerring’s pheasant has a tail
thirty-seven inches in length, whilst that of the female is only eight inches;
the tail of the male common pheasant is about twenty inches, and that of the
female twelve inches long. Now if the female Sœmmerring pheasant with
her short tail were crossed with the male common pheasant, there can be no
doubt that the male hybrid offspring would have a much longer tail than
that of the pure offspring of the common pheasant. On the other hand, if the
female common pheasant, with her tail nearly twice as long as that of the
female Sœmmerring pheasant, were crossed with the male of the latter, the
male hybrid offspring would have a much shorter tail than that of the pure
offspring of Sœmmerring’s pheasant.200
Our fancier, in order to make his new breed with the males of a decided
pale-blue tint, and the females unchanged, would have to continue selecting

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the males during many generations; and each stage of paleness would have
to be fixed in the males, and rendered latent in the females. The task would
be an extremely difficult one, and has never been tried, but might possibly
succeed. The chief obstacle would be the early and complete loss of the
pale-blue tint, from the necessity of reiterated crosses with the slaty female,
the latter not having at first any latent tendency to produce pale-blue
offspring.
On the other hand, if one or two males were to vary ever so slightly in
paleness, and the variations were from the first limited in their transmission
to the male sex, the task of making a new breed of the desired kind would
be easy, for such males would simply have to be selected and matched with
ordinary females. An analogous case has actually occurred, for there are
breeds of the pigeon in Belgium201 in which the males alone are marked
with black striæ. In the case of the fowl, variations of colour limited in their
transmission to the male sex habitually occur. Even when this form of
inheritance prevails, it might well happen that some of the successive steps
in the process of variation might be transferred to the female, who would
then come to resemble in a slight degree the male, as occurs in some breeds
of the fowl. Or again, the greater number, but not all, of the successive steps
might be transferred to both sexes, and the female would then closely
resemble the male. There can hardly be a doubt that this is the cause of the
male pouter pigeon having a somewhat larger crop, and of the male carrier
pigeon having somewhat larger wattles, than their respective females; for
fanciers have not selected one sex more than the other, and have had no
wish that these characters should be more strongly displayed in the male
than in the female, yet this is the case with both breeds.
The same process would have to be followed, and the same difficulties
would be encountered, if it were desired to make a breed with the females
alone of some new colour.
Lastly, our fancier might wish to make a breed with the two sexes differing
from each other, and both from the parent-species. Here the difficulty would
be extreme, unless the successive variations were from the first sexually
limited on both sides, and then there would be no difficulty. We see this
with the fowl; thus the two sexes of the pencilled Hamburghs differ greatly
from each other, and from the two sexes of the aboriginal Gallus bankiva;

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and both are now kept constant to their standard of excellence by continued
selection, which would be impossible unless the distinctive characters of
both were limited in their transmission. The Spanish fowl offers a more
curious case; the male has an immense comb, but some of the successive
variations, by the accumulation of which it was acquired, appear to have
been transferred to the female; for she has a comb many times larger than
that of the females of the parent-species. But the comb of the female differs
in one respect from that of the male, for it is apt to lop over; and within a
recent period it has been ordered by the fancy that this should always be the
case, and success has quickly followed the order. Now the lopping of the
comb must be sexually limited in its transmission, otherwise it would
prevent the comb of the male from being perfectly upright, which would be
abhorrent to every fancier. On the other hand the uprightness of the comb in
the male must likewise be a sexually-limited character, otherwise it would
prevent the comb of the female from lopping over.
From the foregoing illustrations, we see that even with almost unlimited
time at command, it would be an extremely difficult and complex process,
though perhaps not impossible, to change through selection one form of
transmission into the other. Therefore, without distinct evidence in each
case, I am unwilling to admit that this has often been effected with natural
species. On the other hand by means of successive variations, which were
from the first sexually limited in their transmission, there would not be the
least difficulty in rendering a male bird widely different in colour or in any
other character from the female; the latter being left unaltered, or slightly
altered, or specially modified for the sake of protection.
As bright colours are of service to the males in their rivalry with other
males, such colours would be selected, whether or not they were transmitted
exclusively to the same sex. Consequently the females might be expected
often to partake of the brightness of the males to a greater or less degree;
and this occurs with a host of species. If all the successive variations were
transmitted equally to both sexes, the females would be undistinguishable
from the males; and this likewise occurs with many birds. If, however, dull
colours were of high importance for the safety of the female during
incubation, as with many ground birds, the females which varied in
brightness, or which received through inheritance from the males any
marked accession of brightness, would sooner or later be destroyed. But the

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tendency in the males to continue for an indefinite period transmitting to
their female offspring their own brightness, would have to be eliminated by
a change in the form of inheritance; and this, as shewn by our previous
illustration, would be extremely difficult. The more probable result of the
long-continued destruction of the more brightly-coloured females,
supposing the equal form of transmission to prevail, would be the lessening
or annihilation of the bright colours of the males, owing to their continually
crossing with the duller females. It would be tedious to follow out all the
other possible results; but I may remind the reader, as shewn in the eighth
chapter, that if sexually-limited variations in brightness occurred in the
females, even if they were not in the least injurious to them and
consequently were not eliminated, yet they would not be favoured or
selected, for the male usually accepts any female, and does not select the
more attractive individuals; consequently these variations would be liable to
be lost, and would have little influence on the character of the race; and this
will aid in accounting for the females being commonly less brightly-
coloured than the males.
In the chapter just referred to, instances were given, and any number might
have been added, of variations occurring at different ages, and inherited at
the same age. It was also shewn that variations which occur late in life are
commonly transmitted to the same sex in which they first appeared; whilst
variations occurring early in life are apt to be transmitted to both sexes; not
that all the cases of sexually-limited transmission can thus be accounted for.
It was further shewn that if a male bird varied by becoming brighter whilst
young, such variations would be of no service until the age for reproduction
had arrived, and there was competition between rival males. If we suppose
that three-fourths of the young males of any species are on an average
destroyed by various enemies; then the chances would be as three to one
against any one individual more brightly-coloured than usual surviving to
propagate its kind. But in the case of birds which live on the ground and
which commonly need the protection of dull colours, bright tints would be
far more dangerous to the young and inexperienced than to the adult males.
Consequently the males which varied in brightness whilst young would
suffer much destruction and be eliminated through natural selection; on the
other hand the males which varied in this manner when nearly mature,
notwithstanding that they were exposed to some additional danger, might

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survive, and from being favoured through sexual selection, would procreate
their kind. The brightly-coloured young males being destroyed and the
mature ones being successful in their courtship, may account, on the
principle of a relation existing between the period of variation and the form
of transmission, for the males alone of many birds, having acquired and
transmitted brilliant colours to their male offspring alone. But I by no
means wish to maintain that the influence of age on the form of
transmission is indirectly the sole cause of the great difference in brilliancy
between the sexes of many birds.
As with all birds in which the sexes differ in colour, it is an interesting
question whether the males alone have been modified through sexual
selection, the females being left, as far as this agency is concerned,
unchanged or only partially changed; or whether the females have been
specially modified through natural selection for the sake of protection, I
will discuss this question at considerable length, even at greater length than
its intrinsic importance deserves; for various curious collateral points may
thus be conveniently considered.
Before we enter on the subject of colour, more especially in reference to Mr.
Wallace’s conclusions, it may be useful to discuss under a similar point of
view some other differences between the sexes. A breed of fowls formerly
existed in Germany202 in which the hens were furnished with spurs; they
were good layers, but they so greatly disturbed their nests with their spurs
that they could not be allowed to sit on their own eggs. Hence at one time it
appeared to me probable that with the females of the wild Gallinaceæ the
development of spurs had been checked through natural selection, from the
injury thus caused to their nests. This seemed all the more probable as the
wing-spurs, which could not be injurious during nidification, are often as
well developed in the female as in the male; though in not a few cases they
are rather larger in the male. When the male is furnished with leg-spurs the
female almost always exhibits rudiments of them,—the rudiment sometimes
consisting of a mere scale, as with the species of Gallus. Hence it might be
argued that the females had aboriginally been furnished with well-
developed spurs, but that these had subsequently been lost either through
disuse or natural selection. But if this view be admitted, it would have to be
extended to innumerable other cases; and it implies that the female

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progenitors of the existing spur-bearing species were once encumbered with
an injurious appendage.
In some few genera and species, as in Galloperdix, Acomus, and the Javan
peacock (Pavo muticus), the females, as well as the males, possess well-
developed spurs. Are we to infer from this fact that they construct a
different sort of nest, not liable to be injured by their spurs, from that made
by their nearest allies, so that there has been no need for the removal of
their spurs? Or are we to suppose that these females especially require spurs
for their defence? It is a more probable conclusion that both the presence
and absence of spurs in the females result from different laws of inheritance
having prevailed, independently of natural selection. With the many females
in which spurs appear as rudiments, we may conclude that some few of the
successive variations, through which they were developed in the males,
occurred very early in life, and were as a consequence transferred to the
females. In the other and much rarer cases, in which the females possess
fully developed spurs, we may conclude that all the successive variations
were transferred to them; and that they gradually acquired the inherited
habit of not disturbing their nests.
The vocal organs and the variously-modified feathers for producing sound,
as well as the proper instincts for using them, often differ in the two sexes,
but are sometimes the same in both. Can such differences be accounted for
by the males having acquired these organs and instincts, whilst the females
have been saved from inheriting them, on account of the danger to which
they would have been exposed by attracting the attention of birds or beasts
of prey? This does not seem to me probable, when we think of the multitude
of birds which with impunity gladden the country with their voices during
the spring.203 It is a safer conclusion that as vocal and instrumental organs
are of special service only to the males during their courtship, these organs
were developed through sexual selection and continued use in this sex alone
—the successive variations and the effects of use having been from the first
limited in their transmission in a greater or less degree to the male
offspring.
Many analogous cases could be advanced; for instance the plumes on the
head, which are generally longer in the male than in the female, sometimes
of equal length in both sexes, and occasionally absent in the female,—these

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several cases sometimes occurring in the same group of birds. It would be
difficult to account for a difference of this kind between the sexes on the
principle of the female having been benefited by possessing a slightly
shorter crest than the male, and its consequent diminution or complete
suppression through natural selection. But I will take a more favourable
case, namely, the length of the tail. The long train of the peacock would
have been not only inconvenient but dangerous to the peahen during the
period of incubation and whilst accompanying her young. Hence there is
not the least à priori improbability in the development of her tail having
been checked through natural selection. But the females of various
pheasants, which apparently are exposed on their open nests to as much
danger as the peahen, have tails of considerable length. The females as well
as the males of the Menura superba have long tails, and they build a domed
nest, which is a great anomaly in so large a bird. Naturalists have wondered
how the female Menura could manage her tail during incubation; but it is
now known204 that she “enters the nest head first, and then turns round with
her tail sometimes over her back, but more often bent round by her side.
Thus in time the tail becomes quite askew, and is a tolerable guide to the
length of time the bird has been sitting.” Both sexes of an Australian
kingfisher (Tanysiptera sylvia) have the middle tail-feathers greatly
lengthened; and as the female makes her nest in a hole, these feathers
become, as I am informed by Mr. R. B. Sharpe, much crumpled during
nidification.
In these two cases the great length of the tail-feathers must be in some
degree inconvenient to the female; and as in both species the tail-feathers of
the female are somewhat shorter than those of the male, it might be argued
that their full development had been prevented through natural selection.
Judging from these cases, if with the peahen, the development of the tail
had been checked only when it became inconveniently or dangerously long,
she would have acquired a much longer tail than she actually possesses; for
her tail is not nearly so long, relatively to the size of her body, as that of
many female pheasants, nor longer than that of the female turkey. It must
also be borne in mind, that in accordance with this view as soon as the tail
of the peahen became dangerously long, and its development was
consequently checked, she would have continually reacted on her male
progeny, and thus have prevented the peacock from acquiring his present

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magnificent train. We may therefore infer that the length of the tail in the
peacock and its shortness in the peahen are the result of the requisite
variations in the male having been from the first transmitted to the male
offspring alone.
We are led to a nearly similar conclusion with respect to the length of the
tail in the various species of pheasants. In the Eared pheasant (Crossoptilon
auritum) the tail is of equal length in both sexes, namely, sixteen or
seventeen inches; in the common pheasant it is about twenty inches long in
the male, and twelve in the female; in Sœmmerring’s pheasant, thirty-seven
inches in the male, and only eight in the female; and lastly in Reeve’s
pheasant it is sometimes actually seventy-two inches long in the male and
sixteen in the female. Thus in the several species, the tail of the female
differs much in length, irrespectively of that of the male; and this can be
accounted for as it seems to me, with much more probability, by the laws of
inheritance,—that is by the successive variations having been from the first
more or less closely limited in their transmission to the male sex,—than by
the agency of natural selection, owing to the length of tail having been
injurious in a greater or less degree to the females of the several species.
We may now consider Mr. Wallace’s arguments, in regard to the sexual
coloration of birds. He believes that the bright tints originally acquired
through sexual selection by the males, would in all or almost all cases have
been transmitted to the females, unless the transference had been checked
through natural selection. I may here remind the reader that various facts
bearing on this view have already been given under reptiles, amphibians,
fishes, and lepidoptera. Mr. Wallace rests his belief chiefly, but not
exclusively, as we shall see in the next chapter, on the following
statement,205 that when both sexes are coloured in a strikingly-conspicuous
manner the nest is of such a nature as to conceal the sitting bird; but when
there is a marked contrast of colour between the sexes, the male being gay
and the female dull-coloured, the nest is open and exposes the sitting bird to
view. This coincidence, as far as it goes, certainly supports the belief that
the females which sit on open nests have been specially modified for the
sake of protection. Mr. Wallace admits that there are, as might have been
expected, some exceptions to his two rules, but it is a question whether the
exceptions are not so numerous as seriously to invalidate them.

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There is in the first place much truth in the Duke of Argyll’s remark206 that a
large domed nest is more conspicuous to an enemy, especially to all tree-
haunting carnivorous animals, than a smaller open nest. Nor must we forget
that with many birds which build open nests the males sit on the eggs and
aid in feeding the young as well as the females: this is the case, for instance,
with Pyranga æstiva,207 one of the most splendid birds in the United States,
the male being vermilion, and the female light brownish-green. Now if
brilliant colours had been extremely dangerous to birds whilst sitting on
their open nests, the males in these cases would have suffered greatly. It
might, however, be of such paramount importance to the male to be
brilliantly coloured, in order to beat his rivals, that this would more than
compensate for some additional danger.
Mr. Wallace admits that with the King-crows (Dicrurus), Orioles, and
Pittidæ, the females are conspicuously coloured, yet they build open nests;
but he urges that the birds of the first group are highly pugnacious and
could defend themselves; that those of the second group take extreme care
in concealing their open nests, but this does not invariably hold good;208 and
that with the birds of the third group the females are brightly coloured
chiefly on the under surface. Besides these cases the whole great family of
pigeons, which are sometimes brightly, and almost always conspicuously
coloured, and which are notoriously liable to the attacks of birds of prey,
offers a serious exception to the rule, for pigeons almost always build open
and exposed nests. In another large family, that of the Humming-birds, all
the species build open nests, yet with some of the most gorgeous species the
sexes are alike; and in the majority, the females, though less brilliant than
the males, are very brightly coloured. Nor can it be maintained that all
female humming-birds, which are brightly coloured, escape detection by
their tints being green, for some display on their upper surfaces red, blue,
and other colours.209
In regard to birds which build in holes or construct domed nests, other
advantages, as Mr. Wallace remarks, besides concealment are gained, such
as shelter from the rain, greater warmth, and in hot countries protection
from the rays of the sun;210 so that it is no valid objection to his view that
many birds having both sexes obscurely coloured build concealed nests.211
The female Hornbills (Buceros), for instance, of India and Africa are
protected, during nidification, with extraordinary care, for the male plaisters

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up the hole in which the female sits on her eggs, and leaves only a small
orifice through which he feeds her; she is thus kept a close prisoner during
the whole period of incubation;212 yet female hornbills are not more
conspicuously coloured than many other birds of equal size which build
open nests. It is a more serious objection to Mr. Wallace’s view, as is
admitted by him, that in some few groups the males are brilliantly coloured
and the females obscure, and yet the latter hatch their eggs in domed nests.
This is the case with the Grallinæ of Australia, the Superb Warblers
(Maluridæ) of the same country, the Sun-birds (Nectariniæ), and with
several of the Australian Honey-suckers or Meliphagidæ.213
If we look to the birds of England we shall see that there is no close and
general relation between the colours of the female and the nature of the nest
constructed by her. About forty of our British birds (excluding those of
large size which could defend themselves) build in holes in banks, rocks, or
trees, or construct domed nests. If we take the colours of the female
goldfinch, bullfinch, or blackbird, as a standard of the degree of
conspicuousness, which is not highly dangerous to the sitting female, then
out of the above forty birds, the females of only twelve can be considered as
conspicuous to a dangerous degree, the remaining twenty-eight being
inconspicuous.214 Nor is there any close relation between a well-pronounced
difference in colour between the two sexes, and the nature of the nest
constructed. Thus the male house-sparrow (Passer domesticus) differs
much from the female, the male tree-sparrow (P. montanus) differs hardly at
all, and yet both build well-concealed nests. The two sexes of the common
fly-catcher (Muscicapa grisola) can hardly be distinguished, whilst the
sexes of the pied fly-catcher (M. luctuosa) differ considerably, and both
build in holes. The female blackbird (Turdus merula) differs much, the
female ring-ouzel (T. torquatus) differs less, and the female common thrush
(T. musicus) hardly at all from their respective males; yet all build open
nests. On the other hand, the not very distantly-allied water-ouzel (Cinclus
aquaticus) builds a domed nest, and the sexes differ about as much as in the
case of the ring-ouzel. The black and red grouse (Tetrao tetrix and T.
Scoticus) build open nests, in equally well-concealed spots, but in the one
species the sexes differ greatly, and in the other very little.
Notwithstanding the foregoing objections, I cannot doubt, after reading Mr.
Wallace’s excellent essay, that looking to the birds of the world, a large

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majority of the species in which the females are conspicuously coloured
(and in this case the males with rare exceptions are equally conspicuous),
build concealed nests for the sake of protection. Mr. Wallace enumerates215
a long series of groups in which this rule holds good; but it will suffice here
to give, as instances, the more familiar groups of kingfishers, toucans,
trogons, puff-birds (Capitonidæ), plaintain-eaters (Musophagæ),
woodpeckers, and parrots. Mr. Wallace believes that in these groups, as the
males gradually acquired through sexual selection their brilliant colours,
these were transferred to the females and were not eliminated by natural
selection, owing to the protection which they already enjoyed from their
manner of nidification. According to this view, their present manner of
nesting was acquired before their present colours. But it seems to me much
more probable that in most cases as the females were gradually rendered
more and more brilliant from partaking of the colours of the male, they
were gradually led to change their instincts (supposing that they originally
built open nests), and to seek protection by building domed or concealed
nests. No one who studies, for instance, Audubon’s account of the
differences in the nests of the same species in the Northern and Southern
United States,216 will feel any great difficulty in admitting that birds, either
by a change (in the strict sense of the word) of their habits, or through the
natural selection of so-called spontaneous variations of instinct, might
readily be led to modify their manner of nesting.
This way of viewing the relation, as far as it holds good, between the bright
colours of female birds and their manner of nesting, receives some support
from certain analogous cases occurring in the Sahara Desert. Here, as in
most other deserts, various birds, and many other animals, have had their
colours adapted in a wonderful manner to the tints of the surrounding
surface. Nevertheless there are, as I am informed by the Rev. Mr. Tristram,
some curious exceptions to the rule; thus the male of the Monticola cyanea
is conspicuous from his bright blue colour, and the female almost equally
conspicuous from her mottled brown and white plumage; both sexes of two
species of Dromolæa are of a lustrous black; so that these three birds are far
from receiving protection from their colours, yet they are able to survive,
for they have acquired the habit, when in danger, of taking refuge in holes
or crevices in the rocks.

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With respect to the above-specified groups of birds, in which the females
are conspicuously coloured and build concealed nests, it is not necessary to
suppose that each separate species had its nidifying instinct specially
modified; but only that the early progenitors of each group were gradually
led to build domed or concealed nests; and afterwards transmitted this
instinct, together with their bright colours, to their modified descendants.
This conclusion, as far as it can be trusted, is interesting, namely, that
sexual selection, together with equal or nearly equal inheritance by both
sexes, have indirectly determined the manner of nidification of whole
groups of birds.
Even in the groups in which, according to Mr. Wallace, the females from
being protected during nidification, have not had their bright colours
eliminated through natural selection, the males often differ in a slight, and
occasionally in a considerable degree, from the females. This is a
significant fact, for such differences in colour must be accounted for on the
principle of some of the variations in the males having been from the first
limited in their transmission to the same sex; as it can hardly be maintained
that these differences, especially when very slight, serve as a protection to
the female. Thus all the species in the splendid group of the Trogons build
in holes; and Mr. Gould gives figures217 of both sexes of twenty-five
species, in all of which, with one partial exception, the sexes differ
sometimes slightly, sometimes conspicuously, in colour,—the males being
always more beautiful than the females, though the latter are likewise
beautiful. All the species of kingfisher build in holes, and with most of the
species the sexes are equally brilliant, and thus far Mr. Wallace’s rule holds
good; but in some of the Australian species the colours of the females are
rather less vivid than those of the male; and in one splendidly-coloured
species, the sexes differ so much that they were at first thought to be
specifically distinct.218 Mr. R. B. Sharpe, who has especially studied this
group, has shewn me some American species (Ceryle) in which the breast
of the male is belted with black. Again, in Carcineutes, the difference
between the sexes is conspicuous: in the male the upper surface is dull-blue
banded with black, the lower surface being partly fawn-coloured, and there
is much red about the head; in the female the upper surface is reddish-
brown banded with black, and the lower surface white with black markings.
It is an interesting fact, as shewing how the same peculiar style of sexual

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colouring often characterises allied forms, that in three species of Dacelo
the male differs from the female only in the tail being dull-blue banded with
black, whilst that of the female is brown with blackish bars; so that here the
tail differs in colour in the two sexes in exactly the same manner as the
whole upper surface in the sexes of Carcineutes.
With parrots, which likewise build in holes, we find analogous cases: in
most of the species both sexes are brilliantly coloured and
undistinguishable, but in not a few species the males are coloured rather
more vividly than the females, or even very differently from them. Thus,
besides other strongly-marked differences, the whole under surface of the
male King Lory (Aprosmictus scapulatus) is scarlet, whilst the throat and
chest of the female is green tinged with red: in the Euphema splendida there
is a similar difference, the face and wing-coverts moreover of the female
being of a paler blue than in the male.219 In the family of the tits (PARINÆ),
which build concealed nests, the female of our common blue tomtit (Parus
cæruleus) is “much less brightly coloured” than the male; and in the
magnificent Sultan yellow tit of India the difference is greater.220
Again in the great group of the woodpeckers,221 the sexes are generally
nearly alike, but in the Megapicus validus all those parts of the head, neck,
and breast, which are crimson in the male are pale brown in the female. As
in several woodpeckers the head of the male is bright crimson, whilst that
of the female is plain, it occurred to me that this colour might possibly
make the female dangerously conspicuous, whenever she put her head out
of the hole containing her nest, and consequently that this colour, in
accordance with Mr. Wallace’s belief, had been eliminated. This view is
strengthened by what Malherbe states with respect to Indopicus carlotta;
namely, that the young females, like the young males, have some crimson
about their heads, but that this colour disappears in the adult female, whilst
it is intensified in the adult male. Nevertheless the following considerations
render this view extremely doubtful: the male takes a fair share in
incubation,222 and would be thus far almost equally exposed to danger; both
sexes of many species have their heads of an equally bright crimson; in
other species the difference between the sexes in the amount of scarlet is so
slight that it can hardly make any appreciable difference in the danger
incurred; and lastly, the colouring of the head in the two sexes often differs
slightly in other ways.

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The cases, as yet given, of slight and graduated differences in colour
between the males and females in the groups, in which as a general rule the
sexes resemble each other, all relate to species which build domed or
concealed nests. But similar gradations may likewise be observed in groups
in which the sexes as a general rule resemble each other, but which build
open nests. As I have before instanced the Australian parrots, so I may here
instance, without giving any details, the Australian pigeons.223 It deserves
especial notice that in all these cases the slight differences in plumage
between the sexes are of the same general nature as the occasionally greater
differences. A good illustration of this fact has already been afforded by
those kingfishers in which either the tail alone or the whole upper surface of
the plumage differs in the same manner in the two sexes. Similar cases may
be observed with parrots and pigeons. The differences in colour between the
sexes of the same species are, also, of the same general nature as the
differences in colour between the distinct species of the same group. For
when in a group in which the sexes are usually alike, the male differs
considerably from the female, he is not coloured in a quite new style. Hence
we may infer that within the same group the special colours of both sexes
when they are alike, and the colours of the male, when he differs slightly or
even considerably from the female, have in most cases been determined by
the same general cause; this being sexual selection.
It is not probable, as has already been remarked, that differences in colour
between the sexes, when very slight, can be of service to the female as a
protection. Assuming, however, that they are of service, they might be
thought to be cases of transition; but we have no reason to believe that
many species at any one time are undergoing change. Therefore we can
hardly admit that the numerous females which differ very slightly in colour
from their males are now all commencing to become obscure for the sake of
protection. Even if we consider somewhat more marked sexual differences,
is it probable, for instance, that the head of the female chaffinch, the
crimson on the breast of the female bullfinch,—the green of the female
greenfinch,—the crest of the female golden-crested wren, have all been
rendered less bright by the slow process of selection for the sake of
protection? I cannot think so; and still less with the slight differences
between the sexes of those birds which build concealed nests. On the other
hand, the differences in colour between the sexes, whether great or small,

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may to a large extent be explained on the principle of the successive
variations, acquired by the males through sexual selection, having been
from the first more or less limited in their transmission to the females. That
the degree of limitation should differ in different species of the same group
will not surprise any one who has studied the laws of inheritance, for they
are so complex that they appear to us in our ignorance to be capricious in
their action.224
As far as I can discover there are very few groups of birds containing a
considerable number of species, in which all have both sexes brilliantly
coloured and alike; but this appears to be the case, as I hear from Mr.
Sclater, with the Musophagæ or plaintain-eaters. Nor do I believe that any
large group exists in which the sexes of all the species are widely dissimilar
in colour: Mr. Wallace informs me that the chatterers of S. America
(COTINGIDÆ) offer one of the best instances; but with some of the
species, in which the male has a splendid red breast, the female exhibits
some red on her breast; and the females of other species shew traces of the
green and other colours of the males. Nevertheless we have a near approach
to close sexual similarity or dissimilarity throughout several groups: and
this, from what has just been said of the fluctuating nature of inheritance, is
a somewhat surprising circumstance. But that the same laws should largely
prevail with allied animals is not surprising. The domestic fowl has
produced a great number of breeds and sub-breeds, and in these the sexes
generally differ in plumage; so that it has been noticed as a remarkable
circumstance when in certain sub-breeds they resemble each other. On the
other hand, the domestic pigeon has likewise produced a vast number of
distinct breeds and sub-breeds, and in these, with rare exceptions, the two
sexes are identically alike. Therefore if other species of Gallus and
Columba were domesticated and varied, it would not be rash to predict that
the same general rules of sexual similarity and dissimilarity, depending on
the form of transmission, would, in both cases, hold good. In a similar
manner the same form of transmission has generally prevailed throughout
the same natural groups, although marked exceptions to this rule occur.
Within the same family or even genus, the sexes may be identically alike or
very different in colour. Instances have already been given relating to the
same genus, as with sparrows, fly-catchers, thrushes and grouse. In the
family of pheasants the males and females of almost all the species are

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wonderfully dissimilar, but are quite similar in the eared pheasant or
Crossoptilon auritum. In two species of Chloephaga, a genus of geese, the
males cannot be distinguished from the females, except by size; whilst in
two others, the sexes are so unlike that they might easily be mistaken for
distinct species.225
The laws of inheritance can alone account for the following cases, in which
the female by acquiring at a late period of life certain characters proper to
the male, ultimately comes to resemble him in a more or less complete
manner. Here protection can hardly have come into play. Mr. Blyth informs
me that the females of Oriolus melanocephalus and of some allied species,
when sufficiently mature to breed, differ considerably in plumage from the
adult males; but after the second or third moults they differ only in their
beaks having a slight greenish tinge. In the dwarf bitterns (Ardetta),
according to the same authority, “the male acquires his final livery at the
first moult, the female not before the third or fourth moult; in the
meanwhile she presents an intermediate garb, which is ultimately
exchanged for the same livery as that of the male.” So again the female
Falco peregrinus acquires her blue plumage more slowly than the male. Mr.
Swinhoe states that with one of the Drongo shrikes (Dicrurus macrocercus)
the male whilst almost a nestling, moults his soft brown plumage and
becomes of a uniform glossy greenish-black; but the female retains for a
long time the white striæ and spots on the axillary feathers; and does not
completely assume the uniform black colour of the male for the first three
years. The same excellent observer remarks that in the spring of the second
year the female spoonbill (Platalea) of China resembles the male of the first
year, and that apparently it is not until the third spring that she acquires the
same adult plumage as that possessed by the male at a much earlier age.
The female Bombycilla carolinensis differs very little from the male, but the
appendages, which like beads of red sealing-wax ornament the wing-
feathers, are not developed in her so early in life as in the male. The upper
mandible in the male of an Indian parrakeet (Palæornis Javanicus) is coral-
red from his earliest youth, but in the female, as Mr. Blyth has observed
with caged and wild birds, it is at first black and does not become red until
the bird is at least a year old, at which age the sexes resemble each other in
all respects. Both sexes of the wild turkey are ultimately furnished with a
tuft of bristles on the breast, but in two-year-old birds the tuft is about four

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inches long in the male and hardly apparent in the female; when, however,
the latter has reached her fourth year, it is from four to five inches in
length.226
In these cases, the females follow a normal course of development in
ultimately becoming like the males; and such cases must not be confounded
with those in which diseased or old females assume masculine characters,
or with those in which perfectly fertile females, whilst young, acquire
through variation or some unknown cause the characters of the male.227 But
all these cases have so much in common that they depend, according to the
hypothesis of pangenesis, on gemmules derived from each part of the male
being present, though latent, in the female; their development following on
some slight change in the elective affinities of her constituent tissues.
A few words must be added on changes of plumage in relation to the season
of the year. From reasons formerly assigned there can be little doubt that the
elegant plumes, long pendant feathers, crests, &c., of egrets, herons, and
many other birds, which are developed and retained only during the
summer, serve exclusively for ornamental or nuptial purposes, though
common to both sexes. The female is thus rendered more conspicuous
during the period of incubation than during the winter; but such birds as
herons and egrets would be able to defend themselves. As, however, plumes
would probably be inconvenient and certainly of no use during the winter, it
is possible that the habit of moulting twice in the year may have been
gradually acquired through natural selection for the sake of casting off
inconvenient ornaments during the winter. But this view cannot be extended
to the many waders, in which the summer and winter plumages differ very
little in colour. With defenceless species, in which either both sexes or the
males alone become extremely conspicuous during the breeding-season,—
or when the males acquire at this season such long wing or tail-feathers as
to impede their flight, as with Cosmetornis and Vidua,—it certainly at first
appears highly probable that the second moult has been gained for the
special purpose of throwing off these ornaments. We must, however,
remember that many birds, such as Birds of Paradise, the Argus pheasant
and peacock, do not cast their plumes during the winter; and it can hardly
be maintained that there is something in the constitution of these birds, at
least of the Gallinaceæ, rendering a double moult impossible, for the
ptarmigan moults thrice in the year.228 Hence it must be considered as

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doubtful whether the many species which moult their ornamental plumes or
lose their bright colours during the winter, have acquired this habit on
account of the inconvenience or danger which they would otherwise have
suffered.

I conclude, therefore, that the habit of moulting twice in the year was in
most or all cases first acquired for some distinct purpose, perhaps for
gaining a warmer winter covering; and that variations in the plumage
occurring during the summer were accumulated through sexual selection,
and transmitted to the offspring at the same season of the year. Such
variations being inherited either by both sexes or by the males alone,
according to the form of inheritance which prevailed. This appears more
probable than that these species in all cases originally tended to retain their
ornamental plumage during the winter, but were saved from this through
natural selection, owing to the inconvenience or danger thus caused.
I have endeavoured in this chapter to shew that the arguments are not
trustworthy in favour of the view that weapons, bright colours, and various
ornaments, are now confined to the males owing to the conversion, by
means of natural selection, of a tendency to the equal transmission of
characters to both sexes into transmission to the male sex alone. It is also
doubtful whether the colours of many female birds are due to the
preservation, for the sake of protection, of variations which were from the
first limited in their transmission to the female sex. But it will be
convenient to defer any further discussion on this subject until I treat, in the
following chapter, on the differences in plumage between the young and
old.

CHAPTER XVI.

Birds—concluded.
The immature plumage in relation to the character of the plumage in both sexes when adult—Six
classes of cases—Sexual differences between the males of closely-allied or representative
species—The female assuming the characters of the male—Plumage of the young in relation to
the summer and winter plumage of the adults—On the increase of beauty in the Birds of the

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World—Protective colouring—Conspicuously-coloured birds—Novelty appreciated—Summary
of the four chapters on Birds.

We must now consider the transmission of characters as limited by age in
reference to sexual selection. The truth and importance of the principle of
inheritance at corresponding ages need not here be discussed, as enough has
already been said on the subject. Before giving the several rather complex
rules or classes of cases, under which all the differences in plumage
between the young and the old, as far as known to me, may be included, it
will be well to make a few preliminary remarks.
With animals of all kinds when the young differ in colour from the adults,
and the colours of the former are not, as far as we can see, of any special
service, they may generally be attributed, like various embryological
structures, to the retention by the young of the character of an early
progenitor. But this view can be maintained with confidence, only when the
young of several species closely resemble each other, and likewise resemble
other adult species belonging to the same group; for the latter are the living
proofs that such a state of things was formerly possible. Young lions and
pumas are marked with feeble stripes or rows of spots, and as many allied
species both young and old are similarly marked, no naturalist, who
believes in the gradual evolution of species, will doubt that the progenitor
of the lion and puma was a striped animal, the young having retained
vestiges of the stripes, like the kittens of black cats, which when grown up
are not in the least striped. Many species of deer, which when mature are
not spotted, are whilst young covered with white spots, as are likewise
some few species in their adult state. So again the young in the whole
family of pigs (Suidæ), and in certain rather distantly-allied animals, such
as the tapir, are marked with dark longitudinal stripes; but here we have a
character apparently derived from an extinct progenitor, and now preserved
by the young alone. In all such cases the old have had their colours changed
in the course of time, whilst the young have remained but little altered, and
this has been effected through the principle of inheritance at corresponding
ages.
This same principle applies to many birds belonging to various groups, in
which the young closely resemble each other, and differ much from their
respective adult parents. The young of almost all the Gallinaceæ, and of
some distantly-allied birds such as ostriches, are whilst covered with down

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longitudinally striped; but this character points back to a state of things so
remote that it hardly concerns us. Young crossbills (Loxia) have at first
straight beaks like those of other finches, and in their immature striated
plumage they resemble the mature redpole and female siskin, as well as the
young of the goldfinch, greenfinch, and some other allied species. The
young of many kinds of buntings (Emberiza) resemble each other, and
likewise the adult state of the common bunting, E. miliaria. In almost the
whole large group of thrushes the young have their breasts spotted—a
character which is retained by many species throughout life, but is quite lost
by others, as by the Turdus migratorius. So again with many thrushes, the
feathers on the back are mottled before they are moulted for the first time,
and this character is retained for life by certain eastern species. The young
of many species of shrikes (Lanius), of some woodpeckers, and of an Indian
pigeon (Chalcophaps indicus), are transversely striped on the under surface;
and certain allied species or genera when adult are similarly marked. In
some closely-allied and resplendent Indian cuckoos (Chrysococcyx), the
species when mature differ considerably from each other in colour, but the
young cannot be distinguished. The young of an Indian goose (Sarkidiornis
melanonotus) closely resemble in plumage an allied genus, Dendrocygna,
when mature.229 Similar facts will hereafter be given in regard to certain
herons. Young black grouse (Tetrao tetrix) resemble the young as well as
the old of certain other species, for instance the red grouse or T. scoticus.
Finally, as Mr. Blyth, who has attended closely to this subject, has well
remarked, the natural affinities of many species are best exhibited in their
immature plumage; and as the true affinities of all organic beings depend on
their descent from a common progenitor, this remark strongly confirms the
belief that the immature plumage approximately shews us the former or
ancestral condition of the species.
Although many young birds belonging to various orders thus give us a
glimpse of the plumage of their remote progenitors, yet there are many
other birds, both dull-coloured and bright-coloured, in which the young
closely resemble their parents. With such species the young of the different
species cannot resemble each other more closely than do the parents; nor
can they present striking resemblances to allied forms in their adult state.
They give us but little insight into the plumage of their progenitors,
excepting in so far that when the young and the old are coloured in the same

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general manner throughout a whole group of species, it is probable that
their progenitors were similarly coloured.
We may now consider the classes of cases or rules under which the
differences and resemblances, between the plumage of the young and the
old, of both sexes or of one sex alone, may be grouped. Rules of this kind
were first enounced by Cuvier; but with the progress of knowledge they
require some modification and amplification. This I have attempted to do,
as far as the extreme complexity of the subject permits, from information
derived from various sources; but a full essay on this subject by some
competent ornithologist is much needed. In order to ascertain to what extent
each rule prevails, I have tabulated the facts given in four great works,
namely, by Macgillivray on the birds of Britain, Audubon on those of North
America, Jerdon on those of India, and Gould on those of Australia. I may
here premise, firstly, that the several cases or rules graduate into each other;
and secondly, that when the young are said to resemble their parents, it is
not meant that they are identically alike, for their colours are almost always
rather less vivid, and the feathers are softer and often of a different shape.

RULES OR CLASSES OF CASES.

I. When the adult male is more beautiful or conspicuous than the adult
female, the young of both sexes in their first plumage closely resemble the
adult female, as with the common fowl and peacock; or, as occasionally
occurs, they resemble her much more closely than they do the adult male.
II. When the adult female is more conspicuous than the adult male, as
sometimes though rarely occurs, the young of both sexes in their first
plumage resemble the adult male.
III. When the adult male resembles the adult female, the young of both
sexes have a peculiar first plumage of their own, as with the robin.
IV. When the adult male resembles the adult female, the young of both
sexes in their first plumage resemble the adults, as with the kingfisher,
many parrots, crows, hedge-warblers.
V. When the adults of both sexes have a distinct winter and summer
plumage, whether or not the male differs from the female, the young

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resemble the adults of both sexes in their winter dress, or much more rarely
in their summer dress, or they resemble the females alone; or the young
may have an intermediate character; or again they may differ greatly from
the adults in both their seasonal plumages.
VI. In some few cases the young in their first plumage differ from each
other according to sex; the young males resembling more or less closely the
adult males, and the young females more or less closely the adult females.
Class I.—In this class, the young of both sexes resemble, more or less
closely, the adult female, whilst the adult male differs, often in the most
conspicuous manner, from the adult female. Innumerable instances in all
Orders could be given; it will suffice to call to mind the common pheasant,
duck, and house-sparrow. The cases under this class graduate into others.
Thus the two sexes when adult may differ so slightly, and the young so
slightly from the adults, that it is doubtful whether such cases ought to
come under the present, or under the third or fourth classes. So again the
young of both sexes, instead of being quite alike, may differ in a slight
degree from each other, as in our sixth class. These transitional cases,
however, are few in number, or at least are not strongly pronounced, in
comparison with those which come strictly under the present class.
The force of the present law is well shewn in those groups, in which, as a
general rule, the two sexes and the young are all alike; for when the male in
these groups does differ from the female, as with certain parrots,
kingfishers, pigeons, &c., the young of both sexes resemble the adult
female.230 We see the same fact exhibited still more clearly in certain
anomalous cases; thus the male of Heliothrix auriculata (one of the
humming-birds) differs conspicuously from the female in having a splendid
gorget and fine ear-tufts, but the female is remarkable from having a much
longer tail than that of the male; now the young of both sexes resemble
(with the exception of the breast being spotted with bronze) the adult
female in all respects including the length of her tail, so that the tail of the
male actually becomes shorter as he reaches maturity, which is a most
unusual circumstance.231 Again, the plumage of the male goosander (Mergus
merganser) is more conspicuously coloured, with the scapular and
secondary wing-feathers much longer than in the female, but differently
from what occurs, as far as I know, in any other bird, the crest of the adult

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male, though broader than that of the female, is considerably shorter, being
only a little above an inch in length; the crest of the female being two and a
half inches long. Now the young of both sexes resemble in all respects the
adult female, so that their crests are actually of greater length though
narrower than in the adult male.232
When the young and the females closely resemble each other and both
differ from the male, the most obvious conclusion is that the male alone has
been modified. Even in the anomalous cases of the Heliothrix and Mergus,
it is probable that originally both adult sexes were furnished, the one
species with a much elongated tail, and the other with a much elongated
crest, these characters having since been partially lost by the adult males
from some unexplained cause, and transmitted in their diminished state to
their male offspring alone, when arrived at the corresponding age of
maturity. The belief that in the present class the male alone has been
modified, as far as the differences between the male and the female together
with her young are concerned, is strongly supported by some remarkable
facts recorded by Mr. Blyth,233 with respect to closely-allied species which
represent each other in distinct countries. For with several of these
representative species the adult males have undergone a certain amount of
change and can be distinguished; the females and the young being
undistinguishable, and therefore absolutely unchanged. This is the case with
certain Indian chats (Thamnobia), with certain honey-suckers (Nectarinia),
shrikes (Tephrodornis), certain kingfishers (Tanysiptera), Kallij pheasants
(Gallophasis), and tree-partridges (Arboricola).
In some analogous cases, namely with birds having a distinct summer and
winter plumage, but with the two sexes nearly alike, certain closely-allied
species can easily be distinguished in their summer or nuptial plumage, yet
are undistinguishable in their winter as well as in their immature plumage.
This is the case with some of the closely-allied Indian wagtails or
Motacillæ. Mr. Swinhoe234 informs me that three species of Ardeola, a
genus of herons, which represent each other on separate continents, are
“most strikingly different” when ornamented with their summer plumes, but
are hardly, if at all, distinguishable during the winter. The young also of
these three species in their immature plumage closely resemble the adults in
their winter dress. This case is all the more interesting because with two
other species of Ardeola both sexes retain, during the winter and summer,

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nearly the same plumage as that possessed by the three first species during
the winter and in their immature state; and this plumage, which is common
to several distinct species at different ages and seasons, probably shews us
how the progenitor of the genus was coloured. In all these cases, the nuptial
plumage which we may assume was originally acquired by the adult males
during the breeding-season, and transmitted to the adults of both sexes at
the corresponding season, has been modified, whilst the winter and
immature plumages have been left unchanged.
The question naturally arises, how is it that in these latter cases the winter
plumage of both sexes, and in the former cases the plumage of the adult
females, as well as the immature plumage of the young, have not been at all
affected? The species which represent each other in distinct countries will
almost always have been exposed to somewhat different conditions, but we
can hardly attribute the modification of the plumage in the males alone to
this action, seeing that the females and the young, though similarly
exposed, have not been affected. Hardly any fact in nature shews us more
clearly how subordinate in importance is the direct action of the conditions
of life, in comparison with the accumulation through selection of indefinite
variations, than the surprising difference between the sexes of many birds;
for both sexes must have consumed the same food and have been exposed
to the same climate. Nevertheless we are not precluded from believing that
in the course of time new conditions may produce some direct effect; we
see only that this is subordinate in importance to the accumulated results of
selection. When, however, a species migrates into a new country, and this
must precede the formation of representative species, the changed
conditions to which they will almost always have been exposed will cause
them to undergo, judging from a widely-spread analogy, a certain amount of
fluctuating variability. In this case sexual selection, which depends on an
element eminently liable to change—namely the taste or admiration of the
female—will have had new shades of colour or other differences to act on
and accumulate; and as sexual selection is always at work, it would
(judging from what we know of the results on domestic animals of man’s
unintentional selection), be a surprising fact if animals inhabiting separate
districts, which can never cross and thus blend their newly-acquired
characters, were not, after a sufficient lapse of time, differently modified.

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These remarks likewise apply to the nuptial or summer plumage, whether
confined to the males or common to both sexes.
Although the females of the above closely-allied species, together with their
young, differ hardly at all from each other, so that the males alone can be
distinguished, yet in most cases the females of the species within the same
genus obviously differ from each other. The differences, however, are rarely
as great as between the males. We see this clearly in the whole family of the
Gallinaceæ: the females, for instance, of the common and Japan pheasant,
and especially of the gold and Amherst pheasant—of the silver pheasant
and the wild fowl—resemble each other very closely in colour, whilst the
males differ to an extraordinary degree. So it is with the females of most of
the Cotingidæ, Fringillidæ, and many other families. There can indeed be
no doubt that, as a general rule, the females have been modified to a less
extent than the males. Some few birds, however, offer a singular and
inexplicable exception; thus the females of Paradisea apoda and P.
papuana differ from each other more than do their respective males;235 the
female of the latter species having the under surface pure white, whilst the
female P. apoda is deep brown beneath. So, again, as I hear from Professor
Newton, the males of two species of Oxynotus (shrikes), which represent
each other in the islands of Mauritius and Bourbon,236 differ but little in
colour, whilst the females differ much. In the Bourbon species the female
appears to have partially retained an immature condition of plumage, for at
first sight she “might be taken for the young of the Mauritian species.”
These differences may be compared with those which occur, independently
of selection by man, and which we cannot explain, in certain sub-breeds of
the game-fowl, in which the females are very different, whilst the males can
hardly be distinguished.237
As I account so largely by sexual selection for the differences between the
males of allied species, how can the differences between the females be
accounted for in all ordinary cases? We need not here consider the species
which belong to distinct genera; for with these, adaptation to different
habits of life, and other agencies, will have come into play. In regard to the
differences between the females within the same genus, it appears to me
almost certain, after looking through various large groups, that the chief
agent has been the transference, in a greater or less degree, to the female of
the characters acquired by the males through sexual selection. In the several

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British finches, the two sexes differ either very slightly or considerably; and
if we compare the females of the greenfinch, chaffinch, goldfinch,
bullfinch, crossbill, sparrow, &c., we shall see that they differ from each
other chiefly in the points in which they partially resemble their respective
males; and the colours of the males may safely be attributed to sexual
selection. With many gallinaceous species the sexes differ to an extreme
degree, as with the peacock, pheasant, and fowl, whilst with other species
there has been a partial or even complete transference of character from the
male to the female. The females of the several species of Polyplectron
exhibit in a dim condition, and chiefly on the tail, the splendid ocelli of
their males. The female partridge differs from the male only in the red mark
on her breast being smaller; and the female wild turkey only in her colours
being much duller. In the guinea-fowl the two sexes are undistinguishable.
There is no improbability in the plain, though peculiar spotted plumage of
this latter bird having been acquired through sexual selection by the males,
and then transmitted to both sexes; for it is not essentially different from the
much more beautifully-spotted plumage, characteristic of the males alone of
the Tragopan pheasants.
It should be observed that, in some instances, the transference of characters
from the male to the female has been effected apparently at a remote period,
the male having subsequently undergone great changes, without transferring
to the female any of his later-gained characters. For instance, the female and
the young of the black-grouse (Tetrao tetrix) resemble pretty closely both
sexes and the young of the red-grouse T. Scoticus; and we may
consequently infer that the black-grouse is descended from some ancient
species, of which both sexes were coloured in nearly the same manner as
the red-grouse. As both sexes of this latter species are more plainly barred
during the breeding-season than at any other time, and as the male differs
slightly from the female in his more strongly-pronounced red and brown
tints,238 we may conclude that his plumage has been, at least to a certain
extent, influenced by sexual selection. If so, we may further infer that the
nearly similar plumage of the female black-grouse was similarly produced
at some former period. But since this period the male black-grouse has
acquired his fine black plumage, with his forked and outwardly-curled tail-
feathers; but of these characters there has hardly been any transference to
the female, excepting that she shews in her tail a trace of the curved fork.

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We may therefore conclude that the females of distinct though allied species
have often had their plumage rendered more or less different by the
transference in various degrees, of characters acquired, both during former
and recent times, by the males through sexual selection. But it deserves
especial attention that brilliant colours have been transferred much more
rarely than other tints. For instance, the male of the red-throated bluebreast
(Cyanecula suecica) has a rich blue breast, including a sub-triangular red
mark; now marks of approximately the same shape have been transferred to
the female, but the central space is fulvous instead of red, and is surrounded
by mottled instead of blue feathers. The Gallinaceæ offer many analogous
cases; for none of the species, such as partridges, quails, guinea-fowls, &c.,
in which the colours of the plumage have been largely transferred from the
male to the female, are brilliantly coloured. This is well exemplified with
the pheasants, in which the male is generally so much more brilliant than
the female; but with the Eared and Cheer pheasants (Crossoptilon auritum
and Phasianus Wallichii) the two sexes closely resemble each other and
their colours are dull. We may go so far as to believe that if any part of the
plumage in the males of these two pheasants had been brilliantly coloured,
this would not have been transferred to the females. These facts strongly
support Mr. Wallace’s view that with birds which are exposed to much
danger during nidification, the transference of bright colours from the male
to the female has been checked through natural selection. We must not,
however, forget that another explanation, before given, is possible; namely,
that the males which varied and became bright, whilst they were young and
inexperienced, would have been exposed to much danger, and would
generally have been destroyed; the older and more cautious males, on the
other hand, if they varied in a like manner, would not only have been able to
survive, but would have been favoured in their rivalry with other males.
Now variations occurring late in life tend to be transmitted exclusively to
the same sex, so that in this case extremely bright tints would not have been
transmitted to the females. On the other hand, ornaments of a less
conspicuous kind, such as those possessed by the Eared and Cheer
pheasants, would not have been dangerous, and if they appeared during
early youth, would generally have been transmitted to both sexes.
In addition to the effects of the partial transference of characters from the
males to the females, some of the differences between the females of

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closely-allied species may be attributed to the direct or definite action of the
conditions of life.239 With the males any such action would generally have
been masked by the brilliant colours gained through sexual selection; but
not so with the females. Each of the endless diversities in plumage, which
we see in our domesticated birds is, of course, the result of some definite
cause; and under natural and more uniform conditions, some one tint,
assuming that it was in no way injurious, would almost certainly sooner or
later prevail. The free intercrossing of the many individuals belonging to the
same species would ultimately tend to make any change of colour, thus
induced, uniform in character.
No one doubts that both sexes of many birds have had their colours adapted
for the sake of protection; and it is possible that the females alone of some
species may have been thus modified. Although it would be a difficult,
perhaps an impossible process, as shewn in the last chapter, to convert
through selection one form of transmission into another, there would not be
the least difficulty in adapting the colours of the female, independently of
those of the male, to surrounding objects, through the accumulation of
variations which were from the first limited in their transmission to the
female sex. If the variations were not thus limited, the bright tints of the
male would be deteriorated or destroyed. Whether the females alone of
many species have been thus specially modified, is at present very doubtful.
I wish I could follow Mr. Wallace to the full extent; for the admission would
remove some difficulties. Any variations which were of no service to the
female as a protection would be at once obliterated, instead of being lost
simply by not being selected, or from free intercrossing, or from being
eliminated when transferred to the male and in any way injurious to him.
Thus the plumage of the female would be kept constant in character. It
would also be a relief if we could admit that the obscure tints of both sexes
of many birds had been acquired and preserved for the sake of protection,—
for example, of the hedge-warbler or kitty-wren (Accentor modularis and
Troglodytes vulgaris), with respect to which we have no sufficient evidence
of the action of sexual selection. We ought, however, to be cautious in
concluding that colours which appear to us dull, are not attractive to the
females of certain species; we should bear in mind such cases as that of the
common house-sparrow, in which the male differs much from the female,
but does not exhibit any bright tints. No one probably will dispute that

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many gallinaceous birds which live on the open ground have acquired their
present colours, at least in part, for the sake of protection. We know how
well they are thus concealed; we know that ptarmigans, whilst changing
from their winter to their summer plumage, both of which are protective,
suffer greatly from birds of prey. But can we believe that the very slight
differences in tints and markings between, for instance, the female black
and red-grouse serve as a protection? Are partridges, as they are now
coloured, better protected than if they had resembled quails? Do the slight
differences between the females of the common pheasant, the Japan and
golden pheasants, serve as a protection, or might not their plumages have
been interchanged with impunity? From what Mr. Wallace has observed of
the habits of certain gallinaceous birds in the East he thinks that such slight
differences are beneficial. For myself, I will only say that I am not
convinced.
Formerly when I was inclined to lay much stress on the principle of
protection, as accounting for the less bright colours of female birds, it
occurred to me that possibly both sexes and the young might aboriginally
have been brightly coloured in an equal degree; but that subsequently, the
females from the danger incurred during incubation, and the young from
being inexperienced, had been rendered dull as a protection. But this view
is not supported by any evidence, and is not probable; for we thus in
imagination expose during past times the females and the young to danger,
from which it has subsequently been necessary to shield their modified
descendants. We have, also, to reduce, through a gradual process of
selection, the females and the young to almost exactly the same tints and
markings, and to transmit them to the corresponding sex and period of life.
It is also a somewhat strange fact, on the supposition that the females and
the young have partaken during each stage of the process of modification of
a tendency to be as brightly coloured as the males, that the females have
never been rendered dull-coloured without the young participating in the
same change; for there are no instances, as far as I can discover, of species
with the females dull-coloured and the young bright-coloured. A partial
exception, however, is offered by the young of certain woodpeckers, for
they have “the whole upper part of the head tinged with red,” which
afterwards either decreases into a mere circular red line in the adults of both
sexes, or quite disappears in the adult females.240

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Finally, with respect to our present class of cases, the most probable view
appears to be that successive variations in brightness or in other ornamental
characters, occurring in the males at a rather late period of life have alone
been preserved; and that most or all of these variations owing to the late
period of life at which they appeared, have been from the first transmitted
only to the adult male offspring. Any variations in brightness which
occurred in the females or in the young would have been of no service to
them, and would not have been selected; moreover, if dangerous, would
have been eliminated. Thus the females and the young will either have been
left unmodified, or, and this has much more commonly occurred, will have
been partially modified by receiving through transference from the males
some of the successive variations. Both sexes have perhaps been directly
acted on by the conditions of life to which they have long been exposed; but
the females from not being otherwise much modified will best exhibit any
such effects. These changes and all others will have been kept uniform by
the free intercrossing of many individuals. In some cases, especially with
ground birds, the females and the young may possibly have been modified,
independently of the males, for the sake of protection, so as to have
acquired the same dull-coloured plumage.
Class II. When the adult female is more conspicuous than the adult male,
the young of both sexes in their first plumage resemble the adult male.—
This class is exactly the reverse of the last, for the females are here more
brightly coloured or more conspicuous than the males; and the young, as far
as they are known, resemble the adult males instead of the adult females.
But the difference between the sexes is never nearly so great as occurs with
many birds in the first class, and the cases are comparatively rare. Mr.
Wallace who first called attention to the singular relation which exists
between the less bright colours of the males and their performing the duties
of incubation, lays great stress on this point,241 as a crucial test that obscure
colours have been acquired for the sake of protection during the period of
nesting. A different view seems to me more probable. As the cases are
curious and not numerous, I will briefly give all that I have been able to
find.
In one section of the genus Turnix, quail-like birds, the female is invariably
larger than the male (being nearly twice as large in one of the Australian
species) and this is an unusual circumstance with the Gallinaceæ. In most of

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the species the female is more distinctly coloured and brighter than the
male,242 but in some few species the sexes are alike. In Turnix taigoor of
India the male “wants the black on the throat and neck, and the whole tone
of the plumage is lighter and less pronounced than that of the female.” The
female appears to be more vociferous, and is certainly much more
pugnacious than the male; so that the females and not the males are often
kept by the natives for fighting, like game-cocks. As male birds are exposed
by the English bird-catchers for a decoy near a trap, in order to catch other
males by exciting their rivalry, so the females of this Turnix are employed
in India. When thus exposed the females soon begin their “loud purring call,
which can be heard a long way off, and any females within ear-shot run
rapidly to the spot, and commence fighting with the caged bird.” In this way
from twelve to twenty birds, all breeding-females, may be caught in the
course of a single day. The natives assert that the females after laying their
eggs associate in flocks, and leave the males to sit on them. There is no
reason to doubt the truth of this assertion, which is supported by some
observations made in China by Mr. Swinhoe.243 Mr. Blyth believes, that the
young of both sexes resemble the adult male.

Fig. 60. Rhynchæa capensis (from Brehm).

The females of the three species of Painted Snipes (Rhynchæa) “are not
only larger, but much more richly coloured than the males.”244 With all other
birds, in which the trachea differs in structure in the two sexes it is more
developed and complex in the male than in the female; but in the Rhynchæa
Australis it is simple in the male, whilst in the female it makes four distinct

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convolutions before entering the lungs.245 The female therefore of this
species has acquired an eminently masculine character. Mr. Blyth
ascertained, by examining many specimens, that the trachea is not
convoluted in either sex of R. Bengalensis, which species so closely
resembles R. Australis that it can hardly be distinguished except by its
shorter toes. This fact is another striking instance of the law that secondary
sexual characters are often widely different in closely-allied forms; though
it is a very rare circumstance when such differences relate to the female sex.
The young of both sexes of R. Bengalensis in their first plumage are said to
resemble the mature male.246 There is also reason to believe that the male
undertakes the duty of incubation, for Mr. Swinhoe247 found the females
before the close of the summer associated in flocks, as occurs with the
females of the Turnix.
The females of Phalaropus fulicarius and P. hyperboreus are larger, and in
their summer plumage “more gaily attired than the males.” But the
difference in colour between the sexes is far from conspicuous. The male
alone of P. fulicarius undertakes, according to Professor Steenstrup, the
duty of incubation, as is likewise shewn by the state of his breast-feathers
during the breeding-season. The female of the dotterel plover (Eudromias
morinellus) is larger than the male, and has the red and black tints on the
lower surface, the white crescent on the breast, and the stripes over the
eyes, more strongly pronounced. The male also takes at least a share in
hatching the eggs; but the female likewise attends to the young.248 I have not
been able to discover whether with these species the young resemble the
adult males more closely than the adult females; for the comparison is
somewhat difficult to make on account of the double moult.
Turning now to the Ostrich order: the male of the common cassowary
(Casuarius galeatus) would be thought by any one to be the female, from
his smaller size and from the appendages and naked skin about his head
being much less brightly coloured; and I am informed by Mr. Bartlett that in
the Zoological Gardens it is certainly the male alone who sits on the eggs
and takes care of the young.249 The female is said by Mr. T. W. Wood250 to
exhibit during the breeding-season a most pugnacious disposition; and her
wattles then become enlarged and more brilliantly coloured. So again the
female of one of the emus (Dromœus irroratus) is considerably larger than
the male, and she possesses a slight top-knot, but is otherwise

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undistinguishable in plumage. She appears, however, “to have greater
power, when angry or otherwise excited, of erecting, like a turkey-cock, the
feathers of her neck and breast. She is usually the more courageous and
pugilistic. She makes a deep hollow guttural boom, especially at night,
sounding like a small gong. The male has a slenderer frame and is more
docile, with no voice beyond a suppressed hiss when angry, or a croak.” He
not only performs the whole duty of incubation, but has to defend the young
from their mother; “for as soon as she catches sight of her progeny she
becomes violently agitated, and notwithstanding the resistance of the father
appears to use her utmost endeavours to destroy them. For months
afterwards it is unsafe to put the parents together, violent quarrels being the
inevitable result, in which the female generally comes off conqueror.”251 So
that with this emu we have a complete reversal not only of the parental and
incubating instincts, but of the usual moral qualities of the two sexes; the
females being savage, quarrelsome and noisy, the males gentle and good.
The case is very different with the African ostrich, for the male is somewhat
larger than the female and has finer plumes with more strongly contrasted
colours; nevertheless he undertakes the whole duty of incubation.252
I will specify the few other cases known to me, in which the female is more
conspicuously coloured than the male, although nothing is known about
their manner of incubation. With the carrion-hawk of the Falkland Islands
(Milvago leucurus) I was much surprised to find by dissection that the
individuals, which had all their tints strongly pronounced, with the cere and
legs orange-coloured, were the adult females; whilst those with duller
plumage and grey legs were the males or the young. In an Australian tree-
creeper (Climacteris erythrops) the female differs from the male in “being
adorned with beautiful, radiated, rufous markings on the throat, the male
having this part quite plain.” Lastly in an Australian nightjar “the female
always exceeds the male in size and in the brilliance of her tints; the males,
on the other hand, have two white spots on the primaries more conspicuous
than in the female.”253
We thus see that the cases in which female birds are more conspicuously
coloured than the males, with the young in their immature plumage
resembling the adult males instead of the adult females, as in the previous
class, are not numerous, though they are distributed in various Orders. The
amount of difference, also, between the sexes is incomparably less than that

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which frequently occurs in the last class; so that the cause of the difference,
whatever it may have been, has acted on the females in the present class
either less energetically or less persistently than on the males in the last
class. Mr. Wallace believes that the males have had their colours rendered
less conspicuous for the sake of protection during the period of incubation;
but the difference between the sexes in hardly any of the foregoing cases
appears sufficiently great for this view to be safely accepted. In some of the
cases the brighter tints of the female are almost confined to the lower
surface, and the males, if thus coloured, would not have been exposed to
danger whilst sitting on the eggs. It should also be borne in mind that the
males are not only in a slight degree less conspicuously coloured than the
females, but are of less size, and have less strength. They have, moreover,
not only acquired the maternal instinct of incubation, but are less
pugnacious and vociferous than the females, and in one instance have
simpler vocal organs. Thus an almost complete transposition of the
instincts, habits, disposition, colour, size, and of some points of structure,
has been effected between the two sexes.
Now if we might assume that the males in the present class have lost some
of that ardour which is usual to their sex, so that they no longer search
eagerly for the females; or, if we might assume that the females have
become much more numerous than the males—and in the case of one
Indian Turnix the females are said to be “much more commonly met with
than the males”254—then it is not improbable that the females would have
been led to court the males, instead of being courted by them. This indeed is
the case to a certain extent, with some birds, as we have seen with the
peahen, wild turkey, and certain kinds of grouse. Taking as our guide the
habits of most male birds, the greater size and strength and the
extraordinary pugnacity of the females of the Turnix and Emu, must mean
that they endeavour to drive away rival females, in order to gain possession
of the male; and on this view, all the facts become clear; for the males
would probably be most charmed or excited by the females which were the
most attractive to them by their brighter colours, other ornaments, or vocal
powers. Sexual selection would then soon do its work, steadily adding to
the attractions of the females; the males and the young being left not at all,
or but little modified.

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Class III. When the adult male resembles the adult female, the young of
both sexes have a peculiar first plumage of their own.—In this class both
sexes when adult resemble each other, and differ from the young. This
occurs with many birds of many kinds. The male robin can hardly be
distinguished from the female, but the young are widely different with their
mottled dusky-olive and brown plumage. The male and female of the
splendid scarlet Ibis are alike, whilst the young are brown; and the scarlet-
colour, though common to both sexes, is apparently a sexual character, for it
is not well developed with birds under confinement, in the same manner as
often occurs in the case of brilliantly coloured male birds. With many
species of herons the young differ greatly from the adults, and their summer
plumage, though common to both sexes, clearly has a nuptial character.
Young swans are slate-coloured, whilst the mature birds are pure white; but
it would be superfluous to give additional instances. These differences
between the young and the old apparently depend, as in the two last classes,
on the young having retained a former or ancient state of plumage, which
has been exchanged for a new plumage by the old of both sexes. When the
adults are brightly coloured, we may conclude from the remarks just made
in relation to the scarlet ibis and to many herons, and from the analogy of
the species in the first class, that such colours have been acquired through
sexual selection by the nearly mature males; but that, differently from what
occurs in the two first classes, the transmission, though limited to the same
age, has not been limited to the same sex. Consequently both sexes when
mature resemble each other and differ from the young.
Class IV. When the adult male resembles the adult female, the young of
both sexes in their first plumage resemble the adults.—In this class the
young and the adults of both sexes, whether brilliantly or obscurely
coloured, resemble each other. Such cases are, I think, more common than
those in the last class. We have in England instances in the kingfisher, some
woodpeckers, the jay, magpie, crow, and many small dull-coloured birds,
such as the hedge-warbler or kitty-wren. But the similarity in plumage
between the young and the old is never absolutely complete, and graduates
away into dissimilarity. Thus the young of some members of the kingfisher
family are not only less vividly coloured than the adults, but many of the
feathers on the lower surface are edged with brown,255—a vestige probably
of a former state of the plumage. Frequently in the same group of birds,

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even within the same genus, for instance in an Australian genus of
parrakeets (Platycercus), the young of some species closely resemble,
whilst the young of other species differ considerably from their parents of
both sexes, which are alike.256 Both sexes and the young of the common jay
are closely similar; but in the Canada jay (Perisoreus canadensis) the young
differ so much from their parents that they were formerly described as
distinct species.257
Before proceeding, I may remark that under the present and two next
classes of cases the facts are so complex, and the conclusions so doubtful,
that any one who feels no especial interest in the subject had better pass
them over.
The brilliant or conspicuous colours which characterise many birds in the
present class, can rarely or never be of service to them as a protection; so
that they have probably been gained by the males through sexual selection,
and then transferred to the females and the young. It is, however, possible
that the males may have selected the more attractive females; and if these
transmitted their characters to their offspring of both sexes, the same results
would follow as from the selection of the more attractive males by the
females. But there is some evidence that this contingency has rarely, if ever,
occurred in any of those groups of birds, in which the sexes are generally
alike; for if even a few of the successive variations had failed to be
transmitted to both sexes, the females would have exceeded to a slight
degree the males in beauty. Exactly the reverse occurs under nature; for in
almost every large group, in which the sexes generally resemble each other,
the males of some few species are in a slight degree more brightly coloured
than the females. It is again possible that the females may have selected the
more beautiful males, these males having reciprocally selected the more
beautiful females; but it is doubtful whether this double process of selection
would be likely to occur, owing to the greater eagerness of one sex than the
other, and whether it would be more efficient than selection on one side
alone. It is, therefore, the most probable view that sexual selection has
acted, in the present class, as far as ornamental characters are concerned, in
accordance with the general rule throughout the animal kingdom, that is, on
the males; and that these have transmitted their gradually-acquired colours,
either equally or almost equally, to their offspring of both sexes.

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Another point is more doubtful, namely, whether the successive variations
first appeared in the males after they had become nearly mature, or whilst
quite young. In either case sexual selection must have acted on the male
when he had to compete with rivals for the possession of the female; and in
both cases the characters thus acquired have been transmitted to both sexes
and all ages. But these characters, if acquired by the males when adult, may
have been transmitted at first to the adults alone, and at some subsequent
period transferred to the young. For it is known that when the law of
inheritance at corresponding ages fails, the offspring often inherit characters
at an earlier age than that at which they first appeared in their parents.258
Cases apparently of this kind have been observed with birds in a state of
nature. For instance Mr. Blyth has seen specimens of Lanius rufus and of
Colymbus glacialis which had assumed whilst young, in a quite anomalous
manner, the adult plumage of their parents.259 Again, the young of the
common swan (Cygnus olor) do not cast off their dark feathers and become
white until eighteen months or two years old; but Dr. F. Forel has described
the case of three vigorous young birds, out of a brood of four, which were
born pure white. These young birds were not albinoes, as shewn by the
colour of their beaks and legs, which nearly resembled the same parts in the
adults.260
It may be worth while to illustrate the above three modes by which, in the
present class, the two sexes and the young may have come to resemble each
other, by the curious case of the genus Passer.261 In the house-sparrow (P.
domesticus) the male differs much from the female and from the young.
These resemble each other, and likewise to a large extent both sexes and the
young of the sparrow of Palestine (P. brachydactylus), as well as of some
allied species. We may therefore assume that the female and young of the
house-sparrow approximately shew us the plumage of the progenitor of the
genus. Now with the tree-sparrow (P. montanus) both sexes and the young
closely resemble the male of the house-sparrow; so that they have all been
modified in the same manner, and all depart from the typical colouring of
their early progenitor. This may have been effected by a male ancestor of
the tree-sparrow having varied, firstly, when nearly mature, or, secondly,
whilst quite young, having in either case transmitted his modified plumage
to the females and the young; or, thirdly, he may have varied when adult
and transmitted his plumage to both adult sexes, and, owing to the failure of

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the law of inheritance at corresponding ages, at some subsequent period to
his young.
It is impossible to decide which of these three modes has generally
prevailed throughout the present class of cases. The belief that the males
varied whilst young, and transmitted their variations to their offspring of
both sexes is perhaps the most probable. I may here add that I have
endeavoured, with little success, by consulting various works, to decide
how far with birds the period of variation has generally determined the
transmission of characters to one sex or to both. The two rules, often
referred to (namely, that variations occurring late in life are transmitted to
one and the same sex, whilst those which occur early in life are transmitted
to both sexes), apparently hold good in the first,262 second, and fourth
classes of cases; but they fail in an equal number, namely, in the third, often
in the fifth,263 and in the sixth small class. They hold good, however, as far
as I can judge, with a considerable majority of the species of birds. Whether
or not this be so, we may conclude from the facts given in the eighth
chapter that the period of variation has been one important element in
determining the form of transmission.
With birds it is difficult to decide by what standard we ought to judge of the
earliness or lateness of the period of variation, whether by the age in
reference to the duration of life, or to the power of reproduction, or to the
number of moults through which the species passes. The moulting of birds,
even within the same family, sometimes differs much without any
assignable cause. Some birds moult so early, that nearly all the body-
feathers are cast off before the first wing-feathers are fully grown; and we
cannot believe that this was the primordial state of things. When the period
of moulting has been accelerated, the age at which the colours of the adult
plumage were first developed would falsely appear to us to have been
earlier than it really was. This may be illustrated by the practice followed by
some bird-fanciers, who pull out a few feathers from the breast of nestling
bullfinches, and from the head or neck of young gold pheasants, in order to
ascertain their sex; for in the males these feathers are immediately replaced
by coloured ones.264 The actual duration of life is known in but few birds, so
that we can hardly judge by this standard. And with reference to the period
at which the powers of reproduction are gained, it is a remarkable fact that
various birds occasionally breed whilst retaining their immature plumage.265

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The fact of birds breeding in their immature plumage seems opposed to the
belief that sexual selection has played as important a part, as I believe it
has, in giving ornamental colours, plumes, &c., to the males, and, by means
of equal transmission, to the females of many species. The objection would
be a valid one, if the younger and less ornamented males were as successful
in winning females and propagating their kind, as the older and more
beautiful males. But we have no reason to suppose that this is the case.
Audubon speaks of the breeding of the immature males of Ibis tantalus as a
rare event, as does Mr. Swinhoe, in regard to the immature males of
Oriolus.266 If the young of any species in their immature plumage were
more successful in winning partners than the adults, the adult plumage
would probably soon be lost, as the males which retained their immature
dress for the longest period would prevail, and thus the character of the
species would ultimately be modified.267 If, on the other hand, the young
never succeeded in obtaining a female, the habit of early reproduction
would perhaps be sooner or later quite eliminated, from being superfluous
and entailing waste of power.
The plumage of certain birds goes on increasing in beauty during many
years after they are fully mature; this is the case with the train of the
peacock, and with the crest and plumes of certain herons; for instance, the
Ardea Ludovicana;268 but it is very doubtful whether the continued
development of such feathers is the result of the selection of successive
beneficial variations, or merely of continuous growth. Most fishes continue
increasing in size, as long as they are in good health and have plenty of
food; and a somewhat similar law may prevail with the plumes of birds.
Class V. When the adults of both sexes have a distinct winter and summer
plumage, whether or not the male differs from the female, the young
resemble the adults of both sexes in their winter dress, or much more rarely
in their summer dress, or they resemble the females alone; or the young
may have an intermediate character; or again, they may differ greatly from
the adults in both their seasonal plumages.—The cases in this class are
singularly complex; nor is this surprising, as they depend on inheritance,
limited in a greater or less degree in three different ways, namely by sex,
age, and the season of the year. In some cases the individuals of the same
species pass through at least five distinct states of plumage. With the
species, in which the male differs from the female during the summer

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season alone, or, which is rarer, during both seasons,269 the young generally
resemble the females,—as with the so-called goldfinch of North America,
and apparently with the splendid Maluri of Australia.270 With the species,
the sexes of which are alike during both the summer and winter, the young
may resemble the adults, firstly, in their winter dress; secondly, which
occurs much more rarely, in their summer dress; thirdly, they may be
intermediate between these two states; and, fourthly, they may differ greatly
from the adults at all seasons. We have an instance of the first of these four
cases in one of the egrets of India (Buphus coromandus), in which the
young and the adults of both sexes are white during the winter, the adults
becoming golden-buff during the summer. With the Gaper (Anastomus
oscitans) of India we have a similar case, but the colours are reversed; for
the young and the adults of both sexes are grey and black during the winter,
the adults becoming white during the summer.271 As an instance of the
second case, the young of the razor-bill (Alca torda, Linn.), in an early state
of plumage, are coloured like the adults during the summer; and the young
of the white-crowned sparrow of North America (Fringilla leucophrys), as
soon as fledged, have elegant white stripes on their heads, which are lost by
the young and the old during the winter.272 With respect to the third case,
namely, that of the young having an intermediate character between the
summer and winter adult plumages, Yarrell273 insists that this occurs with
many waders. Lastly, in regard to the young differing greatly from both
sexes in their adult summer and winter plumages, this occurs with some
herons and egrets of North America and India,—the young alone being
white.
I will make only a few remarks on these complicated cases. When the
young resemble the female in her summer dress, or the adults of both sexes
in their winter dress, the cases differ from those given under Classes I. and
III. only in the characters originally acquired by the males during the
breeding-season, having been limited in their transmission to the
corresponding season. When the adults have a distinct summer and winter
plumage, and the young differ from both, the case is more difficult to
understand. We may admit as probable that the young have retained an
ancient state of plumage; we can account through sexual selection for the
summer or nuptial plumage of the adults, but how are we to account for
their distinct winter plumage? If we could admit that this plumage serves in

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all cases as a protection, its acquirement would be a simple affair; but there
seems no good reason for this admission. It may be suggested that the
widely different conditions of life during the winter and summer have acted
in a direct manner on the plumage; this may have had some effect, but I
have not much confidence in so great a difference, as we sometimes see,
between the two plumages having been thus caused. A more probable
explanation is, that an ancient style of plumage, partially modified through
the transference of some characters from the summer plumage, has been
retained by the adults during the winter. Finally, all the cases in our present
class apparently depend on characters acquired by the adult males, having
been variously limited in their transmission according to age, season, and
sex; but it would not be worth while to attempt to follow out these complex
relations.
Class VI. The young in their first plumage differ from each other according
to sex; the young males resembling more or less closely the adult males,
and the young females more or less closely the adult females.—The cases in
the present class, though occurring in various groups, are not numerous;
yet, if experience had not taught us to the contrary, it seems the most natural
thing that the young should at first always resemble to a certain extent, and
gradually become more and more like, the adults of the same sex. The adult
male blackcap (Sylvia atricapilla) has a black head, that of the female being
reddish-brown; and I am informed by Mr. Blyth, that the young of both
sexes can be distinguished by this character even as nestlings. In the family
of thrushes an unusual number of similar cases have been noticed; the male
blackbird (Turdus merula) can be distinguished in the nest from the female,
as the main wing-feathers, which are not moulted so soon as the body-
feathers, retain a brownish tint until the second general moult.274 The two
sexes of the mocking bird (Turdus polyglottus, Linn.) differ very little from
each other, yet the males can easily be distinguished at a very early age
from the females by shewing more pure white.275 The males of a forest-
thrush and of a rock-thrush (viz. Orocetes erythrogastra and Petrocincla
cyanea) have much of their plumage of a fine blue, whilst the females are
brown; and the nestling males of both species have their main wing and tail-
feathers edged with blue, whilst those of the female are edged with
brown.276 So that the very same feathers which in the young blackbird
assume their mature character and become black after the others, in these

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two species assume this character and become blue before the others. The
most probable view with reference to these cases is that the males,
differently from what occurs in Class I., have transmitted their colours to
their male offspring at an earlier age than that at which they themselves first
acquired them; for if they had varied whilst quite young, they would
probably have transmitted all their characters to their offspring of both
sexes.277
In Aïthurus polytmus (one of the humming-birds) the male is splendidly
coloured black and green, and two of the tail-feathers are immensely
lengthened; the female has an ordinary tail and inconspicuous colours; now
the young males, instead of resembling the adult female, in accordance with
the common rule, begin from the first to assume the colours proper to their
sex, and their tail-feathers soon become elongated. I owe this information to
Mr. Gould, who has given me the following more striking and as yet
unpublished case. Two humming-birds belonging to the genus Eustephanus,
both beautifully coloured, inhabit the small island of Juan Fernandez, and
have always been ranked as specifically distinct. But it has lately been
ascertained that the one, which is of a rich chesnut-brown colour with a
golden-red head, is the male, whilst the other, which is elegantly variegated
with green and white with a metallic-green head, is the female. Now the
young from the first resemble to a certain extent the adults of the
corresponding sex, the resemblance gradually becoming more and more
complete.
In considering this last case, if as before we take the plumage of the young
as our guide, it would appear that both sexes have been independently
rendered beautiful; and not that the one sex has partially transferred its
beauty to the other. The male apparently has acquired his bright colours
through sexual selection in the same manner as, for instance, the peacock or
pheasant in our first class of cases; and the female in the same manner as
the female Rhynchæa or Turnix in our second class of cases. But there is
much difficulty in understanding how this could have been effected at the
same time with the two sexes of the same species. Mr. Salvin states, as we
have seen in the eighth chapter, that with certain humming-birds the males
greatly exceed in number the females, whilst with other species inhabiting
the same country the females greatly exceed the males. If, then, we might
assume that during some former lengthened period the males of the Juan

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Fernandez species had greatly exceeded the females in number, but that
during another lengthened period the females had greatly exceeded the
males, we could understand how the males at one time, and the females at
another time, might have been rendered beautiful by the selection of the
brighter-coloured individuals of either sex; both sexes transmitting their
characters to their young at a rather earlier age than usual. Whether this is
the true explanation I will not pretend to say; but the case is too remarkable
to be passed over without notice.

We have now seen in numerous instances under all six classes, that an
intimate relation exists between the plumage of the young and that of the
adults, either of one sex or both sexes. These relations are fairly well
explained on the principle that one sex—this being in the great majority of
cases the male—first acquired through variation and sexual selection bright
colours or other ornaments, and transmitted them in various ways, in
accordance with the recognised laws of inheritance. Why variations have
occurred at different periods of life, even sometimes with the species of the
same group, we do not know; but with respect to the form of transmission,
one important determining cause seems to have been the age at which the
variations first appeared.
From the principle of inheritance at corresponding ages, and from any
variations in colour which occurred in the males at an early age not being
then selected, on the contrary being often eliminated as dangerous, whilst
similar variations occurring at or near the period of reproduction have been
preserved, it follows that the plumage of the young will often have been left
unmodified, or but little modified. We thus get some insight into the
colouring of the progenitors of our existing species. In a vast number of
species in five out of our six classes of cases, the adults of one sex or both
are brightly coloured, at least during the breeding-season, whilst the young
are invariably less brightly coloured than the adults, or are quite dull-
coloured; for no instance is known, as far as I can discover, of the young of
dull-coloured species displaying bright colours, or of the young of brightly-
coloured species being more brilliantly coloured than their parents. In the
fourth class, however, in which the young and the old resemble each other,
there are many species (though by no means all) brightly-coloured, and as
these form whole groups, we may infer that their early progenitors were

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likewise brightly-coloured. With this exception, if we look to the birds of
the world, it appears that their beauty has been greatly increased since that
period, of which we have a partial record in their immature plumage.

On the Colour of the Plumage in relation to Protection.—It will have been
seen that I cannot follow Mr. Wallace in the belief that dull colours when
confined to the females have been in most cases specially gained for the
sake of protection. There can, however, be no doubt, as formerly remarked,
that both sexes of many birds have had their colours modified for this
purpose, so as to escape the notice of their enemies; or, in some instances,
so as to approach their prey unobserved, in the same manner as owls have
had their plumage rendered soft, that their flight may not be overheard. Mr.
Wallace remarks278 that “it is only in the tropics, among forests which never
lose their foliage, that we find whole groups of birds, whose chief colour is
green.” It will be admitted by every one, who has ever tried, how difficult it
is to distinguish parrots in a leaf-covered tree. Nevertheless, we must
remember that many parrots are ornamented with crimson, blue, and orange
tints, which can hardly be protective. Woodpeckers are eminently arboreal,
but, besides green species, there are many black, and black-and-white kinds
—all the species being apparently exposed to nearly the same dangers. It is
therefore probable that strongly-pronounced colours have been acquired by
tree-haunting birds through sexual selection, but that green tints have had
an advantage through natural selection over other colours for the sake of
protection.
In regard to birds which live on the ground, everyone admits that they are
coloured so as to imitate the surrounding surface. How difficult it is to see a
partridge, snipe, woodcock, certain plovers, larks, and nightjars when
crouched on the ground. Animals inhabiting deserts offer the most striking
instances, for the bare surface affords no concealment, and all the smaller
quadrupeds, reptiles, and birds depend for safety on their colours. As Mr.
Tristram has remarked,279 in regard to the inhabitants of the Sahara, all are
protected by their “isabelline or sand-colour.” Calling to my recollection the
desert-birds which I had seen in South America, as well as most of the
ground-birds in Great Britain, it appeared to me that both sexes in such
cases are generally coloured nearly alike. Accordingly I applied to Mr.
Tristram, with respect to the birds of the Sahara, and he has kindly given me

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the following information. There are twenty-six species, belonging to
fifteen genera, which manifestly have had their plumage coloured in a
protective manner; and this colouring is all the more striking, as with most
of these birds it is different from that of their congeners. Both sexes of
thirteen out of the twenty-six species are coloured in the same manner; but
these belong to genera in which this rule commonly prevails, so that they
tell us nothing about the protective colours being the same in both sexes of
desert-birds. Of the other thirteen species, three belong to genera in which
the sexes usually differ from each other, yet they have the sexes alike. In the
remaining ten species, the male differs from the female; but the difference is
confined chiefly to the under surface of the plumage, which is concealed
when the bird crouches on the ground; the head and back being of the same
sand-coloured hue in both sexes. So that in these ten species the upper
surfaces of both sexes have been acted on and rendered alike, through
natural selection, for the sake of protection; whilst the lower surfaces of the
males alone have been diversified through sexual selection, for the sake of
ornament. Here, as both sexes are equally well protected, we clearly see that
the females have not been prevented through natural selection from
inheriting the colours of their male parents: we must look to the law of
sexually limited transmission, as before explained.
In all parts of the world both sexes of many soft-billed birds, especially
those which frequent reeds or sedges, are obscurely coloured. No doubt if
their colours had been brilliant, they would have been much more
conspicuous to their enemies; but whether their dull tints have been
specially gained for the sake of protection seems, as far as I can judge,
rather doubtful. It is still more doubtful whether such dull tints can have
been gained for the sake of ornament. We must, however, bear in mind that
male birds, though dull-coloured, often differ much from their females, as
with the common sparrow, and this leads to the belief that such colours
have been gained through sexual selection, from being attractive. Many of
the soft-billed birds are songsters; and a discussion in a former chapter
should not be forgotten, in which it was shewn that the best songsters are
rarely ornamented with bright tints. It would appear that female birds, as a
general rule, have selected their mates either for their sweet voices or gay
colours, but not for both charms combined. Some species which are
manifestly coloured for the sake of protection, such as the jack-snipe,

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woodcock, and nightjar, are likewise marked and shaded, according to our
standard of taste, with extreme elegance. In such cases we may conclude
that both natural and sexual selection have acted conjointly for protection
and ornament. Whether any bird exists which does not possess some special
attraction, by which to charm the opposite sex, may be doubted. When both
sexes are so obscurely coloured, that it would be rash to assume the agency
of sexual selection, and when no direct evidence can be advanced shewing
that such colours serve as a protection, it is best to own complete ignorance
of the cause, or, which comes to nearly the same thing, to attribute the result
to the direct action of the conditions of life.
There are many birds both sexes of which are conspicuously, though not
brilliantly coloured, such as the numerous black, white, or piebald species;
and these colours, are probably the result of sexual selection. With the
common blackbird, capercailzie, black-cock, black Scoter-duck (Oidemia),
and even with one of the Birds of Paradise (Lophorina atra), the males
alone are black, whilst the females are brown or mottled; and there can
hardly be a doubt that blackness in these cases has been a sexually selected
character. Therefore it is in some degree probable that the complete or
partial blackness of both sexes in such birds as crows, certain cockatoos,
storks, and swans, and many marine birds, is likewise the result of sexual
selection, accompanied by equal transmission to both sexes; for blackness
can hardly serve in any case as a protection. With several birds, in which
the male alone is black, and in others in which both sexes are black, the
beak or skin about the head is brightly coloured, and the contrast thus
afforded adds greatly to their beauty; we see this in the bright yellow beak
of the male blackbird, in the crimson skin over the eyes of the black-cock
and capercailzie, in the variously and brightly-coloured beak of the Scoter-
drake (Oidemia), in the red beak of the chough (Corvus graculus, Linn.), of
the black swan, and black stork. This leads me to remark that it is not at all
incredible that toucans may owe the enormous size of their beaks to sexual
selection, for the sake of displaying the diversified and vivid stripes of
colour, with which these organs are ornamented.280 The naked skin at the
base of the beak and round the eyes is likewise often brilliantly coloured;
and Mr. Gould, in speaking of one species,281 says that the colours of the
beak “are doubtless in the finest and most brilliant state during the time of
pairing.” There is no greater improbability in toucans being encumbered

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with immense beaks, though rendered as light as possible by their
cancellated structure, for an object falsely appearing to us unimportant,
namely, the display of fine colours, than that the male Argus pheasant and
some other birds should be encumbered with plumes so long as to impede
their flight.
In the same manner, as the males alone of various species are black, the
females being dull-coloured; so in a few cases the males alone are either
wholly or partially white, as with the several Bell-birds of South America
(Chasmorhynchus), the Antarctic goose (Bernicla antarctica), the silver
pheasant, &c., whilst the females are brown or obscurely mottled.
Therefore, on the same principle as before, it is probable that both sexes of
many birds, such as white cockatoos, several egrets with their beautiful
plumes, certain ibises, gulls, terns, &c., have acquired their more or less
completely white plumage through sexual selection. The species which
inhabit snowy regions of course come under a different head. The white
plumage of some of the above-named birds appears in both sexes only
when they are mature. This is likewise the case with certain gannets, tropic-
birds, &c., and with the snow-goose (Anser hyperboreus). As the latter
breeds on the “barren grounds,” when not covered with snow, and as it
migrates southward during the winter, there is no reason to suppose that its
snow-white adult plumage serves as a protection. In the case of the
Anastomus oscitans previously alluded to, we have still better evidence that
the white plumage is a nuptial character, for it is developed only during the
summer; the young in their immature state, and the adults in their winter
dress, being grey and black. With many kinds of gulls (Larus), the head and
neck become pure white during the summer, being grey or mottled during
the winter and in the young state. On the other hand, with the smaller gulls,
or sea-mews (Gavia), and with some terns (Sterna), exactly the reverse
occurs; for the heads of the young birds during the first year, and of the
adults during the winter, are either pure white, or much paler-coloured than
during the breeding-season. These latter cases offer another instance of the
capricious manner in which sexual selection appears often to have acted.282
The cause of aquatic birds having acquired a white plumage so much more
frequently than terrestrial birds, probably depends on their large size and
strong powers of flight, so that they can easily defend themselves or escape
from birds of prey, to which moreover they are not much exposed.

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Consequently sexual selection has not here been interfered with or guided
for the sake of protection. No doubt, with birds which roam over the open
ocean, the males and females could find each other much more easily when
made conspicuous either by being perfectly white, or intensely black; so
that these colours may possibly serve the same end as the call-notes of
many land-birds. A white or black bird, when it discovers and flies down to
a carcase floating on the sea or cast up on the beach, will be seen from a
great distance, and will guide other birds of the same and of distinct species,
to the prey; but as this would be a disadvantage to the first finders, the
individuals which were the whitest or blackest would not thus have
procured more food than the less strongly coloured individuals. Hence
conspicuous colours cannot have been gradually acquired for this purpose
through natural selection.283
As sexual selection depends on so fluctuating an element as taste, we can
understand how it is that within the same group of birds, with habits of life
nearly the same, there should exist white or nearly white, as well as black,
or nearly black species,—for instance, white and black cockatoos, storks,
ibises, swans, terns, and petrels. Piebald birds likewise sometimes occur in
the same groups, for instance, the black-necked swan, certain terns, and the
common magpie. That a strong contrast in colour is agreeable to birds, we
may conclude, by looking through any large collection of specimens or
series of coloured plates, for the sexes frequently differ from each other in
the male having the pale parts of a purer white, and the variously coloured
dark parts of still darker tints than in the female.
It would even appear that mere novelty, or change for the sake of change,
has sometimes acted like a charm on female birds, in the same manner as
changes of fashion with us. The Duke of Argyll says,284—and I am glad to
have the unusual satisfaction of following for even a short distance in his
footsteps—“I am more and more convinced that variety, mere variety, must
be admitted to be an object and an aim in Nature.” I wish the Duke had
explained what he here means by Nature. Is it meant that the Creator of the
universe ordained diversified results for His own satisfaction, or for that of
man? The former notion seems to me as much wanting in due reverence as
the latter in probability. Capriciousness of taste in the birds themselves
appears a more fitting explanation. For example; the males of some parrots
can hardly be said to be more beautiful, at least according to our taste, than

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the females, but they differ from them in such points, as the male having a
rose-coloured collar instead of, as in the female, “a bright emeraldine
narrow green collar;” or in the male having a black collar instead of “a
yellow demi-collar in front,” with a pale roseate instead of a plum-blue
head.285 As so many male birds have for their chief ornament elongated tail-
feathers or elongated crests, the shortened tail, formerly described in the
male of a humming-bird, and the shortened crest of the male goosander
almost seem like one of the many opposite changes of fashion which we
admire in our own dresses.
Some members of the heron family offer a still more curious case of
novelty in colouring having apparently been appreciated for the sake of
novelty. The young of the Ardea asha are white, the adults being dark slate-
coloured; and not only the young, but the adults of the allied Buphus
coromandus in their winter plumage are white, this colour changing into a
rich golden-buff during the breeding-season. It is incredible that the young
of these two species, as well as of some other members of the same
family,286 should have been specially rendered pure white and thus made
conspicuous to their enemies; or that the adults of one of these two species
should have been specially rendered white during the winter in a country
which is never covered with snow. On the other hand we have reason to
believe that whiteness has been gained by many birds as a sexual ornament.
We may therefore conclude that an early progenitor of the Ardea asha and
the Buphus acquired a white plumage for nuptial purposes, and transmitted
this colour to their young; so that the young and the old became white like
certain existing egrets; the whiteness having afterwards been retained by the
young whilst exchanged by the adults for more strongly pronounced tints.
But if we could look still further backwards in time to the still earlier
progenitors of these two species, we should probably see the adults dark-
coloured. I infer that this would be the case, from the analogy of many other
birds, which are dark whilst young, and when adult are white; and more
especially from the case of the Ardea gularis, the colours of which are the
reverse of those of A. asha, for the young are dark-coloured and the adults
white, the young having retained a former state of plumage. It appears
therefore that the progenitors in their adult condition of the Ardea asha, the
Buphus, and of some allies, have undergone, during a long line of descent,
the following changes of colour: firstly a dark shade, secondly pure white,

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and thirdly, owing to another change of fashion (if I may so express
myself), their present slaty, reddish, or golden-buff tints. These successive
changes are intelligible only on the principle of novelty having been
admired by birds for the sake of novelty.

Summary of the Four Chapters on Birds.—Most male birds are highly
pugnacious during the breeding-season, and some possess weapons
especially adapted for fighting with their rivals. But the most pugnacious
and the best-armed males rarely or never depend for success solely on their
power to drive away or kill their rivals, but have special means for
charming the female. With some it is the power of song, or of emitting
strange cries, or of producing instrumental music, and the males in
consequence differ from the females in their vocal organs, or in the
structure of certain feathers. From the curiously diversified means for
producing various sounds we gain a high idea of the importance of this
means of courtship. Many birds endeavour to charm the females by love-
dances or antics, performed on the ground or in the air, and sometimes at
prepared places. But ornaments of many kinds, the most brilliant tints,
combs and wattles, beautiful plumes, elongated feathers, top-knots, and so
forth, are by far the commonest means. In some cases mere novelty appears
to have acted as a charm. The ornaments of the males must be highly
important to them, for they have been acquired in not a few cases at the cost
of increased danger from enemies, and even at some loss of power in
fighting with their rivals. The males of very many species do not assume
their ornamental dress until they arrive at maturity, or they assume it only
during the breeding-season, or the tints then become more vivid. Certain
ornamental appendages become enlarged, turgid, and brightly-coloured
during the very act of courtship. The males display their charms with
elaborate care and to the best effect; and this is done in the presence of the
females. The courtship is sometimes a prolonged affair, and many males
and females congregate at an appointed place. To suppose that the females
do not appreciate the beauty of the males is to admit that their splendid
decorations, all their pomp and display, are useless; and this is incredible.
Birds have fine powers of discrimination, and in some few instances it can
be shewn that they have a taste for the beautiful. The females, moreover, are
known occasionally to exhibit a marked preference or antipathy for certain
individual males.

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If it be admitted that the females prefer, or are unconsciously excited by the
more beautiful males, then the males would slowly but surely be rendered
more and more attractive through sexual selection. That it is this sex which
has been chiefly modified we may infer from the fact that in almost every
genus in which the sexes differ, the males differ much more from each other
than do the females; this is well shewn in certain closely-allied
representative species in which the females can hardly be distinguished,
whilst the males are quite distinct. Birds in a state of nature offer individual
differences which would amply suffice for the work of sexual selection; but
we have seen that they occasionally present more strongly-marked
variations which recur so frequently that they would immediately be fixed,
if they served to allure the female. The laws of variation will have
determined the nature of the initial changes, and largely influenced the final
result. The gradations, which may be observed between the males of allied
species, indicate the nature of the steps which have been passed through,
and explain in the most interesting manner certain characters, such as the
indented ocelli of the tail-feathers of the peacock, and the wonderfully-
shaded ocelli of the wing-feathers of the Argus pheasant. It is evident that
the brilliant colours, top-knots, fine plumes, &c., of many male birds cannot
have been acquired as a protection; indeed they sometimes lead to danger.
That they are not due to the direct and definite action of the conditions of
life, we may feel assured, because the females have been exposed to the
same conditions, and yet often differ from the males to an extreme degree.
Although it is probable that changed conditions acting during a lengthened
period have produced some definite effect on both sexes, the more
important result will have been an increased tendency to fluctuating
variability or to augmented individual differences; and such differences will
have afforded an excellent groundwork for the action of sexual selection.
The laws of inheritance, irrespectively of selection, appear to have
determined whether the characters acquired by the males for the sake of
ornament, for producing various sounds, and for fighting together, have
been transmitted to the males alone or to both sexes, either permanently or
periodically during certain seasons of the year. Why various characters
should sometimes have been transmitted in one way and sometimes in
another is, in most cases, not known; but the period of variability seems
often to have been the determining cause. When the two sexes have

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inherited all characters in common they necessarily resemble each other;
but as the successive variations may be differently transmitted, every
possible gradation may be found, even within the same genus, from the
closest similarity to the widest dissimilarity between the sexes. With many
closely-allied species, following nearly the same habits of life, the males
have come to differ from each other chiefly through the action of sexual
selection; whilst the females have come to differ chiefly from partaking in a
greater or lesser degree of the characters thus acquired by the males. The
effects, moreover, of the definite action of the conditions of life, will not
have been masked in the females, as in the case of the males, by the
accumulation through sexual selection of strongly-pronounced colours and
other ornaments. The individuals of both sexes, however affected, will have
been kept at each successive period nearly uniform by the free intercrossing
of many individuals.
With the species, in which the sexes differ in colour, it is possible that at
first there existed a tendency to transmit the successive variations equally to
both sexes; and that the females were prevented from acquiring the bright
colours of the males, on account of the danger to which they would have
been exposed during incubation. But it would be, as far as I can see, an
extremely difficult process to convert, by means of natural selection, one
form of transmission into another. On the other hand there would not be the
least difficulty in rendering a female dull-coloured, the male being still kept
bright-coloured, by the selection of successive variations, which were from
the first limited in their transmission to the same sex. Whether the females
of many species have actually been thus modified, must at present remain
doubtful. When, through the law of the equal transmission of characters to
both sexes, the females have been rendered as conspicuously coloured as
the males, their instincts have often been modified, and they have been led
to build domed or concealed nests.
In one small and curious class of cases the characters and habits of the two
sexes have been completely transposed, for the females are larger, stronger,
more vociferous and brightly-coloured than their males. They have, also,
become so quarrelsome that they often fight together like the males of the
most pugnacious species. If, as seems probable, they habitually drive away
rival females, and by the display of their bright colours or other charms
endeavour to attract the males, we can understand how it is that they have

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gradually been rendered, by means of sexual selection and sexually-limited
transmission, more beautiful than the males—the latter being left
unmodified or only slightly modified.
Whenever the law of inheritance at corresponding ages prevails, but not that
of sexually-limited transmission, then if the parents vary late in life—and
we know that this constantly occurs with our poultry, and occasionally with
other birds—the young will be left unaffected, whilst the adults of both
sexes will be modified. If both these laws of inheritance prevail and either
sex varies late in life, that sex alone will be modified, the other sex and the
young being left unaffected. When variations in brightness or in other
conspicuous characters occur early in life, as no doubt often happens, they
will not be acted on through sexual selection until the period of
reproduction arrives; consequently they will be liable to be lost by the
accidental deaths of the young, and if dangerous will be eliminated through
natural selection. Thus we can understand how it is that variations arising
late in life have chiefly been preserved for the ornamentation and arming of
the males, the females and the young being left almost unaffected, and
therefore like each other. With species having a distinct summer and winter
plumage, the males of which either resemble or differ from the females
during both seasons or during the summer alone, the degrees and kinds of
resemblance between the young and the old are exceedingly complex; and
this complexity apparently depends on characters, first acquired by the
males, being transmitted in various ways and degrees, as limited by age,
sex, and season.
As the young of so many species have been but little modified in colour and
in other ornaments, we are enabled to form some judgment with respect to
the plumage of their early progenitors; and we may infer that the beauty of
our existing species, if we look to the whole class, has been largely
increased since that period of which the immature plumage gives us an
indirect record. Many birds, especially those which live much on the
ground, have undoubtedly been obscurely coloured for the sake of
protection. In some instances the upper exposed surface of the plumage has
been thus coloured in both sexes, whilst the lower surface in the males
alone has been variously ornamented through sexual selection. Finally, from
the facts given in these four chapters, we may conclude that weapons for
battle, organs for producing sound, ornaments of many kinds, bright and

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conspicuous colours, have generally been acquired by the males through
variation and sexual selection, and have been transmitted in various ways
according to the several laws of inheritance—the females and the young
being left comparatively but little modified.287

CHAPTER XVII.

Secondary Sexual Characters of Mammals.

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The law of battle—Special weapons, confined to the males—Cause of absence of weapons in the
female—Weapons common to both sexes, yet primarily acquired by the male—Other uses of
such weapons—Their high importance—Greater size of the male—Means of defence—On the
preference shewn by either sex in the pairing of quadrupeds.

With mammals the male appears to win the female much more through the
law of battle than through the display of his charms. The most timid
animals, not provided with any special weapons for fighting, engage in
desperate conflicts during the season of love. Two male hares have been
seen to fight together until one was killed; male moles often fight, and
sometimes with fatal results; male squirrels “engage in frequent contests,
and often wound each other severely;” as do male beavers, so that “hardly a
skin is without scars.”288 I observed the same fact with the hides of the
guanacoes in Patagonia; and on one occasion several were so absorbed in
fighting that they fearlessly rushed close by me. Livingstone speaks of the
males of the many animals in Southern Africa as almost invariably shewing
the scars received in former contests.
The law of battle prevails with aquatic as with terrestrial mammals. It is
notorious how desperately male seals fight, both with their teeth and claws,
during the breeding-season; and their hides are likewise often covered with
scars. Male sperm-whales are very jealous at this season; and in their battles
“they often lock their jaws together, and turn on their sides and twist
about;” so that it is believed by some naturalists that the frequently
deformed state of their lower jaws is caused by these struggles.289
All male animals which are furnished with special weapons for fighting, are
well known to engage in fierce battles. The courage and the desperate
conflicts of stags have often been described; their skeletons have been
found in various parts of the world, with the horns inextricably locked
together, shewing how miserably the victor and vanquished had perished.290
No animal in the world is so dangerous as an elephant in must. Lord
Tankerville has given me a graphic description of the battles between the
wild bulls in Chillingham Park, the descendants, degenerated in size but not
in courage, of the gigantic Bos primigenius. In 1861 several contended for
mastery; and it was observed that two of the younger bulls attacked in
concert the old leader of the herd, overthrew and disabled him, so that he
was believed by the keepers to be lying mortally wounded in a
neighbouring wood. But a few days afterwards one of the young bulls

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singly approached the wood; and then the “monarch of the chase,” who had
been lashing himself up for vengeance, came out and, in a short time killed
his antagonist. He then quietly joined the herd, and long held undisputed
sway. Admiral Sir B. J. Sulivan informs me that when he resided in the
Falkland Islands he imported a young English stallion, which, with eight
mares, frequented the hills near Port William. On these hills there were two
wild stallions, each with a small troop of mares; “and it is certain that these
stallions would never have approached each other without fighting. Both
had tried singly to fight the English horse and drive away his mares, but had
failed. One day they came in together and attacked him. This was seen by
the capitan who had charge of the horses, and who, on riding to the spot,
found one of the two stallions engaged with the English horse, whilst the
other was driving away the mares, and had already separated four from the
rest. The capitan settled the matter by driving the whole party into the
corral, for the wild stallions would not leave the mares.”
Male animals already provided with efficient cutting or tearing teeth for the
ordinary purposes of life, as in the carnivora, insectivora, and rodents, are
seldom furnished with weapons especially adapted for fighting with their
rivals. The case is very different with the males of many other animals. We
see this in the horns of stags and of certain kinds of antelopes in which the
females are hornless. With many animals the canine teeth in the upper or
lower jaw, or in both, are much larger in the males than in the females; or
are absent in the latter, with the exception sometimes of a hidden rudiment.
Certain antelopes, the musk-deer, camel, horse, boar, various apes, seals,
and the walrus, offer instances of these several cases. In the females of the
walrus the tusks are sometimes quite absent.291 In the male elephant of India
and in the male dugong292 the upper incisors form offensive weapons. In the
male narwhal one alone of the upper teeth is developed into the well-
known, spirally-twisted, so called horn, which is sometimes from nine to
ten feet in length. It is believed that the males use these horns for fighting
together; for “an unbroken one can rarely be got, and occasionally one may
be found with the point of another jammed into the broken place.”293 The
tooth on the opposite side of the head in the male consists of a rudiment
about ten inches in length, which is embedded in the jaw. It is not, however,
very uncommon to find double-horned male narwhals in which both teeth
are well developed. In the females both teeth are rudimentary. The male

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cachalot has a larger head than that of the female, and it no doubt aids these
animals in their aquatic battles. Lastly, the adult male ornithorhynchus is
provided with a remarkable apparatus, namely a spur on the foreleg, closely
resembling the poison-fang of a venomous snake; its use is not known, but
we may suspect that it serves as a weapon of offence.294 It is represented by
a mere rudiment in the female.
When the males are provided with weapons which the females do not
possess, there can hardly be a doubt that they are used for fighting with
other males, and that they have been acquired through sexual selection.
It is not probable, at least in most cases, that the females have actually been
saved from acquiring such weapons, owing to their being useless and
superfluous, or in some way injurious. On the contrary, as they are often
used by the males of many animals for various purposes, more especially as
a defence against their enemies, it is a surprising fact that they are so poorly
developed or quite absent in the females. No doubt with female deer the
development during each recurrent season of great branching horns, and
with female elephants the development of immense tusks, would have been
a great waste of vital power, on the admission that they were of no use to
the females. Consequently variations in the size of these organs, leading to
their suppression, would have come under the control of natural selection,
and if limited in their transmission to the female offspring would not have
interfered with their development through sexual selection in the males. But
how on this view can we explain the presence of horns in the females of
certain antelopes, and of tusks in the females of many animals, which are
only of slightly less size than in the males? The explanation in almost all
cases must, I believe, be sought in the laws of transmission.
As the reindeer is the single species in the whole family of Deer in which
the female is furnished with horns, though somewhat smaller, thinner, and
less branched than in the male, it might naturally be thought that they must
be of some special use to her. There is, however, some evidence opposed to
this view. The female retains her horns from the time when they are fully
developed, namely in September, throughout the winter, until May, when
she brings forth her young; whilst the male casts his horns much earlier,
towards the end of November. As both sexes have the same requirements
and follow the same habits of life, and as the male sheds his horns during

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the winter, it is very improbable that they can be of any special service to
the female at this season, which includes the larger proportion of the time
during which she bears horns. Nor is it probable that she can have inherited
horns from some ancient progenitor of the whole family of deer, for, from
the fact of the males alone of so many species in all quarters of the globe
possessing horns, we may conclude that this was the primordial character of
the group. Hence it appears that horns must have been transferred from the
male to the female at a period subsequent to the divergence of the various
species from a common stock; but that this was not effected for the sake of
giving her any special advantage.295
We know that the horns are developed at a most unusually early age in the
reindeer; but what the cause of this may have been is not known. The effect,
however, has apparently been the transference of the horns to both sexes. It
is intelligible on the hypothesis of pangenesis, that a very slight change in
the constitution of the male, either in the tissues of the forehead or in the
gemmules of the horns, might lead to their early development; and as the
young of both sexes have nearly the same constitution before the period of
reproduction, the horns, if developed at an early age in the male, would tend
to be developed equally in both sexes. In support of this view, we should
bear in mind that the horns are always transmitted through the female, and
that she has a latent capacity for their development, as we see in old or
diseased females.296 Moreover the females of some other species of deer
either normally or occasionally exhibit rudiments of horns; thus the female
of Cervulus moschatus has “bristly tufts, ending in a knob, instead of a
horn;” and “in most specimens of the female Wapiti (Cervus Canadensis)
there is a sharp bony protuberance in the place of the horn.”297 From these
several considerations we may conclude that the possession of fairly well-
developed horns by the female reindeer, is due to the males having first
acquired them as weapons for fighting with other males; and secondarily to
their development from some unknown cause at an unusually early age in
the males, and their consequent transmission to both sexes.
Turning to the sheath-horned ruminants: with antelopes a graduated series
can be formed, beginning with the species, the females of which are
completely destitute of horns—passing to those which have horns so small
as to be almost rudimentary, as in Antilocapra Americana—to those which
have fairly well-developed horns, but manifestly smaller and thinner than in

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the male, and sometimes of a different shape,298 and ending with those in
which both sexes have horns of equal size. As with the reindeer, so with
antelopes there exists a relation between the period of the development of
the horns and their transmission to one or both sexes; it is therefore
probable that their presence or absence in the females of some species, and
their more or less perfect condition in the females of other species, depend,
not on their being of some special use, but simply on the form of
inheritance which has prevailed. It accords with this view that even in the
same restricted genus both sexes of some species, and the males alone of
other species, are thus provided. It is a remarkable fact that, although the
females of Antilope bezoartica are normally destitute of horns, Mr. Blyth
has seen no less than three females thus furnished; and there was no reason
to suppose that they were old or diseased. The males of this species have
long straight spirated horns, nearly parallel to each other, and directed
backwards. Those of the female, when present, are very different in shape,
for they are not spirated, and spreading widely bend round, so that their
points are directed forwards. It is a still more remarkable fact that in the
castrated male, as Mr. Blyth informs me, the horns are of the same peculiar
shape as in the female, but longer and thicker. In all cases the differences
between the horns of the males and females, and of castrated and entire
males, probably depend on various causes,—on the more or less complete
transference of male characters to the females,—on the former state of the
progenitors of the species,—and partly perhaps on the horns being
differently nourished, in nearly the same manner as the spurs of the
domestic cock, when inserted into the comb or other parts of the body,
assume various abnormal forms from being differently nourished.
In all the wild species of goats and sheep the horns are larger in the male
than in the female, and are sometimes quite absent in the latter.299 In several
domestic breeds of the sheep and goat, the males alone are furnished with
horns; and it is a significant fact, that in one such breed of sheep on the
Guinea coast, the horns are not developed, as Mr. Winwood Reade informs
me, in the castrated male; so that they are affected in this respect like the
horns of stags. In some breeds, as in that of N. Wales, in which both sexes
are properly horned, the ewes are very liable to be hornless. In these same
sheep, as I have been informed by a trustworthy witness who purposely
inspected a flock during the lambing-season, the horns at birth are generally

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more fully developed in the male than in the female. With the adult musk-
ox (Ovibos moschatus) the horns of the male are larger than those of the
female, and in the latter the bases do not touch.300 In regard to ordinary
cattle Mr. Blyth remarks: “In most of the wild bovine animals the horns are
both longer and thicker in the bull than in the cow, and in the cow-banteng
(Bos sondaicus) the horns are remarkably small, and inclined much
backwards. In the domestic races of cattle, both of the humped and
humpless types, the horns are short and thick in the bull, longer and more
slender in the cow and ox; and in the Indian buffalo, they are shorter and
thicker in the bull, longer and more slender in the cow. In the wild gaour (B.
gaurus) the horns are mostly both longer and thicker in the bull than in the
cow.”301 Hence with most sheath-horned ruminants the horns of the male are
either longer or stronger than those of the female. With the Rhinoceros
simus, as I may here add, the horns of the female are generally longer but
less powerful than in the male; and in some other species of rhinoceros they
are said to be shorter in the female.302 From these various facts we may
conclude that horns of all kinds, even when they are equally developed in
both sexes, were primarily acquired by the males in order to conquer other
males, and have been transferred more or less completely to the female, in
relation to the force of the equal form of inheritance.
The tusks of the elephant, in the different species or races, differ according
to sex, in nearly the same manner as the horns of ruminants. In India and
Malacca the males alone are provided with well-developed tusks. The
elephant of Ceylon is considered by most naturalists as a distinct race, but
by some as a distinct species, and here “not one in a hundred is found with
tusks, the few that possess them being exclusively males.”303 The African
elephant is undoubtedly distinct, and the female has large, well-developed
tusks, though not so large as those of the male. These differences in the
tusks of the several races and species of elephants—the great variability of
the horns of deer, as notably in the wild reindeer—the occasional presence
of horns in the female Antilope bezoartica—the presence of two tusks in
some few male narwhals—the complete absence of tusks in some female
walruses;—are all instances of the extreme variability of secondary sexual
characters, and of their extreme liability to differ in closely-allied forms.
Although tusks and horns appear in all cases to have been primarily
developed as sexual weapons, they often serve for other purposes. The

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elephant uses his tusks in attacking the tiger; according to Bruce, he scores
the trunks of trees until they can be easily thrown down, and he likewise
thus extracts the farinaceous cores of palms; in Africa he often uses one
tusk, this being always the same, to probe the ground and thus to ascertain
whether it will bear his weight. The common bull defends the herd with his
horns; and the elk in Sweden has been known, according to Lloyd, to strike
a wolf dead with a single blow of his great horns. Many similar facts could
be given. One of the most curious secondary uses to which the horns of any
animal are occasionally put, is that observed by Captain Hutton304 with the
wild goat (Capra ægagrus) of the Himalayas, and as it is said with the ibex,
namely, that when the male accidentally falls from a height he bends
inwards his head, and, by alighting on his massive horns, breaks the shock.
The female cannot thus use her horns, which are smaller, but from her more
quiet disposition she does not so much need this strange kind of shield.
Each male animal uses his weapons in his own peculiar fashion. The
common ram makes a charge and butts with such force with the bases of his
horns, that I have seen a powerful man knocked over as easily as a child.
Goats and certain species of sheep, for instance the Ovis cycloceros of
Afghanistan,305 rear on their hind legs, and then not only butt, but “make a
cut down and a jerk up, with the ribbed front of their scimitar-shaped horn,
as with a sabre. When the O. cycloceros attacked a large domestic ram, who
was a noted bruiser, he conquered him by the sheer novelty of his mode of
fighting, always closing at once with his adversary, and catching him across
the face and nose with a sharp drawing jerk of his head, and then bounding
out of the way before the blow could be returned.” In Pembrokeshire a male
goat, the master of a flock which during several generations had run wild,
was known to have killed several other males in single combat; this goat
possessed enormous horns, measuring 39 inches in a straight line from tip
to tip. The common bull, as every one knows, gores and tosses his
opponent; but the Italian buffalo is said never to use his horns, he gives a
tremendous blow with his convex forehead, and then tramples on his fallen
enemy with his knees—an instinct which the common bull does not
possess.306 Hence a dog who pins a buffalo by the nose is immediately
crushed. We must, however, remember that the Italian buffalo has long been
domesticated, and it is by no means certain that the wild parent-form had
similarly shaped horns. Mr. Bartlett informs me that when a female Cape

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buffalo (Bubalus caffer) was turned into an enclosure with a bull of the
same species, she attacked him, and he in return pushed her about with
great violence. But it was manifest to Mr. Bartlett that had not the bull
shewn dignified forbearance, he could easily have killed her by a single
lateral thrust with his immense horns. The giraffe uses his short hair-
covered horns, which are rather longer in the male than in the female, in a
curious manner; for with his long neck he swings his head to either side,
almost upside down, with such force, that I have seen a hard plank deeply
indented by a single blow.

Fig. 61. Oryx leucoryx, male (from the Knowsley Menagerie).

With antelopes it is sometimes difficult to imagine how they can possibly
use their curiously-shaped horns; thus the spring-boc (Ant. euchore) has
rather short upright horns, with the sharp points bent inwards almost at a
right angle, so as to face each other; Mr. Bartlett does not know how they
are used, but suggests that they would inflict a fearful wound down each
side of the face of an antagonist. The slightly-curved horns of the Oryx
leucoryx (fig. 61) are directed backwards, and are of such length that their
points reach beyond the middle of the back, over which they stand in an
almost parallel line. Thus they seem singularly ill-fitted for fighting; but Mr.
Bartlett informs me that when two of these animals prepare for battle, they
kneel down, with their heads between their front legs, and in this attitude

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the horns stand nearly parallel and close to the ground, with the points
directed forwards and a little upwards. The combatants then gradually
approach each other and endeavour to get the upturned points under each
other’s bodies; if one succeeds in doing this, he suddenly springs up,
throwing up his head at the same time, and can thus wound or perhaps even
transfix his antagonist. Both animals always kneel down so as to guard as
far as possible against this manœuvre. It has been recorded that one of these
antelopes has used his horns with effect even against a lion; yet from being
forced to place his head between the forelegs in order to bring the points of
the horns forward, he would generally be under a great disadvantage when
attacked by any other animal. It is, therefore, not probable that the horns
have been modified into their present great length and peculiar position, as
a protection against beasts of prey. We can, however, see that as soon as
some ancient male progenitor of the Oryx acquired moderately long horns,
directed a little backwards, he would be compelled in his battles with rival
males to bend his head somewhat inwards or downwards, as is now done by
certain stags; and it is not improbable that he might have acquired the habit
of at first occasionally and afterwards of regularly kneeling down. In this
case it is almost certain that the males which possessed the longest horns
would have had a great advantage over others with shorter horns; and then
the horns would gradually have been rendered longer and longer, through
sexual selection, until they acquired their present extraordinary length and
position.
With stags of many kinds the branching of the horns offers a curious case of
difficulty; for certainly a single straight point would inflict a much more
serious wound than several diverging points. In Sir Philip Egerton’s
museum there is a horn of the red-deer (Cervus elaphus) thirty inches in
length, with “not fewer than fifteen snags or branches;” and at Moritzburg
there is still preserved a pair of antlers of a red-deer, shot in 1699 by
Frederick I., each of which bears the astonishing number of thirty-three
branches. Richardson figures a pair of antlers of the wild reindeer with
twenty-nine points.307 From the manner in which the horns are branched,
and more especially from deer being known occasionally to fight together
by kicking with their fore-feet,308 M. Bailly actually came to the conclusion
that their horns were more injurious than useful to them! But this author
overlooks the pitched battles between rival males. As I felt much perplexed

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about the use or advantage of the branches, I applied to Mr. McNeill of
Colinsay, who has long and carefully observed the habits of red-deer, and
he informs me that he has never seen some of the branches brought into
action, but that the brow-antlers, from inclining downwards, are a great
protection to the forehead, and their points are likewise used in attack. Sir
Philip Egerton also informs me in regard both to red-deer and fallow-deer,
that when they fight they suddenly dash together, and getting their horns
fixed against each other’s bodies a desperate struggle ensues. When one is
at last forced to yield and turn round, the victor endeavours to plunge his
brow-antlers into his defeated foe. It thus appears that the upper branches
are used chiefly or exclusively for pushing and fencing. Nevertheless with
some species the upper branches are used as weapons of offence; when a
man was attacked by a Wapiti deer (Cervus Canadensis) in Judge Caton’s
park in Ottawa, and several men tried to rescue him, the stag “never raised
his head from the ground; in fact he kept his face almost flat on the ground,
with his nose nearly between his fore-feet, except when he rolled his head
to one side to take a new observation preparatory to a plunge.” In this
position the terminal points of the horns were directed against his
adversaries. “In rolling his head he necessarily raised it somewhat, because
his antlers were so long that he could not roll his head without raising them
on one side, while on the other side they touched the ground.” The stag by
this procedure gradually drove the party of rescuers backwards, to a
distance of 150 or 200 feet; and the attacked man was killed.309
Although the horns of stags are efficient weapons, there can, I think, be no
doubt that a single point would have been much more dangerous than a
branched antler; and Judge Caton, who has had large experience with deer,
fully concurs in this conclusion. Nor do the branching horns, though highly
important as a means of defence against rival stags, appear perfectly well
adapted for this purpose, as they are liable to become interlocked. The
suspicion has therefore crossed my mind that they may serve partly as
ornaments. That the branched antlers of stags, as well as the elegant lyrated
horns of certain antelopes, with their graceful double curvature, (fig. 62),
are ornamental in our eyes, no one will dispute. If, then, the horns, like the
splendid accoutrements of the knights of old, add to the noble appearance of
stags and antelopes, they may have been partly modified for this purpose,

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though mainly for actual service in battle; but I have no evidence in favour
of this belief.

Fig. 62. Strepsiceros Kudu (from Andrew Smith’s ‘Zoology of South Africa’).

An interesting case has lately been published, from which it appears that the
horns of a deer in one district in the United States are now being modified
through sexual and natural selection. A writer in an excellent American
Journal310 says, that he has hunted for the last twenty-one years in the
Adirondacks, where the Cervus Virginianus abounds. About fourteen years
ago he first heard of spike-horn bucks. These became from year to year
more common; about five years ago he shot one, and subsequently another,

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and now they are frequently killed. “The spike-horn differs greatly from the
common antler of the C. Virginianus. It consists of a single spike, more
slender than the antler, and scarcely half so long, projecting forward from
the brow, and terminating in a very sharp point. It gives a considerable
advantage to its possessor over the common buck. Besides enabling him to
run more swiftly through the thick woods and underbrush (every hunter
knows that does and yearling bucks run much more rapidly than the large
bucks when armed with their cumbrous antlers), the spike-horn is a more
effective weapon than the common antler. With this advantage the spike-
horn bucks are gaining upon the common bucks, and may, in time, entirely
supersede them in the Adirondacks. Undoubtedly the first spike-horn buck
was merely an accidental freak of nature. But his spike-horns gave him an
advantage, and enabled him to propagate his peculiarity. His descendants,
having a like advantage, have propagated the peculiarity in a constantly
increasing ratio, till they are slowly crowding the antlered deer from the
region they inhabit.”
Male quadrupeds which are furnished with tusks use them in various ways,
as in the case of horns. The boar strikes laterally and upwards; the musk-
deer with serious effect downwards.311 The walrus, though having so short a
neck and so unwieldy a body, “can strike either upwards, or downwards, or
sideways, with equal dexterity.”312 The Indian elephant fights, as I was
informed by the late Dr. Falconer, in a different manner according to the
position and curvature of his tusks. When they are directed forwards and
upwards he is able to fling a tiger to a great distance—it is said to even
thirty feet; when they are short and turned downwards he endeavours
suddenly to pin the tiger to the ground, and in consequence is dangerous to
the rider, who is liable to be jerked off the hoodah.313
Very few male quadrupeds possess weapons of two distinct kinds specially
adapted for fighting with rival males. The male muntjac-deer (Cervulus),
however, offers an exception, as he is provided with horns and exserted
canine teeth. But one form of weapon, has often been replaced in the course
of ages by another form, as we may infer from what follows. With
ruminants the development of horns generally stands in an inverse relation
with that of even moderately well-developed canine teeth. Thus camels,
guanacoes, chevrotains and musk-deer, are hornless, and they have efficient
canines; these teeth being “always of smaller size in the females than in the

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males.” The Camelidæ have in their upper jaws, in addition to their true
canines, a pair of canine-shaped incisors.314 Male deer and antelopes, on the
other hand, possess horns, and they rarely have canine teeth; and these
when present are always of small size, so that it is doubtful whether they are
of any service in their battles. With Antilope montana they exist only as
rudiments in the young male, disappearing as he grows old; and they are
absent in the female at all ages; but the females of certain other antelopes
and deer have been known occasionally to exhibit rudiments of these
teeth.315 Stallions have small canine teeth, which are either quite absent or
rudimentary in the mare; but they do not appear to be used in fighting, for
stallions bite with their incisors, and do not open their mouths widely like
camels and guanacoes. Whenever the adult male possesses canines now in
an inefficient state, whilst the female has either none or mere rudiments, we
may conclude that the early male progenitor of the species was provided
with efficient canines, which had been partially transferred to the females.
The reduction of these teeth in the males seems to have followed from some
change in their manner of fighting, often caused (but not in the case of the
horse) by the development of new weapons.
Tusks and horns are manifestly of high importance to their possessors, for
their development consumes much organised matter. A single tusk of the
Asiatic elephant,—one of the extinct woolly species,—and of the African
elephant, have been known to weigh respectively 150, 160, and 180
pounds; and even greater weights have been assigned by some authors.316
With deer, in which the horns are periodically renewed, the drain on the
constitution must be greater; the horns, for instance, of the moose weigh
from fifty to sixty pounds, and those of the extinct Irish elk from sixty to
seventy pounds,—the skull of the latter weighing on an average only five
and a quarter pounds. With sheep, although the horns are not periodically
renewed, yet their development, in the opinion of many agriculturists,
entails a sensible loss to the breeder. Stags, moreover, in escaping from
beasts of prey are loaded with an additional weight for the race, and are
greatly retarded in passing through a woody country. The moose, for
instance, with horns extending five and a half feet from tip to tip, although
so skilful in their use that he will not touch or break a dead twig when
walking quietly, cannot act so dexterously whilst rushing away from a pack
of wolves. “During his progress he holds his nose up, so as to lay the horns

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horizontally back; and in this attitude cannot see the ground distinctly.”317
The tips of the horns of the great Irish elk were actually eight feet apart!
Whilst the horns are covered with velvet, which lasts with the red-deer for
about twelve weeks, they are extremely sensitive to a blow; so that in
Germany the stags at this time change their habits to a certain extent, and
avoid dense forests, frequenting young woods and low thickets.318 These
facts remind us, that male birds have acquired ornamental plumes at the
cost of retarded flight, and other ornaments at the cost of some loss of
power in their battles with rival males.
With quadrupeds, when, as is often the case, the sexes differ in size, the
males are, I believe, always larger and stronger. This holds good in a
marked manner, as I am informed by Mr. Gould, with the marsupials of
Australia, the males of which appear to continue growing until an unusually
late age. But the most extraordinary case is that of one of the seals
(Callorhinus ursinus), a full-grown female weighing less than one-sixth of a
full-grown male.319 The greater strength of the male is invariably displayed,
as Hunter long ago remarked,320 in those parts of the body which are
brought into action in fighting with rival males,—for instance, in the
massive neck of the bull. Male quadrupeds are also more courageous and
pugnacious than the females. There can be little doubt that these characters
have been gained, partly through sexual selection, owing to a long series of
victories by the stronger and more courageous males over the weaker, and
partly through the inherited effects of use. It is probable that the successive
variations in strength, size, and courage, whether due to so-called
spontaneous variability or to the effects of use, by the accumulation of
which male quadrupeds have acquired these characteristic qualities,
occurred rather late in life, and were consequently to a large extent limited
in their transmission to the same sex.
Under this point of view I was anxious to obtain information in regard to
the Scotch deerhound, the sexes of which differ more in size than those of
any other breed (though blood-hounds differ considerably), or than in any
wild canine species known to me.
Accordingly, I applied to Mr. Cupples, a well-known breeder of these dogs,
who has weighed and measured many of his own dogs, and who, with great
kindness, has collected for me the following facts from various sources.

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Superior male dogs, measured at the shoulder, range from twenty-eight
inches, which is low, to thirty-three, or even thirty-four inches in height;
and in weight from eighty pounds, which is low, to 120, or even more
pounds. The females range in height from twenty-three to twenty-seven, or
even to twenty-eight inches; and in weight from fifty to seventy, or even
eighty pounds.321 Mr. Cupples concludes that from ninety-five to one
hundred pounds for the male, and seventy for the female, would be a safe
average; but there is reason to believe that formerly both sexes attained a
greater weight. Mr. Cupples has weighed puppies when a fortnight old; in
one litter the average weight of four males exceeded that of two females by
six and a half ounces; in another litter the average weight of four males
exceeded that of one female by less than one ounce; the same males, when
three weeks old, exceeded the female by seven and a half ounces, and at the
age of six weeks by nearly fourteen ounces. Mr. Wright of Yeldersley
House, in a letter to Mr. Cupples, says: “I have taken notes on the sizes and
weights of puppies of many litters, and as far as my experience goes, dog-
puppies as a rule differ very little from bitches till they arrive at about five
or six months old; and then the dogs begin to increase, gaining upon the
bitches both in weight and size. At birth, and for several weeks afterwards,
a bitch-puppy will occasionally be larger than any of the dogs, but they are
invariably beaten by them later.” Mr. McNeill, of Colinsay, concludes that
“the males do not attain their full growth till over two years old, though the
females attain it sooner.” According to Mr. Cupples’ experience, male dogs
go on growing in stature till they are from twelve to eighteen months old,
and in weight till from eighteen to twenty-four months old; whilst the
females cease increasing in stature at the age of from nine to fourteen or
fifteen months, and in weight at the age of from twelve to fifteen months.
From these various statements it is clear that the full difference in size
between the male and female Scotch deerhound is not acquired until rather
late in life. The males are almost exclusively used for coursing, for, as Mr.
McNeill informs me, the females have not sufficient strength and weight to
pull down a full-grown deer. From the names used in old legends, it
appears, as I hear from Mr. Cupples, that at a very ancient period the males
were the most celebrated, the females being mentioned only as the mothers
of famous dogs. Hence during many generations, it is the male which has
been chiefly tested for strength, size, speed, and courage, and the best will
have been bred from. As, however, the males do not attain their full

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dimensions until a rather late period in life, they will have tended, in
accordance with the law often indicated, to transmit their characters to their
male offspring alone; and thus the great inequality in size between the sexes
of the Scotch deerhound may probably be accounted for.
The males of some few quadrupeds possess organs or parts developed
solely as a means of defence against the attacks of other males. Some kinds
of deer use, as we have seen, the upper branches of their horns chiefly or
exclusively for defending themselves; and the Oryx antelope, as I am
informed by Mr. Bartlett, fences most skilfully with his long, gently curved
horns; but these are likewise used as organs of offence. Rhinoceroses, as the
same observer remarks, in fighting parry each other’s sidelong blows with
their horns, which loudly clatter together, as do the tusks of boars. Although
wild boars fight desperately together, they seldom, according to Brehm,
receive fatal blows, as these fall on each other’s tusks, or on the layer of
gristly skin covering the shoulder, which the German hunters call the shield;
and here we have a part specially modified for defence. With boars in the
prime of life (see fig. 63) the
tusks in the lower jaw are used
for fighting but they become in
old age, as Brehm states, so
much curved inwards and
upwards, over the snout, that they
can no longer be thus used. They
may, however, still continue to
serve, and even in a still more
effective manner, as a means of
defence. In compensation for the
loss of the lower tusks as
weapons of offence, those in the Fig. 63. Head of common wild boar, in prime of life
upper jaw, which always project (from Brehm).
a little laterally, increase so much
in length during old age, and curve so much upwards, that they can be used
as a means of attack. Nevertheless an old boar is not so dangerous to man as
one at the age of six or seven years.322

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Fig. 64. Skull of the Babirusa Pig (from Wallace’s ‘Malay
Archipelago’).

In the full-grown male Babirusa pig of Celebes (fig. 64), the lower tusks are
formidable weapons, like those of the European boar in the prime of life,
whilst the upper tusks are so long and have their points so much curled
inwards, sometimes even touching the forehead, that they are utterly useless
as weapons of attack. They more nearly resemble horns than teeth, and are
so manifestly useless as teeth that the animal was formerly supposed to rest
his head by hooking them on to a branch. Their convex surfaces would,
however, if the head were held a little laterally, serve as an excellent guard;
and hence, perhaps it is that in old animals they “are generally broken off,
as if by fighting.”323 Here, then, we have the curious case of the upper tusks
of the Babirusa regularly assuming during the prime of life, a structure
which apparently renders them fitted only for defence; whilst in the
European boar the lower and opposite tusks assume in a less degree and
only during old age nearly the same form, and then serve in like manner
solely for defence.

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Fig. 65. Head of Æthiopian Wart-hog, from ‘Proc. Zool. Soc.’ 1869. (I now find that this drawing
represents the head of a female, but it serves to shew, on a reduced scale, the characters of the
male.)

In the wart-hog (Phacochœrus æthiopicus, fig. 65) the tusks in the upper
jaw of the male curve upwards during the prime of life, and from being
pointed, serve as formidable weapons. The tusks in the lower jaw are
sharper than those in the upper, but from their shortness it seems hardly
possible that they can be used as weapons of attack. They must, however,
greatly strengthen those in the upper jaw, from being ground so as to fit
closely against their bases. Neither the upper nor the lower tusks appear to
have been specially modified to act as guards, though, no doubt, they are
thus used to a certain extent. But the wart-hog is not destitute of other
special means of protection, for there exists, on each side of the face,
beneath the eyes, a rather stiff, yet flexible, cartilaginous, oblong pad (fig.
65), which projects two or three inches outwards; and it appeared to Mr.
Bartlett and myself, when viewing the living animal, that these pads, when
struck from beneath by the tusks of an opponent, would be turned upwards,
and would thus protect in an admirable manner the somewhat prominent
eyes. These boars, as I may add on the authority of Mr. Bartlett, when
fighting together, stand directly face to face.
Lastly, the African river-hog (Potamochoerus penicillatus) has a hard
cartilaginous knob on each side of the face beneath the eyes, which answers
to the flexible pad of the wart-hog; it has also two bony prominences on the
upper jaw above the nostrils. A boar of this species in the Zoological
Gardens recently broke into the cage of the wart-hog. They fought all night-

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long, and were found in the morning much exhausted, but not seriously
wounded. It is a significant fact, as shewing the purpose of the above-
described projections and excrescences, that these were covered with blood,
and were scored and abraded in an extraordinary manner.
The mane of the lion forms a good defence against the one danger to which
he is liable, namely the attacks of rival lions: for the males, as Sir. A. Smith
informs me, engage in terrible battles, and a young lion dares not approach
an old one. In 1857 a tiger at Bromwich broke into the cage of a lion, and a
fearful scene ensued; “the lion’s mane saved his neck and head from being
much injured, but the tiger at last succeeded in ripping up his belly, and in a
few minutes he was dead.”324 The broad ruff round the throat and chin of the
Canadian lynx (Felis Canadensis) is much longer in the male than in the
female; but whether it serves as a defence I do not know. Male seals are
well known to fight desperately together, and the males of certain kinds
(Otaria jubata)325 have great manes, whilst the females have small ones or
none. The male baboon of the Cape of Good Hope (Cynocephalus
porcarius) has a much longer mane and larger canine teeth than the female;
and the mane probably serves as a protection, for on asking the keepers in
the Zoological Gardens, without giving them any clue to my object,
whether any of the monkeys especially attacked each other by the nape of
the neck, I was answered that this was not the case, excepting with the
above baboon. In the Hamadryas baboon, Ehrenberg compares the mane of
the adult male to that of a young lion, whilst in the young of both sexes and
in the female the mane is almost absent.
It appeared to me probable that the immense woolly mane of the male
American bison, which reaches almost to the ground, and is much more
developed in the males than in the females, served as a protection to them in
their terrible battles; but an experienced hunter told Judge Caton that he had
never observed anything which favoured this belief. The stallion has a
thicker and fuller mane than the mare; and I have made particular inquiries
of two great trainers and breeders who have had charge of many entire
horses, and am assured that they “invariably endeavour to seize one another
by the neck.” It does not, however, follow from the foregoing statements,
that when the hair on the neck serves as a defence, that it was originally
developed for this purpose, though this is probable in some cases, as in that
of the lion. I am informed by Mr. McNeill that the long hairs on the throat

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of the stag (Cervus elephas) serve as a great protection to him when hunted,
for the dogs generally endeavour to seize him by the throat; but it is not
probable that these hairs were specially developed for this purpose;
otherwise the young and the females would, as we may feel assured, have
been equally protected.

On Preference or Choice in Pairing, as shewn by either sex of Quadrupeds.
—Before describing, in the next chapter, the differences between the sexes
in voice, odour emitted, and ornamentation, it will be convenient here to
consider whether the sexes exert any choice in their unions. Does the
female prefer any particular male, either before or after the males may have
fought together for supremacy; or does the male, when not a polygamist,
select any particular female? The general impression amongst breeders
seems to be that the male accepts any female; and this, owing to his
eagerness, is, in most cases, probably the truth. Whether the female as a
general rule indifferently accepts any male is much more doubtful. In the
fourteenth chapter, on Birds, a considerable body of direct and indirect
evidence was advanced, shewing that the female selects her partner; and it
would be a strange anomaly if female quadrupeds, which stand higher in the
scale of organisation and have higher mental powers, did not generally, or at
least often, exert some choice. The female could in most cases escape, if
wooed by a male that did not please or excite her; and when pursued, as so
incessantly occurs, by several males, she would often have the opportunity,
whilst they were fighting together, of escaping with, or at least of
temporarily pairing with, some one male. This latter contingency has often
been observed in Scotland with female red-deer, as I have been informed by
Sir Philip Egerton.326
It is scarcely possible that much should be known about female quadrupeds
exerting in a state of nature any choice in their marriage unions. The
following very curious details on the courtship of one of the eared seals,
Callorhinus ursinus, are given327 on the authority of Capt. Bryant, who had
ample opportunities for observation. He says, “Many of the females on their
arrival at the island where they breed appear desirous of returning to some
particular male, and frequently climb the outlying rocks to overlook the
rookeries, calling out and listening as if for a familiar voice. Then changing
to another place they do the same again.... As soon as a female reaches the

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shore, the nearest male goes down to meet her, making meanwhile a noise
like the clucking of a hen to her chickens. He bows to her and coaxes her
until he gets between her and the water so that she cannot escape him. Then
his manner changes, and with a harsh growl he drives her to a place in his
harem. This continues until the lower row of harems is nearly full. Then the
males higher up select the time when their more fortunate neighbours are
off their guard to steal their wives. This they do by taking them in their
mouths and lifting them over the heads of the other females, and carefully
placing them in their own harem, carrying them as cats do their kittens.
Those still higher up pursue the same method until the whole space is
occupied. Frequently a struggle ensues between two males for the
possession of the same female, and both seizing her at once pull her in two
or terribly lacerate her with their teeth. When the space is all filled, the old
male walks around complacently reviewing his family, scolding those who
crowd or disturb the others, and fiercely driving off all intruders. This
surveillance always keeps him actively occupied.”
As so little is known about the courtship of animals in a state of nature, I
have endeavoured to discover how far our domesticated quadrupeds evince
any choice in their unions. Dogs offer the best opportunity for observation,
as they are carefully attended to and well understood. Many breeders have
expressed a strong opinion on this head. Thus Mr. Mayhew remarks, “The
females are able to bestow their affections; and tender recollections are as
potent over them as they are known to be in other cases, where higher
animals are concerned. Bitches are not always prudent in their loves, but are
apt to fling themselves away on curs of low degree. If reared with a
companion of vulgar appearance, there often springs up between the pair a
devotion which no time can afterwards subdue. The passion, for such it
really is, becomes of a more than romantic endurance.” Mr. Mayhew, who
attended chiefly to the smaller breeds, is convinced that the females are
strongly attracted by males of large size.328 The well-known veterinary
Blaine states329 that his own female pug became so attached to a spaniel,
and a female setter to a cur, that in neither case would they pair with a dog
of their own breed until several weeks had elapsed. Two similar and
trustworthy accounts have been given me in regard to a female retriever and
a spaniel, both of which became enamoured with terrier-dogs.

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Mr. Cupples informs me that he can personally vouch for the accuracy of
the following more remarkable case, in which a valuable and wonderfully-
intelligent female terrier loved a retriever, belonging to a neighbour, to such
a degree that she had often to be dragged away from him. After their
permanent separation, although repeatedly shewing milk in her teats, she
would never acknowledge the courtship of any other dog, and to the regret
of her owner, never bore puppies. Mr. Cupples also states that a female
deerhound now (1868) in his kennel has thrice produced puppies, and on
each occasion shewed a marked preference for one of the largest and
handsomest, but not the most eager, of four deerhounds living with her, all
in the prime of life. Mr. Cupples has observed that the female generally
favours a dog whom she has associated with and knows; her shyness and
timidity at first incline her against a strange dog. The male, on the contrary,
seems rather inclined towards strange females. It appears to be rare when
the male refuses any particular female, but Mr. Wright, of Yeldersley
House, a great breeder of dogs, informs me that he has known some
instances; he cites the case of one of his own deerhounds, who would not
take any notice of a particular female mastiff, so that another deerhound had
to be employed. It would be superfluous to give other cases, and I will only
add that Mr. Barr, who has carefully bred many blood-hounds, states that in
almost every instance particular individuals of the opposite sex shew a
decided preference for each other. Finally Mr. Cupples, after attending to
this subject for another year, has recently written to me, “I have had full
confirmation of my former statement, that dogs in breeding form decided
preferences for each other, being often influenced by size, bright colour, and
individual character, as well as by the degree of their previous familiarity.”
In regard to horses, Mr. Blenkiron, the greatest breeder of race-horses in the
world, informs me that stallions are so frequently capricious in their choice,
rejecting one mare and without any apparent cause taking to another, that
various artifices have to be habitually used. The famous Monarque, for
instance, would never consciously look at the dam of Gladiateur, and a trick
had to be practised. We can partly see the reason why valuable race-horse
stallions, which are in such demand, should be so particular in their choice.
Mr. Blenkiron has never known a mare to reject a horse; but this has
occurred in Mr. Wright’s stable, so that the mare had to be cheated. Prosper
Lucas330 quotes various statements from French authorities, and remarks,

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“On voit des étalons qui s’éprennent d’une jument, et négligent toutes les
autres.” He gives, on the authority of Baëlen, similar facts in regard to bulls.
Hoffberg, in describing the domesticated reindeer of Lapland, says,
“Fœmina majores et fortiores mares præ cæteris admittunt, ad eos
confugiunt, a junioribus agitatæ, qui hos in fugam conjiciunt.”331 A clergy-
man, who has bred many pigs, assures me that sows often reject one boar
and immediately accept another.
From these facts there can be no doubt that with most of our domesticated
quadrupeds strong individual antipathies and preferences are frequently
exhibited, and much more commonly by the female than by the male. This
being the case, it is improbable that the unions of quadrupeds in a state of
nature should be left to mere chance. It is much more probable that the
females are allured or excited by particular males, who possess certain
characters in a higher degree than other males; but what these characters
are, we can seldom or never discover with certainty.

CHAPTER XVIII.

Secondary Sexual Characters of Mammals—continued.
Voice—Remarkable sexual peculiarities in seals—Odour—Development of the hair—Colour of the
hair and skin—Anomalous case of the female being more ornamented than the male—Colour
and ornaments due to sexual selection—Colour acquired for the sake of protection—Colour,
though common to both sexes, often due to sexual selection—On the disappearance of spots and
stripes in adult quadrupeds—On the colours and ornaments of the Quadrumana—Summary.

Quadrupeds use their voices for various purposes, as a signal of danger, as a
call from one member of a troop to another, or from the mother to her lost
offspring, or from the latter for protection to their mother; but such uses
need not here be considered. We are concerned only with the difference
between the voices of the two sexes, for instance between that of the lion
and lioness, or of the bull and cow. Almost all male animals use their voices
much more during the rutting-season than at any other time; and some, as
the giraffe and porcupine,332 are said to be completely mute excepting at this
season. As the throats (i.e. the larynx and thyroid bodies333) of stags become
periodically enlarged at the commencement of the breeding-season, it might

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be thought that their powerful voices must be then in some way of high
importance to them; but this is very doubtful. From information given to me
by two experienced observers, Mr. McNeill and Sir P. Egerton, it seems that
young stags under three years old do not roar or bellow; and that the old
ones begin bellowing at the commencement of the breeding-season, at first
only occasionally and moderately, whilst they restlessly wander about in
search of the females. Their battles are prefaced by loud and prolonged
bellowing, but during the actual conflict they are silent. Animals of all kinds
which habitually use their voices, utter various noises under any strong
emotion, as when enraged and preparing to fight; but this may merely be
the result of their nervous excitement, which leads to the spasmodic
contraction of almost all the muscles of the body, as when a man grinds his
teeth and clenches his hands in rage or agony. No doubt stags challenge
each other to mortal combat by bellowing; but it is not likely that this habit
could have led through sexual selection, that is by the loudest-voiced males
having been the most successful in their conflicts, to the periodical
enlargement of the vocal organs; for the stags with the most powerful
voices, unless at the same time the strongest, best-armed, and most
courageous, would not have gained any advantage over their rivals with
weaker voices. The stags, moreover, which had weaker voices, though not
so well able to challenge other stags, would have been drawn to the place of
combat as certainly as those with stronger voices.
It is possible that the roaring of the lion may be of some actual service to
him in striking terror into his adversary; for when enraged he likewise
erects his mane and thus instinctively tries to make himself appear as
terrible as possible. But it can hardly be supposed that the bellowing of the
stag, even if it be of any service to him in this way, can have been important
enough to have led to the periodical enlargement of the throat. Some writers
suggest that the bellowing serves as a call to the female; but the experienced
observers above quoted inform me that female deer do not search for the
male, though the males search eagerly for the females, as indeed might be
expected from what we know of the habits of other male quadrupeds. The
voice of the female, on the other hand, quickly brings to her one or more
stags,334 as is well known to the hunters who in wild countries imitate her
cry. If we could believe that the male had the power to excite or allure the
female by his voice, the periodical enlargement of his vocal organs would

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be intelligible on the principle of sexual selection, together with inheritance
limited to the same sex and season of the year; but we have no evidence in
favour of this view. As the case stands, the loud voice of the stag during the
breeding season does not seem to be of any special service to him, either
during his courtship or battles, or in any other way. But may we not believe
that the frequent use of the voice, under the strong excitement of love,
jealousy, and rage, continued during many generations, may at last have
produced an inherited effect on the vocal organs of the stag, as well as of
other male animals? This appears to me, with our present state of
knowledge, the most probable view.
The male gorilla has a tremendous voice, and when adult is furnished with a
laryngeal sack, as is likewise the adult male orang.335 The gibbons rank
amongst the noisiest of monkeys, and the Sumatra species (Hylobates
syndactylus) is also furnished with a laryngeal sack; but Mr. Blyth, who has
had opportunities for observation, does not believe that the male is more
noisy than the female. Hence, these latter monkeys probably use their
voices as a mutual call; and this is certainly the case with some quadrupeds,
for instance with the beaver.336 Another gibbon, the H. agilis, is highly
remarkable, from having the power of emitting a complete and correct
octave of musical notes,337 which we may reasonably suspect serves as a
sexual charm; but I shall have to recur to this subject in the next chapter.
The vocal organs of the American Mycetes caraya are one-third larger in
the male than in the female, and are wonderfully powerful. These monkeys,
when the weather is warm, make the forests resound during the morning
and evening with their overwhelming voices. The males begin the dreadful
concert, in which the females, with their less powerful voices, sometimes
join, and which is often continued during many hours. An excellent
observer, Rengger,338 could not perceive that they were excited to begin
their concert by any special cause; he thinks that like many birds, they
delight in their own music, and try to excel each other. Whether most of the
foregoing monkeys have acquired their powerful voices in order to beat
their rivals and to charm the females—or whether the vocal organs have
been strengthened and enlarged through the inherited effects of long-
continued use without any particular good being gained—I will not pretend
to say; but the former view, at least in the case of the Hylobates agilis,
seems the most probable.

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I may here mention two very curious sexual peculiarities occurring in seals,
because they have been supposed by some writers to affect the voice. The
nose of the male sea-elephant (Macrorhinus proboscideus), when about
three years old, is greatly elongated during the breeding-season, and can
then be erected. In this state it is sometimes a foot in length. The female at
no period of life is thus provided, and her voice is different. That of the
male consists of a wild, hoarse, gurgling noise, which is audible at a great
distance, and is believed to be strengthened by the proboscis. Lesson
compares the erection of the proboscis, to the swelling of the wattles of
male gallinaceous birds, whilst they court the females. In another allied
kind of seal, namely, the bladder-nose (Cystophora cristata), the head is
covered by a great hood or bladder. This is internally supported by the
septum of the nose, which is produced far backwards and rises into a crest
seven inches in height. The hood is clothed with short hair, and is muscular;
it can be inflated until it more than equals the whole head in size! The males
when rutting fight furiously on the ice, and their roaring “is said to be
sometimes so loud as to be heard four miles off.” When attacked by man
they likewise roar or bellow; and whenever irritated the bladder is inflated.
Some naturalists believe that the voice is thus strengthened, but various
other uses have been assigned to this extraordinary structure. Mr. R. Brown
thinks that it serves as a protection against accidents of all kinds. This latter
view is not probable, if what the sealers have long maintained is correct,
namely, that the hood or bladder is very poorly developed in the females
and in the males whilst young.339

Odour.—With some animals, as with the notorious skunk of America, the
overwhelming odour which they emit appears to serve exclusively as a
means of defence. With shrew-mice (Sorex) both sexes possess abdominal
scent-glands, and there can be little doubt, from the manner in which their
bodies are rejected by birds and beasts of prey, that their odour is
protective; nevertheless the glands become enlarged in the males during the
breeding-season. In many quadrupeds the glands are of the same size in
both sexes;340 but their use is not known. In other species the glands are
confined to the males, or are more developed in them than in the females;
and they almost always become more active during the rutting-season. At
this period the glands on the sides of the face of the male elephant enlarge
and emit a secretion having a strong musky odour.

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The rank effluvium of the male goat is well known, and that of certain male
deer is wonderfully strong and persistent. On the banks of the Plata I have
perceived the whole air tainted with the odour of the male Cervus
campestris, at the distance of half a mile to leeward of a herd; and a silk
handkerchief, in which I carried home a skin, though repeatedly used and
washed, retained, when first unfolded, traces of the odour for one year and
seven months. This animal does not emit its strong odour until more than a
year old, and if castrated whilst young never emits it.341 Besides the general
odour, with which the whole body of certain ruminants seems to be
permeated during the breeding-season, many deer, antelopes, sheep, and
goats, possess odoriferous glands in various situations, more especially on
their faces. The so-called tear-sacks or suborbital pits come under this head.
These glands secrete a semi-fluid fetid matter, which is sometimes so
copious as to stain the whole face, as I have seen in the case of an antelope.
They are “usually larger in the male than in the female, and their
development is checked by castration.”342 According to Desmarest they are
altogether absent in the female of Antilope subgutturosa. Hence, there can
be no doubt that they stand in some close relation with the reproductive
functions. They are also sometimes present, and sometimes absent, in
nearly-allied forms. In the adult male musk-deer (Moschus moschiferus), a
naked space round the tail is bedewed with an odoriferous fluid, whilst in
the adult female, and in the male, until two years old, this space is covered
with hair and is not odoriferous. The proper musk-sack, from its position, is
necessarily confined to the male, and forms an additional scent-organ. It is a
singular fact that the matter secreted by this latter gland does not, according
to Pallas, change in consistence, or increase in quantity, during the rutting-
season; nevertheless this naturalist admits that its presence is in some way
connected with the act of reproduction. He gives, however, only a
conjectural and unsatisfactory explanation of its use.343
In most cases, when during the breeding-season the male alone emits a
strong odour, this probably serves to excite or allure the female. We must
not judge on this head by our own taste, for it is well known that rats are
enticed by certain essential oils, and cats by valerian, substances which are
far from agreeable to us; and that dogs, though they will not eat carrion,
sniff and roll in it. From the reasons given when discussing the voice of the
stag, we may reject the idea that the odour serves to bring the females from

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a distance to the males. Active and long-continued use cannot here have
come into play, as in the case of the vocal organs. The odour emitted must
be of considerable importance to the male, inasmuch as large and complex
glands, furnished with muscles for everting the sack, and for closing or
opening the orifice, have in some cases been developed. The development
of these organs is intelligible through sexual selection, if the more
odoriferous males are the most successful in winning the females, and in
leaving offspring to inherit their gradually-perfected glands and odours.

Development of the Hair.—We have seen that male quadrupeds often have
the hair on their necks and shoulders much more developed than in the
females; and many additional instances could be given. This sometimes
serves as a defence to the male during his battles; but whether the hair in
most cases has been specially developed for this purpose is very doubtful.
We may feel almost certain that this is not the case, when a thin and narrow
crest runs along the whole length of the back; for a crest of this kind would
afford scarcely any protection, and the ridge of the back is not a likely place
to be injured; nevertheless such crests are sometimes confined to the males,
or are much more developed in them than in the females. Two antelopes,
the Tragelaphus scriptus344 (see fig. 68, p. 300) and Portax picta, may be
given as instances. The crests of certain stags and of the male wild goat
stand erect, when these animals are enraged or terrified;345 but it can hardly
be supposed that they have been acquired for the sake of exciting fear in
their enemies. One of the above-named antelopes, the Portax picta, has a
large well-defined brush of black hair on the throat, and this is much larger
in the male than in the female. In the Ammotragus tragelaphus of North
Africa, a member of the sheep-family, the front-legs are almost concealed
by an extraordinary growth of hair, which descends from the neck and
upper halves of the legs; but Mr. Bartlett does not believe that this mantle is
of the least use to the male, in whom it is much more developed than in the
female.
Male quadrupeds of many kinds differ from the females in having more
hair, or hair of a different character, on certain parts of their faces. The bull
alone has curled hair on the forehead.346 In three closely-allied sub-genera
of the goat family, the males alone possess beards, sometimes of large size;
in two other sub-genera both sexes have a beard, but this disappears in

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some of the domestic breeds of the common goat; and neither sex of the
Hemitragus has a beard. In the ibex the beard is not developed during the
summer, and is so small at other seasons that it may be called
rudimentary.347 With some monkeys the beard is confined to the male, as in
the Orang, or is much larger in the male than in the female, as in the
Mycetes caraya and Pithecia satanas (fig. 66). So it is with the whiskers of
some species of Macacus,348 and, as we have seen, with the manes of some
species of baboons. But with most kinds of monkeys the various tufts of
hair about the face and head are alike in both sexes.

Fig. 66. Pithecia Satanas, male (from Brehm).

The males of various members of the Ox family (Bovidæ), and of certain
antelopes, are furnished with a dewlap, or great fold of skin on the neck,
which is much less developed in the female.
Now, what must we conclude with respect to such sexual differences as
these? No one will pretend that the beards of certain male-goats, or the
dewlap of the bull, or the crests of hair along the backs of certain male
antelopes, are of any direct or ordinary use to them. It is possible that the
immense beard of the male Pithecia, and the large beard of the male Orang,
may protect their throats when fighting; for the keepers in the Zoological
Gardens inform me that many monkeys attack each other by the throat: but
it is not probable that the beard has been developed for a distinct purpose
from that which the whiskers, moustache, and other tufts of hair on the face
serve; and no one will suppose that these are useful as a protection. Must
we attribute to mere purposeless variability in the male all these appendages

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of hair or skin? It cannot be denied that this is possible; for with many
domesticated quadrupeds, certain characters, apparently not derived through
reversion from any wild parent-form, have appeared in, and are confined to,
the males, or are more largely developed in them than in the females,—for
instance the hump in the male zebu-cattle of India, the tail in fat-tailed
rams, the arched outline of the forehead in the males of several breeds of
sheep, the mane in the ram of an African breed, and, lastly, the mane, long
hairs on the hinder legs, and the dewlap in the male alone of the Berbura
goat.349 The mane which occurs in the rams alone of the above-mentioned
African breed of sheep, is a true secondary sexual character, for it is not
developed, as I hear from Mr. Winwood Reade, if the animal be castrated.
Although we ought to be extremely cautious, as shewn in my work on
‘Variation under Domestication,’ in concluding that any character, even
with animals kept by semi-civilised people, has not been subjected to
selection by man, and thus augmented; yet in the cases just specified this is
improbable, more especially as the characters are confined to the males, or
are more strongly developed in them than in the females. If it were
positively known that the African ram with a mane was descended from the
same primitive stock with the other breeds of sheep, or the Berbura male-
goat with his mane, dewlap, &c., from the same stock with other goats; and
if selection has not been applied to these characters, then they must be due
to simple variability, together with sexually-limited inheritance.
t would appear reasonable to extend the same view to the many analogous
characters occurring in animals under a state of nature. Nevertheless I
cannot persuade myself that this view is applicable in many cases, as in that
of the extraordinary development of hair on the throat and forelegs of the
male Ammotragus, or of the immense beard of the male Pithecia. With
those antelopes in which the male when adult is more strongly-coloured
than the female, and with those monkeys in which this is likewise the case,
and in which the hair on the face is of a different colour from that on the
rest of the head, being arranged in the most diversified and elegant manner,
it seems probable that the crests and tufts of hair have been acquired as
ornaments; and this I know is the opinion of some naturalists. If this view
be correct, there can be little doubt that they have been acquired, or at least
modified, through sexual selection.

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Colour of the Hair and of the Naked Skin.—I will first give briefly all the
cases known to me, of male quadrupeds differing in colour from the
females. With Marsupials, as I am informed by Mr. Gould, the sexes rarely
differ in this respect; but the great red kangaroo offers a striking exception,
“delicate blue being the prevailing tint in those parts of the female, which in
the male are red.”350 In the Didelphis opossum of Cayenne the female is said
to be a little more red than the male. With Rodents Dr. Gray remarks:
“African squirrels, especially those found in the tropical regions, have the
fur much brighter and more vivid at some seasons of the year than at others,
and the fur of the male is generally brighter than that of the female.”351 Dr.
Gray informs me that he specified the African squirrels, because, from their
unusually bright colours, they best exhibit this difference. The female of the
Mus minutus of Russia is of a paler and dirtier tint than the male. In some
few bats the fur of the male is lighter and brighter than in the female.352
The terrestrial Carnivora and Insectivora rarely exhibit sexual differences of
any kind, and their colours are almost always exactly the same in both
sexes. The ocelot (Felis pardalis), however, offers an exception, for the
colours of the female, compared with those of the male, are “moins
apparentes, le fauve étant plus terne, le blanc moins pur, les raies ayant
moins de largeur et les taches moins de diamètre.”353 The sexes of the allied
Felis mitis also differ, but even in a less degree, the general hues of the
female being rather paler than in the male, with the spots less black. The
marine Carnivora or Seals, on the other hand, sometimes differ considerably
in colour, and they present, as we have already seen, other remarkable
sexual differences. Thus the male of the Otaria nigrescens of the southern
hemisphere is of a rich brown shade above; whilst the female, who acquires
her adult tints earlier in life than the male, is dark-grey above, the young of
both sexes being of a very deep chocolate colour. The male of the northern
Phoca groenlandica is tawny grey, with a curious saddle-shaped dark mark
on the back; the female is much smaller, and has a very different
appearance, being “dull white or yellowish straw-colour, with a tawny hue
on the back;” the young at first are pure white, and can “hardly be
distinguished among the icy hummocks and snow, their colour thus acting
as a protection.”354
With Ruminants sexual differences of colour occur more commonly than in
any other order. A difference of this kind is general with the Strepsicerene

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antelopes; thus the male nilghau (Portax picta) is bluish-grey and much
darker than the female, with the square white patch on the throat, the white
marks on the fetlocks, and the black spots on the ears, all much more
distinct. We have seen that in this species the crests and tufts of hair are
likewise more developed in the male than in the hornless female. The male,
as I am informed by Mr. Blyth, without shedding his hair, periodically
becomes darker during the breeding-season. Young males cannot be
distinguished from young females until above twelve months old; and if the
male is emasculated before this period, he never, according to the same
authority, changes colour. The importance of this latter fact, as distinctive of
sexual colouring, becomes obvious, when we hear355 that neither the red
summer-coat nor the blue winter-coat of the Virginian deer is at all affected
by emasculation. With most or all of the highly-ornamented species of
Tragelaphus the males are darker than the hornless females, and their crests
of hair are more fully developed. In the male of that magnificent antelope,
the Derbyan Eland, the body is redder, the whole neck much blacker, and
the white band which separates these colours, broader, than in the female. In
the Cape Eland also, the male is slightly darker than the female.356
In the Indian Black-buck (A. bezoartica), which belongs to another tribe of
antelopes, the male is very dark, almost black; whilst the hornless female is
fawn-coloured. We have in this species, as Mr. Blyth informs me, an exactly
parallel series of facts, as with the Portax picta, namely in the male
periodically changing colour during the breeding season, in the effects of
emasculation on this change, and in the young of both sexes being
undistinguishable from each other. In the Antilope niger the male is black,
the female as well as the young being brown; in A. sing-sing the male is
much brighter coloured than the hornless female, and his chest and belly are
blacker; in the male A. caama, the marks and lines which occur on various
parts of the body are black instead of as in the female brown; in the brindled
gnu (A. gorgon) “the colours of the male are nearly the same as those of the
female, only deeper and of a brighter hue.”357 Other analogous cases could
be added.
The Banteng bull (Bos sondaicus) of the Malayan archipelago is almost
black, with white legs and buttocks; the cow is of a bright dun, as are the
young males until about the age of three years, when they rapidly change
colour. The emasculated bull reverts to the colour of the female. The female

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Kemas goat is paler, and the female Capra ægagrus is said to be more
uniformly tinted than their respective males. Deer rarely present any sexual
differences in colour. Judge Caton, however, informs me that with the males
of the Wapiti deer (Cervus Canadensis) the neck, belly, and legs are much
darker than the same parts in the female; but during the winter the darker
tints gradually fade away and disappear. I may here mention that Judge
Caton has in his park three races of the Virginian deer, which differ slightly
in colour, but the differences are almost exclusively confined to the blue
winter or breeding coat; so that this case may be compared with those given
in a previous chapter of closely-allied or representative species of birds
which differ from each other only in their nuptial plumage.358 The females
of Cervus paludosus of S. America, as well as the young of both sexes, do
not possess the black stripes on the nose, and the blackish-brown line on the
breast which characterise the adult males.359 Lastly, the mature male of the
beautifully coloured and spotted Axis deer is considerably darker, as I am
informed by Mr. Blyth, than the female; and this hue the castrated male
never acquires.
The last Order which we have to consider—for I am not aware that sexual
differences in colour occur in the other mammalian groups—is that of the
Primates. The male of the Lemur macaco is coal-black, whilst the female is
reddish-yellow, but highly variable in colour.360 Of the Quadrumana of the
New World, the females and young of Mycetes caraya are greyish-yellow
and alike; in the second year the young male becomes reddish-brown, in the
third year black, excepting the stomach, which, however, becomes quite
black in the fourth or fifth year. There is also a strongly-marked difference
in colour between the sexes in Mycetes seniculus and Cebus capucinus; the
young of the former and I believe of the latter species resembling the
females. With Pithecia leucocephala the young likewise resemble the
females, which are brownish-black above and light rusty-red beneath, the
adult males being black. The ruff of hair round the face of Ateles
marginatus is tinted yellow in the male and white in the female. Turning to
the Old World, the males of Hylobates hoolock are always black, with the
exception of a white band over the brows; the females vary from whity-
brown to a dark tint mixed with black, but are never wholly black.361 In the
beautiful Cercopithecus diana the head of the adult male is of an intense
black, whilst that of the female is dark grey; in the former the fur between

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the thighs is of an elegant fawn-colour, in the latter it is paler. In the equally
beautiful and curious moustache monkey (Cercopithecus cephus) the only
difference between the sexes is that the tail of the male is chesnut and that
of the female grey; but Mr. Bartlett informs me that all the hues become
more strongly pronounced in the male when adult, whilst in the female they
remain as they were during youth. According to the coloured figures given
by Solomon Müller, the male of Semnopithecus chrysomelas is nearly
black, the female being pale brown. In the Cercopithecus cynosurus and
griseo-viridis one part of the body which is confined to the male sex is of
the most brilliant blue or green, and contrasts strikingly with the naked skin
on the hinder part of the body, which is vivid red.

Fig. 67. Head of male Mandrill (from Gervais, ‘Hist. Nat
des Mammifères’).

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Lastly, in the Baboon family, the adult male of Cynocephalus hamadryas
differs from the female not only by his immense mane, but slightly in the
colour of the hair and of the naked callosities. In the drill (Cynocephalus
leucophœus) the females and young are much paler-coloured, with less
green, than the adult males. No other member of the whole class of
mammals is coloured in so extraordinary a manner as the adult male
mandrill (Cynocephalus mormon). The face at this age becomes of a fine
blue, with the ridge and tip of the nose of the most brilliant red. According
to some authors the face is also marked with whitish stripes, and is shaded
in parts with black, but the colours appear to be variable. On the forehead
there is a crest of hair, and on the chin a yellow beard. “Toutes les parties
supérieures de leurs cuisses et le grand espace nu de leurs fesses sont
également colorés du rouge le plus vif, avec un mélange de bleu qui ne
manque réellement pas d’élégance.”362 When the animal is excited all the
naked parts become much more vividly tinted. Several authors have used
the strongest expressions in describing these resplendent colours, which
they compare with those of the most brilliant birds. Another most
remarkable peculiarity is that when the great canine teeth are fully
developed, immense protuberances of bone are formed on each cheek,
which are deeply furrowed longitudinally, and the naked skin over them is
brilliantly-coloured, as just described. (Fig. 67.) In the adult females and in
the young of both sexes these protuberances are scarcely perceptible; and
the naked parts are much less brightly coloured, the face being almost
black, tinged with blue. In the adult female, however, the nose at certain
regular intervals of time becomes tinted with red.
In all the cases hitherto given the male is more strongly or brightly coloured
than the female, and differs in a greater degree from the young of both
sexes. But as a reversed style of colouring is characteristic of the two sexes
with some few birds, so with the Rhesus monkey (Macacus rhesus) the
female has a large surface of naked skin round the tail, of a brilliant carmine
red, which periodically becomes, as I was assured by the keepers in the
Zoological Gardens, even more vivid, and her face is also pale red. On the
other hand with the adult male and with the young of both sexes, as I saw in
the Gardens, neither the naked skin at the posterior end of the body, nor the
face, shew a trace of red. It appears, however, from some published
accounts, that the male does occasionally, or during certain seasons, exhibit

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some traces of the red. Although he is thus less ornamented than the female,
yet in the larger size of his body, larger canine teeth, more developed
whiskers, more prominent superciliary ridges, he follows the common rule
of the male excelling the female.

I have now given all the cases known to me of a difference in colour
between the sexes of mammals. The colours of the female either do not
differ in a sufficient degree from those of the male, or are not of a suitable
nature, to afford her protection, and therefore cannot be explained on this
principle. In some, perhaps in many cases, the differences may be the result
of variations confined to one sex and transmitted to the same sex, without
any good having been thus gained, and therefore without the aid of
selection. We have instances of this kind with our domesticated animals, as
in the males of certain cats being rusty-red, whilst the females are tortoise-
shell coloured. Analogous cases occur under nature; Mr. Bartlett has seen
many black varieties of the jaguar, leopard, vulpine phalanger and wombat;
and he is certain that all, or nearly all, were males. On the other hand, both
sexes of wolves, foxes, and apparently of American squirrels, are
occasionally born black. Hence it is quite possible that with some mammals
the blackness of the males, especially when this colour is congenital, may
simply be the result, without the aid of selection, of one or more variations
having occurred, which from the first were sexually limited in their
transmission. Nevertheless it can hardly be admitted that the diversified,
vivid, and contrasted colours of certain quadrupeds, for instance of the
above-mentioned monkeys and antelopes, can thus be accounted for. We
should bear in mind that these colours do not appear in the male at birth, as
in the case of most ordinary variations, but only at or near maturity; and that
unlike ordinary variations, if the male be emasculated, they never appear or
subsequently disappear. It is on the whole a much more probable conclusion
that the strongly-marked colours and other ornamental characters of male
quadrupeds are beneficial to them in their rivalry with other males, and
have consequently been acquired through sexual selection. The probability
of this view is strengthened by the differences in colour between the sexes
occurring almost exclusively, as may be observed by going through the
previous details, in those groups and subgroups of mammals, which present
other and distinct secondary sexual characters; these being likewise due to
the action of sexual selection.

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Quadrupeds manifestly take notice of colour. Sir S. Baker repeatedly
observed that the African elephant and rhinoceros attacked with special fury
white or grey horses. I have elsewhere shewn363 that half-wild horses
apparently prefer pairing with those of the same colour, and that herds of
fallow-deer of a different colour, though living together, have long kept
distinct. It is a more significant fact that a female zebra would not admit the
addresses of a male ass until he was painted so as to resemble a zebra, and
then, as John Hunter remarks, “she received him very readily. In this
curious fact, we have instinct excited by mere colour, which had so strong
an effect as to get the better of everything else. But the male did not require
this, the female being an animal somewhat similar to himself, was sufficient
to rouse him.”364
In an early chapter we have seen that the mental powers of the higher
animals do not differ in kind, though so greatly in degree, from the
corresponding powers of man, especially of the lower and barbarous races;
and it would appear that even their taste for the beautiful is not widely
different from that of the Quadrumana. As the negro of Africa raises the
flesh on his face into parallel ridges “or cicatrices, high above the natural
surface, which unsightly deformities, are considered great personal
attractions;”365—as negroes, as well as savages in many parts of the world,
paint their faces with red, blue, white, or black bars,—so the male mandrill
of Africa appears to have acquired his deeply-furrowed and gaudily-
coloured face from having been thus rendered attractive to the female. No
doubt it is to us a most grotesque notion that the posterior end of the body
should have been coloured for the sake of ornament even more brilliantly
than the face; but this is really not more strange than that the tails of many
birds should have been especially decorated.
e do not at present possess any evidence that the males take pains to display
their charms before the female; and the elaborate manner in which this is
performed by male birds, is the strongest argument in favour of the belief
that the females admire, or are excited by, the ornaments and colours
displayed before them. There is, however, a striking parallelism between
mammals and birds in all their secondary sexual characters, namely in their
weapons for fighting with rival males, in their ornamental appendages, and
in their colours. In both classes, when the male differs from the female, the
young of both sexes almost always resemble each other, and in a large

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majority of cases resemble the adult female. In both classes the male
assumes the characters proper to his sex shortly before the age for
reproduction; if emasculated he either never acquires such characters or
subsequently loses them. In both classes the change of colour is sometimes
seasonal, and the tints of the naked parts sometimes become more vivid
during the act of courtship. In both classes the male is almost always more
vividly or strongly coloured than the female, and is ornamented with larger
crests either of hair or feathers, or other appendages. In a few exceptional
cases the female in both classes is more highly ornamented than the male.
With many mammals, and at least in the case of one bird, the male is more
odoriferous than the female. In both classes the voice of the male is more
powerful than that of the female. Considering this parallelism there can be
little doubt that the same cause, whatever it may be, has acted on mammals
and birds; and the result, as far as ornamental characters are concerned, may
safely be attributed, as it appears to me, to the long-continued preference of
the individuals of one sex for certain individuals of the opposite sex,
combined with their success in leaving a larger number of offspring to
inherit their superior attractions.

Equal transmission of ornamental characters to both sexes.—With many
birds, ornaments, which analogy leads us to believe were primarily acquired
by the males, have been transmitted equally, or almost equally, to both
sexes; and we may now enquire how far this view may be extended to
mammals. With a considerable number of species, especially the smaller
kinds, both sexes have been coloured, independently of sexual selection, for
the sake of protection; but not, as far as I can judge, in so many cases, nor
in nearly so striking a manner as in most of the lower classes. Audubon
remarks that he often mistook the musk-rat,366 whilst sitting on the banks of
a muddy stream, for a clod of earth, so complete was the resemblance. The
hare on her form is a familiar instance of concealment through colour; yet
this principle partly fails in a closely-allied species, namely the rabbit, for
as this animal runs to its burrow, it is made conspicuous to the sportsman
and no doubt to all beasts of prey, by its upturned pure-white tail. No one
has ever doubted that the quadrupeds which inhabit snow-clad regions, have
been rendered white to protect them from their enemies, or to favour their
stealing on their prey. In regions where snow never lies long on the ground
a white coat would be injurious; consequently species thus coloured are

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extremely rare in the hotter parts of the world. It deserves notice that many
quadrupeds, inhabiting moderately cold regions, although they do not
assume a white winter dress, become paler during this season; and this
apparently is the direct result of the conditions to which they have long
been exposed. Pallas367 states that in Siberia a change of this nature occurs
with the wolf, two species of Mustela, the domestic horse, the Equus
hemionus, the domestic cow, two species of antelopes, the musk-deer, the
roe, the elk, and reindeer. The roe, for instance, has a red summer and a
greyish-white winter coat; and the latter may perhaps serve as a protection
to the animal whilst wandering through the leafless thickets, sprinkled with
snow and hoar-frost. If the above named animals were gradually to extend
their range into regions perpetually covered with snow, their pale winter-
coats would probably be rendered, through natural selection, whiter and
whiter by degrees, until they became as white as snow.

Fig. 68. Tragelaphus scriptus, male (from the Knowsley Menagerie).

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Fig. 69. Damalis pygarga, male (from the Knowsley
Menagerie).

Although we must admit that many quadrupeds have received their present
tints as a protection, yet with a host of species, the colours are far too
conspicuous and too singularly arranged to allow us to suppose that they
serve for this purpose. We may take as an illustration certain antelopes:
when we see that the square white patch on the throat, the white marks on
the fetlocks, and the round black spots on the ears, are all more distinct in
the male of the Portax picta, than in the female;—when we see that the
colours are more vivid, that the narrow white lines on the flank and the
broad white bar on the shoulder are more distinct in the male Oreas
derbyanus than in the female;—when we see a similar difference between
the sexes of the curiously-ornamented Tragelaphus scriptus (fig. 68),—we
may conclude that these colours and various marks have been at least
intensified through sexual selection. It is inconceivable that such colours
and marks can be of any direct or ordinary service to these animals; and as
they have almost certainly been intensified through sexual selection, it is

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probable that they were originally gained through this same process, and
then partially transferred to the females. If this view be admitted, there can
be little doubt that the equally singular colours and marks of many other
antelopes, though common to both sexes, have been gained and transmitted
in a like manner. Both sexes, for instance, of the Koodoo (Strepsiceros
Kudu, fig. 62) have narrow white vertical lines on their hinder flanks, and
an elegant angular white mark on their foreheads. Both sexes in the genus
Damalis are very oddly coloured; in D. pygarga the back and neck are
purplish-red, shading on the flanks into black, and abruptly separated from
the white belly and a large white space on the buttocks; the head is still
more oddly coloured, a large oblong white mask, narrowly-edged with
black, covers the face up to the eyes (fig. 69); there are three white stripes
on the forehead, and the ears are marked with white. The fawns of this
species are of a uniform pale yellowish-brown. In Damalis albifrons the
colouring of the head differs from that in the last species in a single white
stripe replacing the three stripes, and in the ears being almost wholly
white.368 After having studied to the best of my ability the sexual differences
of animals belonging to all classes, I cannot avoid the conclusion that the
curiously-arranged colours of many antelopes, though common to both
sexes, are the result of sexual selection primarily applied to the male.
The same conclusion may perhaps be extended to the tiger, one of the most
beautiful animals in the world, the sexes of which cannot be distinguished
by colour, even by the dealers in wild beasts. Mr. Wallace believes369 that
the striped coat of the tiger “so assimilates with the vertical stems of the
bamboo, as to assist greatly in concealing him from his approaching prey.”
But this view does not appear to me satisfactory. We have some slight
evidence that his beauty may be due to sexual selection, for in two species
of Felis analogous marks and colours are rather brighter in the male than in
the female. The zebra is conspicuously striped, and stripes on the open
plains of South Africa cannot afford any protection. Burchell370 in
describing a herd says, “their sleek ribs glistened in the sun, and the
brightness and regularity of their striped coats presented a picture of
extraordinary beauty, in which probably they are not surpassed by any other
quadruped.” Here we have no evidence of sexual selection, as throughout
the whole group of the Equidæ the sexes are identical in colour.
Nevertheless he who attributes the white and dark vertical stripes on the

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flanks of various antelopes to sexual selection, will probably extend the
same view to the Royal Tiger and beautiful Zebra.
We have seen in a former chapter that when young animals belonging to
any class follow nearly the same habits of life with their parents, and yet are
coloured in a different manner, it may be inferred that they have retained
the colouring of some ancient and extinct progenitor. In the family of pigs,
and in the genus Tapir, the young are marked with longitudinal stripes, and
thus differ from every existing adult species in these two groups. With
many kinds of deer the young are marked with elegant white spots, of
which their parents exhibit not a trace. A graduated series can be followed
from the Axis deer, both sexes of which at all ages and during all seasons
are beautifully spotted (the male being rather more strongly coloured than
the female)—to species in which neither the old nor the young are spotted. I
will specify some of the steps in this series. The Mantchurian deer (Cervus
Mantchuricus) is spotted during the whole year, but the spots are much
plainer, as I have seen in the Zoological Gardens, during the summer, when
the general colour of the coat is lighter, than during the winter, when the
general colour is darker and the horns are fully developed. In the hog-deer
(Hyelaphus porcinus) the spots are extremely conspicuous during the
summer when the coat is reddish-brown, but quite disappear during the
winter when the coat is brown.371 In both these species the young are
spotted. In the Virginian deer the young are likewise spotted, and about five
per cent. of the adult animals living in Judge Caton’s park, as I am informed
by him, temporarily exhibit at the period when the red summer coat is being
replaced by the bluish winter coat, a row of spots on each flank, which are
always the same in number, though very variable in distinctness. From this
condition there is but a very small step to the complete absence of spots at
all seasons in the adults; and lastly, to their absence at all ages, as occurs
with certain species. From the existence of this perfect series, and more
especially from the fawns of so many species being spotted, we may
conclude that the now living members of the deer family are the
descendants of some ancient species which, like the Axis deer, was spotted
at all ages and seasons. A still more ancient progenitor probably resembled
to a certain extent the Hyomoschus aquaticus—for this animal is spotted,
and the hornless males have large exserted canine teeth, of which some few
true deer still retain rudiments. It offers, also, one of those interesting cases

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of a form linking together two groups, as it is intermediate in certain
osteological characters between the pachyderms and ruminants, which were
formerly thought to be quite distinct.372
A curious difficulty here arises. If we admit that coloured spots and stripes
have been acquired as ornaments, how comes it that so many existing deer,
the descendants of an aboriginally spotted animal, and all the species of
pigs and tapirs, the descendants of an aboriginally striped animal, have lost
in their adult state their former ornaments? I cannot satisfactorily answer
this question. We may feel nearly sure that the spots and stripes disappeared
in the progenitors of our existing species at or near maturity, so that they
were retained by the young and, owing to the law of inheritance at
corresponding ages, by the young of all succeeding generations. It may
have been a great advantage to the lion and puma from the open nature of
the localities which they commonly haunt, to have lost their stripes, and to
have been thus rendered less conspicuous to their prey; and if the
successive variations, by which this end was gained, occurred rather late in
life, the young would have retained their stripes, as we know to be the case.
In regard to deer, pigs, and tapirs, Fritz Müller has suggested to me that
these animals by the removal through natural selection of their spots or
stripes would have been less easily seen by their enemies; and they would
have especially required this protection, as soon as the carnivora increased
in size and number during the Tertiary periods. This may be the true
explanation, but it is rather strange that the young should not have been
equally well protected, and still more strange that with some species the
adults should have retained their spots, either partially or completely, during
part of the year. We know, though we cannot explain the cause, that when
the domestic ass varies and becomes reddish-brown, grey or black, the
stripes on the shoulders and even on the spine frequently disappear. Very
few horses, except dun-coloured kinds, exhibit stripes on any part of their
bodies, yet we have good reason to believe that the aboriginal horse was
striped on the legs and spine, and probably on the shoulders.373 Hence the
disappearance of the spots and stripes in our adult existing deer, pigs, and
tapirs, may be due to a change in the general colour of their coats; but
whether this change was effected through sexual or natural selection, or was
due to the direct action of the conditions of life, or some other unknown
cause, it is impossible to decide. An observation made by Mr. Sclater well

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illustrates our ignorance of the laws which regulate the appearance and
disappearance of stripes; the species of Asinus which inhabit the Asiatic
continent are destitute of stripes, not having even the cross shoulder-stripe,
whilst those which inhabit Africa are conspicuously striped, with the partial
exception of A. tæniopus, which has only the cross shoulder-stripe and
generally some faint bars on the legs; and this species inhabits the almost
intermediate region of Upper Egypt and Abyssinia.374

Fig. 70. Head of Semnopithecus rubicundus. This and the
following figures (from Prof. Gervais) are given to shew
the odd arrangement and development of the hair on the
head.

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Fig. 71. Head of Semnopithecus comatus. Fig. 72. Head of Cebus capucinus.

Fig. 73. Head of Ateles marginatus. Fig. 74. Head of Cebus vellerosus.

Quadrumana.—Before we conclude, it will be advisable to add a few
remarks to those already given on the ornamental characters of monkeys. In
most of the species the sexes resemble each other in colour, but in some, as
we have seen, the males differ from the females, especially in the colour of
the naked parts of the skin, in the development of the beard, whiskers, and
mane. Many species are coloured either in so extraordinary or beautiful a
manner, and are furnished with such curious and elegant crests of hair, that
we can hardly avoid looking at these characters as having been gained for

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the sake of ornament. The accompanying figures (figs. 70 to 74) serve to
shew the arrangement of the hair on the face and head in several species. It
is scarcely conceivable that these crests of hair and the strongly-contrasted
colours of the fur and skin can be the result of mere variability without the
aid of selection; and it is inconceivable that they can be of any ordinary use
to these animals. If so, they have probably been gained through sexual
selection, though transmitted equally, or almost equally, to both sexes. With
many of the Quadrumana, we have additional evidence of the action of
sexual selection in the greater size and strength of the males, and in the
greater development of their canine teeth, in comparison with the females.
With respect to the strange manner in which both sexes of some species are
coloured, and of the beauty of others, a few instances will suffice. The face
of the Cercopithecus petaurista (fig. 75) is black, the whiskers and beard
being white, with a defined, round, white spot on the nose, covered with
short white hair, which gives to the animal an almost ludicrous aspect. The
Semnopithecus frontatus likewise, has a blackish face with a long black
beard, and a large naked spot on the forehead of a bluish-white colour. The
face of Macacus lasiotus is dirty flesh-coloured, with a defined red spot on
each cheek. The appearance of Cercocebus æthiops is grotesque, with its
black face, white whiskers and collar, chesnut head, and a large naked white
spot over each eyelid. In very many species, the beard, whiskers, and crests
of hair round the face are of a different colour from the rest of the head, and
when different, are always of a lighter tint,375 being often pure white,
sometimes bright yellow, or reddish. The whole face of the South American
Brachyurus calvus is of a “glowing scarlet hue;” but this colour does not
appear until the animal is nearly mature.376 The naked skin of the face
differs wonderfully in colour in the various species. It is often brown or
flesh-colour, with parts perfectly white, and often as black as that of the
most sooty negro. In the Brachyurus the scarlet tint is brighter than that of
the most blushing Caucasian damsel. It is sometimes more distinctly orange
than in any Mongolian, and in several species it is blue, passing into violet
or grey. In all the species known to Mr. Bartlett, in which the adults of both
sexes have strongly-coloured faces, the colours are dull or absent during
early youth. This likewise holds good with the Mandrill and Rhesus, in
which the face and the posterior parts of the body are brilliantly coloured in
one sex alone. In these latter cases we have every reason to believe that the

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colours were acquired through sexual selection; and we are naturally led to
extend the same view to the foregoing species, though both sexes when
adult have their faces coloured in the same manner.

Fig. 75. Cercopithecus petaurista (from Brehm)

Although, according to our taste, many kinds of monkeys are far from
beautiful, other species are universally admired for their elegant appearance
and bright colours. The Semnopithecus nemæus, though peculiarly
coloured, is described as extremely pretty; the orange-tinted face is
surrounded by long whiskers of glossy whiteness, with a line of chesnut-red
over the eyebrows; the fur on the back is of a delicate grey, with a square
patch on the loins, the tail and the fore-arms all of a pure white; a gorget of
chesnut surmounts the chest; the hind thighs are black, with the legs
chesnut-red. I will mention only two other monkeys on account of their
beauty; and I have selected these as they present slight sexual differences in
colour, which renders it in some degree probable that both sexes owe their
elegant appearance to sexual selection. In the moustache-monkey
(Cercopithecus cephus) the general colour of the fur is mottled-greenish,
with the throat white; in the male the end of the tail is chesnut; but the face
is the most ornamented part, the skin being chiefly bluish-grey, shading into
a blackish tint beneath the eyes, with the upper lip of a delicate blue,
clothed on the lower edge with a thin black moustache; the whiskers are
orange-coloured, with the upper part black, forming a band which extends
backwards to the ears, the latter being clothed with whitish hairs. In the

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Zoological Society’s Gardens I have often overheard visitors admiring the
beauty of another monkey, deservedly called Cercopithecus Diana (fig. 76);
the general colour of the fur is grey; the chest and inner surface of the
forelegs are white; a large triangular defined space on the hinder part of the
back is rich chesnut; in the male the inner sides of the thighs and the
abdomen are delicate fawn-coloured, and the top of the head is black; the
face and ears are intensely black, finely contrasted with a white transverse
crest over the eyebrows and with a long white peaked beard, of which the
basal portion is black.377

Fig. 76. Cercopithecus Diana (from Brehm).

In these and many other monkeys, the beauty and singular arrangement of
their colours, and still more the diversified and elegant arrangement of the
crests and tufts of hair on their heads, force the conviction on my mind that
these characters have been acquired through sexual selection exclusively as
ornaments.

Summary.—The law of battle for the possession of the female appears to
prevail throughout the whole great class of mammals. Most naturalists will
admit that the greater size, strength, courage, and pugnacity of the male, his
special weapons of offence, as well as his special means of defence, have
all been acquired or modified through that form of selection which I have
called sexual selection. This does not depend on any superiority in the
general struggle for life, but on certain individuals of one sex, generally the

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male sex, having been successful in conquering other males, and on their
having left a larger number of offspring to inherit their superiority, than the
less successful males.
There is another and more peaceful kind of contest, in which the males
endeavour to excite or allure the females by various charms. This may be
effected by the powerful odours emitted by the males during the breeding-
season; the odoriferous glands having been acquired through sexual
selection. Whether the same view can be extended to the voice is doubtful,
for the vocal organs of the males may have been strengthened by use during
maturity, under the powerful excitements of love, jealousy, or rage, and
transmitted to the same sex. Various crests, tufts, and mantles of hair, which
are either confined to the male, or are more developed in this sex than in the
females, seem in most cases to be merely ornamental, though they
sometimes serve as a defence against rival males. There is even reason to
suspect that the branching horns of stags, and the elegant horns of certain
antelopes, though properly serving as weapons of offence or of defence,
have been partly modified for the sake of ornament.
When the male differs in colour from the female he generally exhibits
darker and more strongly-contrasted tints. We do not in this class meet with
the splendid red, blue, yellow, and green colours, so common with male
birds and many other animals. The naked parts, however, of certain
Quadrumana must be excepted; for such parts, often oddly situated, are
coloured in some species in the most brilliant manner. The colours of the
male in other cases may be due to simple variation, without the aid of
selection. But when the colours are diversified and strongly pronounced,
when they are not developed until near maturity, and when they are lost
after emasculation, we can hardly avoid the conclusion that they have been
acquired through sexual selection for the sake of ornament, and have been
transmitted exclusively, or almost exclusively, to the same sex. When both
sexes are coloured in the same manner, and the colours are conspicuous or
curiously arranged, without being of the least apparent use as a protection,
and especially when they are associated with various other ornamental
appendages, we are led by analogy to the same conclusion, namely, that
they have been acquired through sexual selection, although transmitted to
both sexes. That conspicuous and diversified colours, whether confined to
the males or common to both sexes, are as a general rule associated in the

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same groups and subgroups with other secondary sexual characters, serving
for war or for ornament, will be found to hold good if we look back to the
various cases given in this and the last chapter.
The law of the equal transmission of characters to both sexes, as far as
colour and other ornaments are concerned, has prevailed far more
extensively with mammals than with birds; but in regard to weapons, such
as horns and tusks, these have often been transmitted either exclusively, or
in a much higher degree to the males than to the females. This is a
surprising circumstance, for as the males generally use their weapons as a
defence against enemies of all kinds, these weapons would have been of
service to the female. Their absence in this sex can be accounted for, as far
as we can see, only by the form of inheritance which has prevailed. Finally
with quadrupeds the contest between the individuals of the same sex,
whether peaceful or bloody, has with the rarest exceptions been confined to
the males; so that these have been modified through sexual selection, either
for fighting with each other or for alluring the opposite sex, far more
commonly than the females.

CHAPTER XIX.

Secondary Sexual Characters of Man.
Differences between man and woman—Causes of such differences and of certain characters common
to both sexes—Law of battle—Differences in mental powers—and voice—On the influence of
beauty in determining the marriages of mankind—Attention paid by savages to ornaments—
Their ideas of beauty in woman—The tendency to exaggerate each natural peculiarity.

With mankind the differences between the sexes are greater than in most
species of Quadrumana, but not so great as in some, for instance, the
mandrill. Man on an average is considerably taller, heavier, and stronger
than woman, with squarer shoulders and more plainly-pronounced muscles.
Owing to the relation which exists between muscular development and the
projection of the brows,378 the superciliary ridge is generally more strongly
marked in man than in woman. His body, and especially his face, is more
hairy, and his voice has a different and more powerful tone. In certain tribes
the women are said, whether truly I know not, to differ slightly in tint from

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the men; and with Europeans, the women are perhaps the more brightly
coloured of the two, as may be seen when both sexes have been equally
exposed to the weather.
Man is more courageous, pugnacious, and energetic than woman, and has a
more inventive genius. His brain is absolutely larger, but whether relatively
to the larger size of his body, in comparison with that of woman, has not, I
believe been fully ascertained. In woman the face is rounder; the jaws and
the base of the skull smaller; the outlines of her body rounder, in parts more
prominent; and her pelvis is broader than in man;379 but this latter character
may perhaps be considered rather as a primary than a secondary sexual
character. She comes to maturity at an earlier age than man.
As with animals of all classes, so with man, the distinctive characters of the
male sex are not fully developed until he is nearly mature; and if
emasculated they never appear. The beard, for instance, is a secondary
sexual character, and male children are beardless, though at an early age
they have abundant hair on their heads. It is probably due to the rather late
appearance in life of the successive variations, by which man acquired his
masculine characters, that they are transmitted to the male sex alone. Male
and female children resemble each other closely, like the young of so many
other animals in which the adult sexes differ; they likewise resemble the
mature female much more closely, than the mature male. The female,
however, ultimately assumes certain distinctive characters, and in the
formation of her skull, is said to be intermediate between the child and the
man.380 Again, as the young of closely allied though distinct species do not
differ nearly so much from each other as do the adults, so it is with the
children of the different races of man. Some have even maintained that
race-differences cannot be detected in the infantile skull.381 In regard to
colour, the newborn negro child is reddish nut-brown, which soon becomes
slaty-grey; the black colour being fully developed within a year in the
Sudan, but not until three years in Egypt. The eyes of the negro are at first
blue, and the hair chesnut-brown rather than black, being curled only at the
ends. The children of the Australians immediately after birth are yellowish-
brown, and become dark at a later age. Those of the Guaranys of Paraguay
are whitish-yellow, but they acquire in the course of a few weeks the
yellowish-brown tint of their parents. Similar observations have been made
in other parts of America.382

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I have specified the foregoing familiar differences between the male and
female sex in mankind, because they are curiously the same as in the
Quadrumana. With these animals the female is mature at an earlier age than
the male; at least this is certainly the case with the Cebus azaræ.383 With
most of the species the males are larger and much stronger than the females,
of which fact the gorilla offers a well-known instance. Even in so trifling a
character as the greater prominence of the superciliary ridge, the males of
certain monkeys differ from the females,384 and agree in this respect with
mankind. In the gorilla and certain other monkeys, the cranium of the adult
male presents a strongly-marked sagittal crest, which is absent in the
female; and Ecker found a trace of a similar difference between the two
sexes in the Australians.385 With monkeys when there is any difference in
the voice, that of the male is the more powerful. We have seen that certain
male monkeys, have a well-developed beard, which is quite deficient, or
much less developed in the female. No instance is known of the beard,
whiskers, or moustache being larger in a female than in the male monkey.
Even in the colour of the beard there is a curious parallelism between man
and the Quadrumana, for when in man the beard differs in colour from the
hair of the head, as is often the case, it is, I believe, invariably of a lighter
tint, being often reddish. I have observed this fact in England, and Dr.
Hooker, who attended to this little point for me in Russia, found no
exception to the rule. In Calcutta, Mr. J. Scott, of the Botanic Gardens, was
so kind as to observe with care the many races of men to be seen there, as
well as in some other parts of India, namely, two races in Sikhim, the
Bhoteas, Hindoos, Burmese, and Chinese. Although most of these races
have very little hair on the face, yet he always found that when there was
any difference in colour between the hair of the head and the beard, the
latter was invariably of a lighter tint. Now with monkeys, as has already
been stated, the beard frequently differs in a striking manner in colour from
the hair of the head, and in such cases it is invariably of a lighter hue, being
often pure white, sometimes yellow or reddish.386
In regard to the general hairiness of the body, the women in all races are
less hairy than the men, and in some few Quadrumana the under side of the
body of the female is less hairy than that of the male.387 Lastly, male
monkeys, like men, are bolder and fiercer than the females. They lead the
troop, and when there is danger, come to the front. We thus see how close is

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the parallelism between the sexual differences of man and the Quadrumana.
With some few species, however, as with certain baboons, the gorilla and
orang, there is a considerably greater difference between the sexes, in the
size of the canine teeth, in the development and colour of the hair, and
especially in the colour of the naked parts of the skin, than in the case of
mankind.
The secondary sexual characters of man are all highly variable, even within
the limits of the same race or sub-species; and they differ much in the
several races. These two rules generally hold good throughout the animal
kingdom. In the excellent observations made on board the Novara,388 the
male Australians were found to exceed the females by only 65 millim. in
height, whilst with the Javanese the average excess was 218 millim., so that
in this latter race the difference in height between the sexes is more than
thrice as great as with the Australians. The numerous measurements of
various other races, with respect to stature, the circumference of the neck
and chest, and the length of the back-bone and arms, which were carefully
made, nearly all shewed that the males differed much more from each other
than did the females. This fact indicates that, as far as these characters are
concerned, it is the male which has been chiefly modified, since the races
diverged from their common and primeval source.
The development of the beard and the hairiness of the body differ
remarkably in the men belonging to distinct races, and even to different
families in the same race. We Europeans see this amongst ourselves. In the
island of St. Kilda, according to Martin,389 the men do not acquire beards,
which are very thin, until the age of thirty or upwards. On the Europæo-
Asiatic continent, beards prevail until we pass beyond India, though with
the natives of Ceylon they are frequently absent, as was noticed in ancient
times by Diodorus.390 Beyond India beards disappear, as with the Siamese,
Malays, Kalmucks, Chinese, and Japanese; nevertheless the Ainos,391 who
inhabit the northernmost islands of the Japan archipelago, are the most
hairy men in the world. With negroes the beard is scanty or absent, and they
have no whiskers; in both sexes the body is almost destitute of fine down.392
On the other hand, the Papuans of the Malay archipelago, who are nearly as
black as negroes, possess well-developed beards.393 In the Pacific Ocean the
inhabitants of the Fiji archipelago have large bushy beards, whilst those of
the not-distant archipelagoes of Tonga and Samoa are beardless; but these

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men belong to distinct races. In the Ellice group all the inhabitants belong
to the same race; yet on one island alone, namely Nunemaya, “the men have
splendid beards;” whilst on the other islands “they have, as a rule, a dozen
straggling hairs for a beard.”394
Throughout the great American continent the men may be said to be
beardless; but in almost all the tribes a few short hairs are apt to appear on
the face, especially during old age. With the tribes of North America, Catlin
estimates that eighteen out of twenty men are completely destitute by nature
of a beard; but occasionally there may be seen a man, who has neglected to
pluck out the hairs at puberty, with a soft beard an inch or two in length.
The Guaranys of Paraguay differ from all the surrounding tribes in having a
small beard, and even some hair on the body, but no whiskers.395 I am
informed by Mr. D. Forbes, who particularly attended to this subject, that
the Aymaras and Quechuas of the Cordillera are remarkably hairless, yet in
old age a few straggling hairs occasionally appear on the chin. The men of
these two tribes have very little hair on the various parts of the body where
hair grows abundantly in Europeans, and the women have none on the
corresponding parts. The hair on the head, however, attains an extraordinary
length in both sexes, often reaching almost to the ground; and this is
likewise the case with some of the N. American tribes. In the amount of
hair, and in the general shape of the body, the sexes of the American
aborigines do not differ from each other so much as with most other races
of mankind.396 This fact is analogous with what occurs with some allied
monkeys; thus the sexes of the chimpanzee are not as different as those of
the gorilla or orang.397
In the previous chapters we have seen that with mammals, birds, fishes,
insects, &c., many characters, which there is every reason to believe were
primarily gained through sexual selection by one sex alone, have been
transferred to both sexes. As this same form of transmission has apparently
prevailed to a large extent with mankind, it will save much useless
repetition if we consider the characters peculiar to the male sex together
with certain other characters common to both sexes.

Law of Battle.—With barbarous nations, for instance with the Australians,
the women are the constant cause of war both between the individuals of

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the same tribe and between distinct tribes. So no doubt it was in ancient
times; “nam fuit ante Helenam mulier teterrima belli causa.” With the North
American Indians, the contest is reduced to a system. That excellent
observer, Hearne,398 says:—“It has ever been the custom among these
people for the men to wrestle for any woman to whom they are attached;
and, of course, the strongest party always carries off the prize. A weak man,
unless he be a good hunter, and well-beloved, is seldom permitted to keep a
wife that a stronger man thinks worth his notice. This custom prevails
throughout all the tribes, and causes a great spirit of emulation among their
youth, who are upon all occasions, from their childhood, trying their
strength and skill in wrestling.” With the Guanas of South America, Azara
states that the men rarely marry till twenty or more years old, as before that
age they cannot conquer their rivals.
Other similar facts could be given; but even if we had no evidence on this
head, we might feel almost sure, from the analogy of the higher
Quadrumana,399 that the law of battle had prevailed with man during the
early stages of his development. The occasional appearance at the present
day of canine teeth which project above the others, with traces of a
diastema or open space for the reception of the opposite canines, is in all
probability a case of reversion to a former state, when the progenitors of
man were provided with these weapons, like so many existing male
Quadrumana. It was remarked in a former chapter that as man gradually
became erect, and continually used his hands and arms for fighting with
sticks and stones, as well as for the other purposes of life, he would have
used his jaws and teeth less and less. The jaws, together with their muscles,
would then have become reduced through disuse, as would the teeth
through the not well understood principles of correlation and the economy
of growth; for we everywhere see that parts which are no longer of service
are reduced in size. By such steps the original inequality between the jaws
and teeth in the two sexes of mankind would ultimately have been quite
obliterated. The case is almost parallel with that of many male Ruminants,
in which the canine teeth have been reduced to mere rudiments, or have
disappeared, apparently in consequence of the development of horns. As the
prodigious difference between the skulls of the two sexes in the Gorilla and
Orang, stands in close relation with the development of the immense canine
teeth in the males, we may infer that the reduction of the jaws and teeth in

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the early male progenitors of man led to a most striking and favourable
change in his appearance.
There can be little doubt that the greater size and strength of man, in
comparison with woman, together with his broader shoulders, more
developed muscles, rugged outline of body, his greater courage and
pugnacity, are all due in chief part to inheritance from some early male
progenitor, who, like the existing anthropoid apes, was thus characterised.
These characters will, however, have been preserved or even augmented
during the long ages whilst man was still in a barbarous condition, by the
strongest and boldest men having succeeded best in the general struggle for
life, as well as in securing wives, and thus having left a large number of
offspring. It is not probable that the greater strength of man was primarily
acquired through the inherited effects of his having worked harder than
woman for his own subsistence and that of his family; for the women in all
barbarous nations are compelled to work at least as hard as the men. With
civilised people the arbitrament of battle for the possession of the women
has long ceased; on the other hand, the men, as a general rule, have to work
harder than the women for their mutual subsistence; and thus their greater
strength will have been kept up.

Difference in the Mental Powers of the two Sexes.—With respect to
differences of this nature between man and woman, it is probable that
sexual selection has played a very important part. I am aware that some
writers doubt whether there is any inherent difference; but this is at least
probable from the analogy of the lower animals which present other
secondary sexual characters. No one will dispute that the bull differs in
disposition from the cow, the wild-boar from the sow, the stallion from the
mare, and, as is well known to the keepers of menageries, the males of the
larger apes from the females. Woman seems to differ from man in mental
disposition, chiefly in her greater tenderness and less selfishness; and this
holds good even with savages, as shewn by a well-known passage in
Mungo Park’s Travels, and by statements made by many other travellers.
Woman, owing to her maternal instincts, displays these qualities towards
her infants in an eminent degree; therefore it is likely that she should often
extend them towards her fellow-creatures. Man is the rival of other men; he
delights in competition, and this leads to ambition which passes too easily

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into selfishness. These latter qualities seem to be his natural and unfortunate
birthright. It is generally admitted that with woman the powers of intuition,
of rapid perception, and perhaps of imitation, are more strongly marked
than in man; but some, at least, of these faculties are characteristic of the
lower races, and therefore of a past and lower state of civilisation.
The chief distinction in the intellectual powers of the two sexes is shewn by
man attaining to a higher eminence, in whatever he takes up, than woman
can attain—whether requiring deep thought, reason, or imagination, or
merely the use of the senses and hands. If two lists were made of the most
eminent men and women in poetry, painting, sculpture, music,—comprising
composition and performance, history, science, and philosophy, with half-a-
dozen names under each subject, the two lists would not bear comparison.
We may also infer, from the law of the deviation of averages, so well
illustrated by Mr. Galton, in his work on ‘Hereditary Genius,’ that if men
are capable of decided eminence over women in many subjects, the average
standard of mental power in man must be above that of woman.
The half-human male progenitors of man, and men in a savage state, have
struggled together during many generations for the possession of the
females. But mere bodily strength and size would do little for victory,
unless associated with courage, perseverance, and determined energy. With
social animals, the young males have to pass through many a contest before
they win a female, and the older males have to retain their females by
renewed battles. They have, also, in the case of man, to defend their
females, as well as their young, from enemies of all kinds, and to hunt for
their joint subsistence. But to avoid enemies, or to attack them with success,
to capture wild animals, and to invent and fashion weapons, requires the aid
of the higher mental faculties, namely, observation, reason, invention, or
imagination. These various faculties will thus have been continually put to
the test, and selected during manhood; they will, moreover, have been
strengthened by use during this same period of life. Consequently, in
accordance with the principle often alluded to, we might expect that they
would at least tend to be transmitted chiefly to the male offspring at the
corresponding period of manhood.
Now, when two men are put into competition, or a man with a woman, who
possess every mental quality in the same perfection, with the exception that

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the one has higher energy, perseverance, and courage, this one will
generally become more eminent, whatever the object may be, and will gain
the victory.400 He may be said to possess genius—for genius has been
declared by a great authority to be patience; and patience, in this sense,
means unflinching, undaunted perseverance. But this view of genius is
perhaps deficient; for without the higher powers of the imagination and
reason, no eminent success in many subjects can be gained. But these latter
as well as the former faculties will have been developed in man, partly
through sexual selection,—that is, through the contest of rival males, and
partly through natural selection,—that is, from success in the general
struggle for life; and as in both cases the struggle will have been during
maturity, the characters thus gained will have been transmitted more fully to
the male than to the female offspring. Thus man has ultimately become
superior to woman. It is, indeed, fortunate that the law of the equal
transmission of characters to both sexes has commonly prevailed
throughout the whole class of mammals; otherwise it is probable that man
would have become as superior in mental endowment to woman, as the
peacock is in ornamental plumage to the peahen.
It must be borne in mind that the tendency in characters acquired at a late
period of life by either sex, to be transmitted to the same sex at the same
age, and of characters acquired at an early age to be transmitted to both
sexes, are rules which, though general, do not always hold good. If they
always held good, we might conclude (but I am here wandering beyond my
proper bounds) that the inherited effects of the early education of boys and
girls would be transmitted equally to both sexes; so that the present
inequality between the sexes in mental power could not be effaced by a
similar course of early training; nor can it have been caused by their
dissimilar early training. In order that woman should reach the same
standard as man, she ought, when nearly adult, to be trained to energy and
perseverance, and to have her reason and imagination exercised to the
highest point; and then she would probably transmit these qualities chiefly
to her adult daughters. The whole body of women, however, could not be
thus raised, unless during many generations the women who excelled in the
above robust virtues were married, and produced offspring in larger
numbers than other women. As before remarked with respect to bodily
strength, although men do not now fight for the sake of obtaining wives,

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and this form of selection has passed away, yet they generally have to
undergo, during manhood, a severe struggle in order to maintain themselves
and their families; and this will tend to keep up or even increase their
mental powers, and, as a consequence, the present inequality between the
sexes.401

Voice and Musical Powers.—In some species of Quadrumana there is a
great difference between the adult sexes, in the power of the voice and in
the development of the vocal organs; and man appears to have inherited this
difference from his early progenitors. His vocal cords are about one-third
longer than in woman, or than in boys; and emasculation produces the same
effect on him as on the lower animals, for it “arrests that prominent growth
of the thyroid, &c., which accompanies the elongation of the cords.”402 With
respect to the cause of this difference between the sexes, I have nothing to
add to the remarks made in the last chapter on the probable effects of the
long-continued use of the vocal organs by the male under the excitement of
love, rage, and jealousy. According to Sir Duncan Gibb,403 the voice differs
in the different races of mankind; and with the natives of Tartary, China,
&c., the voice of the male is said not to differ so much from that of the
female, as in most other races.
The capacity and love for singing or music, though not a sexual character in
man, must not here be passed over. Although the sounds emitted by animals
of all kinds serve many purposes, a strong case can be made out, that the
vocal organs were primarily used and perfected in relation to the
propagation of the species. Insects and some few spiders are the lowest
animals which voluntarily produce any sound; and this is generally effected
by the aid of beautifully constructed stridulating organs, which are often
confined to the males alone. The sounds thus produced consist, I believe in
all cases, of the same note, repeated rhythmically;404 and this is sometimes
pleasing even to the ears of man. Their chief, and in some cases exclusive
use appears to be either to call or to charm the opposite sex.
The sounds produced by fishes are said in some cases to be made only by
the males during the breeding season. All the air-breathing Vertebrata
necessarily possess an apparatus for inhaling and expelling air, with a pipe
capable of being closed at one end. Hence when the primeval members of

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this class were strongly excited and their muscles violently contracted,
purposeless sounds would almost certainly have been produced; and these,
if they proved in any way serviceable, might readily have been modified or
intensified by the preservation of properly adapted variations. The
Amphibians are the lowest Vertebrates which breathe air; and many of these
animals, namely, frogs and toads, possess vocal organs, which are
incessantly used during the breeding-season, and which are often more
highly developed in the male than in the female. The male alone of the
tortoise utters a noise, and this only during the season of love. Male
alligators roar or bellow during the same season. Every one knows how
largely birds use their vocal organs as a means of courtship; and some
species likewise perform what may be called instrumental music.
In the class of Mammals, with which we are here more particularly
concerned, the males of almost all the species use their voices during the
breeding-season much more than at any other time; and some are absolutely
mute excepting at this season. Both sexes of other species, or the females
alone, use their voices as a love-call. Considering these facts, and that the
vocal organs of some quadrupeds are much more largely developed in the
male than in the female, either permanently or temporarily during the
breeding season; and considering that in most of the lower classes the
sounds produced by the males, serve not only to call but to excite or allure
the female, it is a surprising fact that we have not as yet any good evidence
that these organs are used by male mammals to charm the females. The
American Mycetes caraya perhaps forms an exception, as does more
probably one of those apes which come nearer to man, namely, the
Hylobates agilis. This gibbon has an extremely loud but musical voice. Mr.
Waterhouse states,405 “It appeared to me that in ascending and descending
the scale, the intervals were always exactly half-tones; and I am sure that
the highest note was the exact octave to the lowest. The quality of the notes
is very musical; and I do not doubt that a good violinist would be able to
give a correct idea of the gibbon’s composition, excepting as regards its
loudness.” Mr. Waterhouse then gives the notes. Professor Owen, who is
likewise a musician, confirms the foregoing statement, and remarks that this
gibbon “alone of brute mammals may be said to sing.” It appears to be
much excited after its performance. Unfortunately its habits have never
been closely observed in a state of nature; but from the analogy of almost

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all other animals, it is highly probable that it utters its musical notes
especially during the season of courtship.
The perception, if not the enjoyment, of musical cadences and of rhythm is
probably common to all animals, and no doubt depends on the common
physiological nature of their nervous systems. Even Crustaceans, which are
not capable of producing any voluntary sound, possess certain auditory
hairs, which have been seen to vibrate when the proper musical notes are
struck.406 It is well known that some dogs howl when hearing particular
tones. Seals apparently appreciate music, and their fondness for it “was well
known to the ancients, and is often taken advantage of by the hunters at the
present day.”407 With all those animals, namely insects, amphibians, and
birds, the males of which during the season of courtship incessantly
produce musical notes or mere rhythmical sounds, we must believe that the
females are able to appreciate them, and are thus excited or charmed;
otherwise the incessant efforts of the males and the complex structures
often possessed exclusively by them would be useless.
With man song is generally admitted to be the basis or origin of
instrumental music. As neither the enjoyment nor the capacity of producing
musical notes are faculties of the least direct use to man in reference to his
ordinary habits of life, they must be ranked amongst the most mysterious
with which he is endowed. They are present, though in a very rude and as it
appears almost latent condition, in men of all races, even the most savage;
but so different is the taste of the different races, that our music gives not
the least pleasure to savages, and their music is to us hideous and
unmeaning. Dr. Seemann, in some interesting remarks on this subject,408
“doubts whether even amongst the nations of Western Europe, intimately
connected as they are by close and frequent intercourse, the music of the
one is interpreted in the same sense by the others. By travelling eastwards
we find that there is certainly a different language of music. Songs of joy
and dance-accompaniments are no longer, as with us, in the major keys, but
always in the minor.” Whether or not the half-human progenitors of man
possessed, like the before-mentioned gibbon, the capacity of producing, and
no doubt of appreciating, musical notes, we have every reason to believe
that man possessed these faculties at a very remote period, for singing and
music are extremely ancient arts. Poetry, which may be considered as the
offspring of song, is likewise so ancient that many persons have felt

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astonishment that it should have arisen during the earliest ages of which we
have any record.
The musical faculties, which are not wholly deficient in any race, are
capable of prompt and high development, as we see with Hottentots and
Negroes, who have readily become excellent musicians, although they do
not practise in their native countries anything that we should esteem as
music. But there is nothing anomalous in this circumstance: some species of
birds which never naturally sing, can without much difficulty be taught to
perform; thus the house-sparrow has learnt the song of a linnet. As these
two species are closely allied, and belong to the order of Insessores, which
includes nearly all the singing-birds in the world, it is quite possible or
probable that a progenitor of the sparrow may have been a songster. It is a
much more remarkable fact that parrots, which belong to a group distinct
from the Insessores, and have differently-constructed vocal organs, can be
taught not only to speak, but to pipe or whistle tunes invented by man, so
that they must have some musical capacity. Nevertheless it would be
extremely rash to assume that parrots are descended from some ancient
progenitor which was a songster. Many analogous cases could be advanced
of organs and instincts originally adapted for one purpose, having been
utilised for some quite distinct purpose.409 Hence the capacity for high
musical development, which the savage races of man possess, may be due
either to our semi-human progenitors having practised some rude form of
music, or simply to their having acquired for some distinct purposes the
proper vocal organs. But in this latter case we must assume that they
already possessed, as in the above instance of the parrots, and as seems to
occur with many animals, some sense of melody.
Music affects every emotion, but does not by itself excite in us the more
terrible emotions of horror, rage, &c. It awakens the gentler feelings of
tenderness and love, which readily pass into devotion. It likewise stirs up in
us the sensation of triumph and the glorious ardour for war. These powerful
and mingled feelings may well give rise to the sense of sublimity. We can
concentrate, as
Dr. Seemann observes, greater intensity of feeling in a single musical note
than in pages of writing. Nearly the same emotions, but much weaker and
less complex, are probably felt by birds when the male pours forth his full

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volume of song, in rivalry with other males, for the sake of captivating the
female. Love is still the commonest theme of our own songs. As Herbert
Spencer remarks, music “arouses dormant sentiments of which we had not
conceived the possibility, and do not know the meaning; or, as Richter says,
tells us of things we have not seen and shall not see.”410 Conversely, when
vivid emotions are felt and expressed by the orator or even in common
speech, musical cadences and rhythm are instinctively used. Monkeys also
express strong feelings in different tones—anger and impatience by low,—
fear and pain by high notes.411 The sensations and ideas excited in us by
music, or by the cadences of impassioned oratory, appear from their
vagueness, yet depth, like mental reversions to the emotions and thoughts of
a long-past age.
All these facts with respect to music become to a certain extent intelligible
if we may assume that musical tones and rhythm were used by the half-
human progenitors of man, during the season of courtship, when animals of
all kinds are excited by the strongest passions. In this case, from the deeply-
laid principle of inherited associations, musical tones would be likely to
excite in us, in a vague and indefinite manner, the strong emotions of a
long-past age. Bearing in mind that the males of some quadrumanous
animals have their vocal organs much more developed than in the females,
and that one anthropomorphous species pours forth a whole octave of
musical notes and may be said to sing, the suspicion does not appear
improbable that the progenitors of man, either the males or females, or both
sexes, before they had acquired the power of expressing their mutual love in
articulate language, endeavoured to charm each other with musical notes
and rhythm. So little is known about the use of the voice by the
Quadrumana during the season of love, that we have hardly any means of
judging whether the habit of singing was first acquired by the male or
female progenitors of mankind. Women are generally thought to possess
sweeter voices than men, and as far as this serves as any guide we may infer
that they first acquired musical powers in order to attract the other sex.412
But if so, this must have occurred long ago, before the progenitors of man
had become sufficiently human to treat and value their women merely as
useful slaves. The impassioned orator, bard, or musician, when with his
varied tones and cadences he excites the strongest emotions in his hearers,
little suspects that he uses the same means by which, at an extremely

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remote period, his half-human ancestors aroused each other’s ardent
passions, during their mutual courtship and rivalry.

On the influence of beauty in determining the marriages of mankind.—In
civilised life man is largely, but by no means exclusively, influenced in the
choice of his wife by external appearance; but we are chiefly concerned
with primeval times, and our only means of forming a judgment on this
subject is to study the habits of existing semi-civilised and savage nations.
If it can be shewn that the men of different races prefer women having
certain characteristics, or conversely that the women prefer certain men, we
have then to enquire whether such choice, continued during many
generations, would produce any sensible effect on the race, either on one
sex or both sexes; this latter circumstance depending on the form of
inheritance which prevails.
It will be well first to shew in some detail that savages pay the greatest
attention to their personal appearance.413 That they have a passion for
ornament is notorious; and an English philosopher goes so far as to
maintain that clothes were first made for ornament and not for warmth. As
Professor Waitz remarks, “however poor and miserable man is, he finds a
pleasure in adorning himself.” The extravagance of the naked Indians of
South America in decorating themselves is shewn “by a man of large stature
gaining with difficulty enough by the labour of a fortnight to procure in
exchange the chica necessary to paint himself red.”414 The ancient
barbarians of Europe during the Reindeer period brought to their caves any
brilliant or singular objects which they happened to find. Savages at the
present day everywhere deck themselves with plumes, necklaces, armlets,
earrings, &c. They paint themselves in the most diversified manner. “If
painted nations,” as Humboldt observes, “had been examined with the same
attention as clothed nations, it would have been perceived that the most
fertile imagination and the most mutable caprice have created the fashions
of painting, as well as those of garments.”
In one part of Africa the eyelids are coloured black; in another the nails are
coloured yellow or purple. In many places the hair is dyed of various tints.
In different countries the teeth are stained black, red, blue, &c., and in the
Malay Archipelago it is thought shameful to have white teeth like those of a

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dog. Not one great country can be named, from the Polar regions in the
north to New Zealand in the south, in which the aborigines do not tattoo
themselves. This practice was followed by the Jews of old and by the
ancient Britons. In Africa some of the natives tattoo themselves, but it is
much more common to raise protuberances by rubbing salt into incisions
made in various parts of the body; and these are considered by the
inhabitants of Kordofan and Darfur “to be great personal attractions.” In the
Arab countries no beauty can be perfect until the cheeks or temples have
been gashed.”415 In South America, as Humboldt remarks, “a mother would
be accused of culpable indifference towards her children, if she did not
employ artificial means to shape the calf of the leg after the fashion of the
country.” In the Old and New World the shape of the skull was formerly
modified during infancy in the most extraordinary manner, as is still the
case in many places, and such deformities are considered ornamental. For
instance, the savages of Colombia416 deem a much flattened head “an
essential point of beauty.”
The hair is treated with especial care in various countries; it is allowed to
grow to full length, so as to reach to the ground, or is combed into “a
compact frizzled mop, which is the Papuan’s pride and glory.”417 In
Northern Africa “a man requires a period of from eight to ten years to
perfect his coiffure.” With other nations the head is shaved, and in parts of
South America and Africa even the eyebrows are eradicated. The natives of
the Upper Nile knock out the four front teeth, saying that they do not wish
to resemble brutes. Further south, the Batokas knock out the two upper
incisors, which, as Livingstone418 remarks, gives the face a hideous
appearance, owing to the growth of the lower jaw; but these people think
the presence of the incisors most unsightly, and on beholding some
Europeans, cried out, “Look at the great teeth!” The great chief Sebituani
tried in vain to alter this fashion. In various parts of Africa and in the Malay
Archipelago the natives file the incisor teeth into points like those of a saw,
or pierce them with holes, into which they insert studs.
As the face with us is chiefly admired for its beauty, so with savages it is
the chief seat of mutilation. In all quarters of the world the septum, and
more rarely the wings of the nose are pierced, with rings, sticks, feathers,
and other ornaments inserted into the holes. The ears are everywhere
pierced and similarly ornamented, and with the Botocudos and Lenguas of

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South America the hole is gradually so much enlarged that the lower edge
touches the shoulder. In North and South America and in Africa either the
upper or lower lip is pierced; and with the Botocudos the hole in the lower
lip is so large that a disc of wood four inches in diameter is placed in it.
Mantegazza gives a curious account of the shame felt by a South American
native, and of the ridicule which he excited, when he sold his tembeta,—the
large coloured piece of wood which is passed through the hole. In central
Africa the women perforate the lower lip and wear a crystal, which, from
the movement of the tongue, has “a wriggling motion indescribably
ludicrous during conversation.” The wife of the chief of Latooka told Sir S.
Baker419 that his “wife would be much improved if she would extract her
four front teeth from the lower jaw, and wear the long pointed polished
crystal in her under lip.” Further south with the Makalolo, the upper lip is
perforated, and a large metal and bamboo ring, called a pelelé, is worn in
the hole. “This caused the lip in one case to project two inches beyond the
tip of the nose; and when the lady smiled the contraction of the muscles
elevated it over the eyes. ‘Why do the women wear these things?’ the
venerable chief, Chinsurdi, was asked. Evidently surprised at such a stupid
question, he replied, ‘For beauty! They are the only beautiful things women
have; men have beards, women have none. What kind of a person would
she be without the pelelé? She would not be a woman at all with a mouth
like a man, but no beard.’”420
Hardly any part of the body, which can be unnaturally modified, has
escaped. The amount of suffering thus caused must have been wonderfully
great, for many of the operations require several years for their completion,
so that the idea of their necessity must be imperative. The motives are
various; the men paint their bodies to make themselves appear terrible in
battle; certain mutilations are connected with religious rites; or they mark
the age of puberty, or the rank of the man, or they serve to distinguish the
tribes. As with savages the same fashions prevail for long periods,421
mutilations, from whatever cause first made, soon come to be valued as
distinctive marks. But self-adornment, vanity, and the admiration of others,
seem to be the commonest motives. In regard to tattooing, I was told by the
missionaries in New Zealand, that when they tried to persuade some girls to
give up the practice, they answered, “We must just have a few lines on our
lips; else when we grow old we shall be so very ugly.” With the men of

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New Zealand, a most capable judge422 says, “to have fine tattooed faces was
the great ambition of the young, both to render themselves attractive to the
ladies, and conspicuous in war.” A star tattooed on the forehead and a spot
on the chin are thought by the women in one part of Africa to be irresistible
attractions.423 In most, but not all parts of the world, the men are more
highly ornamented than the women, and often in a different manner;
sometimes, though rarely, the women are hardly at all ornamented. As the
women are made by savages to perform the greatest share of the work, and
as they are not allowed to eat the best kinds of food, so it accords with the
characteristic selfishness of man that they should not be allowed to obtain,
or to use, the finest ornaments. Lastly it is a remarkable fact, as proved by
the foregoing quotations, that the same fashions in modifying the shape of
the head, in ornamenting the hair, in painting, tattooing, perforating the
nose, lips, or ears, in removing or filing the teeth, &c., now prevail and
have long prevailed in the most distant quarters of the world. It is extremely
improbable that these practices which are followed by so many distinct
nations are due to tradition from any common source. They rather indicate
the close similarity of the mind of man, to whatever race he may belong, in
the same manner as the almost universal habits of dancing, masquerading,
and making rude pictures.

Having made these preliminary remarks on the admiration felt by savages
for various ornaments, and for deformities most unsightly in our eyes, let us
see how far the men are attracted by the appearance of their women, and
what are their ideas of beauty. As I have heard it maintained that savages
are quite indifferent about the beauty of their women, valuing them solely
as slaves, it may be well to observe that this conclusion does not at all agree
with the care which the women take in ornamenting themselves, or with
their vanity. Burchell424 gives an amusing account of a Bushwoman, who
used so much grease, red ochre, and shining powder, “as would have ruined
any but a very rich husband.” She displayed also “much vanity and too
evident a consciousness of her superiority.” Mr. Winwood Reade informs
me that the negroes of the West Coast often discuss the beauty of their
women. Some competent observers have attributed the fearfully common
practice of infanticide partly to the desire felt by the women to retain their
good looks.425 In several regions the women wear charms and love-philters

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to gain the affections of the men; and Mr. Brown enumerates four plants
used for this purpose by the women of North-Western America.426
Hearne,427 who lived many years with the American Indians, and who was
an excellent observer, says, in speaking of the women, “Ask a Northern
Indian what is beauty, and he will answer, a broad flat face, small eyes, high
cheek-bones, three or four broad black lines across each cheek, a low
forehead, a large broad chin, a clumsy hook nose, a tawny hide, and breasts
hanging down to the belt.” Pallas, who visited the northern parts of the
Chinese empire, says “those women are preferred who have the Mandschú
form; that is to say, a broad face, high cheek-bones, very broad noses, and
enormous ears;”428 and Vogt remarks that the obliquity of the eye, which is
proper to the Chinese and Japanese, is exaggerated in their pictures for the
purpose, as “it seems, of exhibiting its beauty, as contrasted with the eye of
the red-haired barbarians.” It is well known, as Huc repeatedly remarks,
that the Chinese of the interior think Europeans hideous with their white
skins and prominent noses. The nose is far from being too prominent,
according to our ideas, in the natives of Ceylon; yet “the Chinese in the
seventh century, accustomed to the flat features of the Mogul races, were
surprised at the prominent noses of the Cingalese; and Thsang described
them as having ‘the beak of a bird, with the body of a man.’”
Finlayson, after minutely describing the people of Cochin China, says that
their rounded heads and faces are their chief characteristics; and, he adds,
“the roundness of the whole countenance is more striking in the women,
who are reckoned beautiful in proportion as they display this form of face.”
The Siamese have small noses with divergent nostrils, a wide mouth, rather
thick lips, a remarkably large face, with very high and broad cheek-bones. It
is, therefore, not wonderful that “beauty, according to our notion is a
stranger to them. Yet they consider their own females to be much more
beautiful than those of Europe.”429
It is well known that with many Hottentot women the posterior part of the
body projects in a wonderful manner; they are steatopygous; and Sir
Andrew Smith is certain that this peculiarity is greatly admired by the
men.430 He once saw a woman who was considered a beauty, and she was so
immensely developed behind, that when seated on level ground she could
not rise, and had to push herself along until she came to a slope. Some of

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the women in various negro tribes are similarly characterised; and,
according to Burton, the Somal men “are said to choose their wives by
ranging them in a line, and by picking her out who projects farthest a tergo.
Nothing can be more hateful to a negro than the opposite form.”431
With respect to colour, the negroes rallied Mungo Park on the whiteness of
his skin and the prominence of his nose, both of which they considered as
“unsightly and unnatural conformations.” He in return praised the glossy jet
of their skins and the lovely depression of their noses; this they said was
“honey-mouth,” nevertheless they gave him food. The African Moors, also,
“knitted their brows and seemed to shudder” at the whiteness of his skin.
On the eastern coast, the negro boys when they saw Burton, cried out “Look
at the white man; does he not look like a white ape?” On the western coast,
as Mr. Winwood Reade informs me, the negroes admire a very black skin
more than one of a lighter tint. But their horror of whiteness may be partly
attributed, according to this same traveller, to the belief held by most
negroes that demons and spirits are white.
The Banyai of the more southern part of the continent are negroes, but “a
great many of them are of a light coffee-and-milk colour, and, indeed, this
colour is considered handsome throughout the whole country;” so that here
we have a different standard of taste. With the Kafirs, who differ much from
negroes, “the skin, except among the tribes near Delagoa Bay, is not usually
black, the prevailing colour being a mixture of black and red, the most
common shade being chocolate. Dark complexions, as being most common
are naturally held in the highest esteem. To be told that he is light-coloured,
or like a white man, would be deemed a very poor compliment by a Kafir. I
have heard of one unfortunate man who was so very fair that no girl would
marry him.” One of the titles of the Zulu king is “You who are black.”432
Mr. Galton, in speaking to me about the natives of S. Africa, remarked that
their ideas of beauty seem very different from ours; for in one tribe two
slim, slight, and pretty girls were not admired by the natives.
Turning to other quarters of the world; in Java, a yellow, not a white girl, is
considered, according to Madame Pfeiffer, a beauty. A man of Cochin-
China “spoke with contempt of the wife of the English Ambassador, that
she had white teeth like a dog, and a rosy colour like that of potato-
flowers.” We have seen that the Chinese dislike our white skin, and that the

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N. Americans admire “a tawny hide.” In S. America, the Yura-caras, who
inhabit the wooded, damp slopes of the eastern Cordillera, are remarkably
pale-coloured, as their name in their own language expresses; nevertheless
they consider European women as very inferior to their own.433
In several of the tribes of North America the hair on the head grows to a
wonderful length; and Catlin gives a curious proof how much this is
esteemed, for the chief of the Crows was elected to this office from having
the longest hair of any man in the tribe, namely ten feet and seven inches.
The Aymaras and Quechuas of S. America, likewise have very long hair;
and this, as Mr. D. Forbes informs me, is so much valued for the sake of
beauty, that cutting it off was the severest punishment which he could inflict
on them. In both halves of the continent the natives sometimes increase the
apparent length of their hair by weaving into it fibrous substances. Although
the hair on the head is thus cherished, that on the face is considered by the
North American Indians “as very vulgar,” and every hair is carefully
eradicated. This practice prevails throughout the American continent from
Vancouver’s Island in the north to Tierra del Fuego in the south. When York
Minster, a Fuegian on board the “Beagle” was taken back to his country, the
natives told him he ought to pull out the few short hairs on his face. They
also threatened a young missionary, who was left for a time with them, to
strip him naked, and pluck the hairs from his face and body, yet he was far
from a hairy man. This fashion is carried to such an extreme that the Indians
of Paraguay eradicate their eyebrows and eyelashes, saying that they do not
wish to be like horses.434
It is remarkable that throughout the world the races which are almost
completely destitute of a beard dislike hairs on the face and body, and take
pains to eradicate them. The Kalmucks are beardless, and they are well
known, like the Americans, to pluck out all straggling hairs; and so it is
with the Polynesians, some of the Malays, and the Siamese. Mr. Veitch
states that the Japanese ladies “all objected to our whiskers, considering
them very ugly, and told us to cut them off, and be like Japanese men.” The
New Zealanders are beardless; they carefully pluck out the hairs on the
face, and have a saying that “There is no woman for a hairy man.”435
On the other hand, bearded races admire and greatly value their beards;
among the Anglo-Saxons every part of the body, according to their laws,

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had a recognised value; “the loss of the beard being estimated at twenty
shillings, while the breaking of a thigh was fixed at only twelve.”436 In the
East men swear solemnly by their beards. We have seen that Chinsurdi, the
chief of the Makalolo in Africa, evidently thought that beards were a great
ornament. With the Fijians in the Pacific the beard is “profuse and bushy,
and is his greatest pride;” whilst the inhabitants of the adjacent
archipelagoes of Tonga and Samoa are “beardless, and abhor a rough chin.”
In one island alone of the Ellice group “the men are heavily bearded, and
not a little proud thereof.”437
We thus see how widely the different races of man differ in their taste for
the beautiful. In every nation sufficiently advanced to have made effigies of
their gods or of their deified rulers, the sculptors no doubt have
endeavoured to express their highest ideal of beauty and grandeur.438 Under
this point of view it is well to compare in our mind the Jupiter or Apollo of
the Greeks with the Egyptian or Assyrian statues; and these with the
hideous bas-reliefs on the ruined buildings of Central America.
I have met with very few statements opposed to the above conclusion. Mr.
Winwood Reade, however, who has had ample opportunities for
observation, not only with the negroes of the West Coast of Africa, but with
those of the interior who have never associated with Europeans, is
convinced that their ideas of beauty are on the whole the same as ours. He
has repeatedly found that he agreed with negroes in their estimation of the
beauty of the native girls; and that their appreciation of the beauty of
European women corresponded with ours. They admire long hair, and use
artificial means to make it appear abundant; they admire also a beard,
though themselves very scantily provided. Mr. Reade feels doubtful what
kind of nose is most appreciated: a girl has been heard to say, “I do not want
to marry him, he has got no nose;” and this shews that a very flat nose is not
an object of admiration. We should, however, bear in mind that the
depressed and very broad noses and projecting jaws of the negroes of the
West Coast are exceptional types with the inhabitants of Africa.
Notwithstanding the foregoing statements, Mr. Reade does not think it
probable that negroes would ever prefer the “most beautiful European
woman, on the mere grounds of physical admiration, to a good-looking
negress.”439

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The truth of the principle, long ago insisted on by Humboldt,440 that man
admires and often tries to exaggerate whatever characters nature may have
given him, is shewn in many ways. The practice of beardless races
extirpating every trace of a beard, and generally all the hairs on the body,
offers one illustration. The skull has been greatly modified during ancient
and modern times by many nations; and there can be little doubt that this
has been practised, especially in N. and S. America, in order to exaggerate
some natural and admired peculiarity. Many American Indians are known to
admire a head flattened to such an extreme degree as to appear to us like
that of an idiot. The natives on the nort-hwestern coast compress the head
into a pointed cone; and it is their constant practice to gather the hair into a
knot on the top of the head, for the sake, as Dr. Wilson remarks, “of
increasing the apparent elevation of the favourite conoid form.” The
inhabitants of Arakhan “admire a broad, smooth forehead, and in order to
produce it, they fasten a plate of lead on the heads of the newborn
children.” On the other hand, “a broad, well-rounded occiput is considered
a great beauty” by the natives of the Fiji islands.441
As with the skull, so with the nose; the ancient Huns during the age of
Attila were accustomed to flatten the noses of their infants with bandages,
“for the sake of exaggerating a natural conformation.” With the Tahitians, to
be called, long-nose is considered as an insult, and they compress the noses
and foreheads of their children for the sake of beauty. So it is with the
Malays of Sumatra, the Hottentots, certain Negroes, and the natives of
Brazil.442 The Chinese have by nature unusually small feet;443 and it is well
known that the women of the upper classes distort their feet to make them
still smaller. Lastly, Humboldt thinks that the American Indians prefer
colouring their bodies with red paint in order to exaggerate their natural
tint; and until recently European women added to their naturally bright
colours by rouge and white cosmetics; but I doubt whether many barbarous
nations have had any such intention in painting themselves.
In the fashions of our own dress we see exactly the same principle and the
same desire to carry every point to an extreme; we exhibit, also, the same
spirit of emulation. But the fashions of savages are far more permanent than
ours; and whenever their bodies are artificially modified this is necessarily
the case. The Arab women of the Upper Nile occupy about three days in
dressing their hair; they never imitate other tribes, “but simply vie with each

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other in the superlativeness of their own style.” Dr. Wilson, in speaking of
the compressed skulls of various American races, adds, “such usages are
among the least eradicable, and long survive the shock of revolutions that
change dynasties and efface more important national peculiarities.”444 The
same principle comes largely into play in the art of selection; and we can
thus understand, as I have elsewhere explained,445 the wonderful
development of all the races of animals and plants which are kept merely
for ornament. Fanciers always wish each character to be somewhat
increased; they do not admire a medium standard; they certainly do not
desire any great and abrupt change in the character of their breeds; they
admire solely what they are accustomed to behold, but they ardently desire
to see each characteristic feature a little more developed.
No doubt the perceptive powers of man and the lower animals are so
constituted that brilliant colours and certain forms, as well as harmonious
and rhythmical sounds, give pleasure and are called beautiful; but why this
should be so, we know no more than why certain bodily sensations are
agreeable and others disagreeable. It is certainly not true that there is in the
mind of man any universal standard of beauty with respect to the human
body. It is, however, possible that certain tastes may in the course of time
become inherited, though I know of no evidence in favour of this belief;
and if so, each race would possess its own innate ideal standard of beauty. It
has been argued446 that ugliness consists in an approach to the structure of
the lower animals, and this no doubt is true with the more civilised nations,
in which intellect is highly appreciated; but a nose twice as prominent, or
eyes twice as large as usual, would not be an approach in structure to any of
the lower animals, and yet would be utterly hideous. The men of each race
prefer what they are accustomed to behold; they cannot endure any great
change; but they like variety, and admire each characteristic point carried to
a moderate extreme.447 Men accustomed to a nearly oval face, to straight
and regular features, and to bright colours, admire, as we Europeans know,
these points when strongly developed. On the other hand, men accustomed
to a broad face, with high cheek-bones, a depressed nose, and a black skin,
admire these points strongly developed. No doubt characters of all kinds
may easily be too much developed for beauty. Hence a perfect beauty,
which implies many characters modified in a particular manner, will in
every race be a prodigy. As the great anatomist Bichat long ago said, if

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every one were cast in the same mould, there would be no such thing as
beauty. If all our women were to become as beautiful as the Venus de
Medici, we should for a time be charmed; but we should soon wish for
variety; and as soon as we had obtained variety, we should wish to see
certain characters in our women a little exaggerated beyond the then
existing common standard.

CHAPTER XX.

Secondary Sexual Characters of Man—continued.

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On the effects of the continued selection of women according to a different standard of beauty in
each race—On the causes which interfere with sexual selection in civilised and savage nations
—Conditions favourable to sexual selection during primeval times—On the manner of action of
sexual selection with mankind—On the women in savage tribes having some power to choose
their husbands—Absence of hair on the body, and development of the beard—Colour of the skin
—Summary.

We have seen in the last chapter that with all barbarous races ornaments,
dress, and external appearance are highly valued; and that the men judge of
the beauty of their women by widely different standards. We must next
inquire whether this preference and the consequent selection during many
generations of those women, which, appear to the men of each race the
most attractive, has altered the character either of the females alone or of
both sexes. With mammals the general rule appears to be that characters of
all kinds are inherited equally by the males and females; we might therefore
expect that with mankind any characters gained through sexual selection by
the females would commonly be transferred to the offspring of both sexes.
If any change has thus been effected it is almost certain that the different
races will have been differently modified, as each has its own standard of
beauty.
With mankind, especially with savages, many causes interfere with the
action of sexual selection as far as the bodily frame is concerned. Civilised
men are largely attracted by the mental charms of women, by their wealth,
and especially by their social position; for men rarely marry into a much
lower rank of life. The men who succeed in obtaining the more beautiful
women, will not have a better chance of leaving a long line of descendants
than other men with plainer wives, with the exception of the few who
bequeath their fortunes according to primogeniture. With respect to the
opposite form of selection, namely of the more attractive men by the
women, although in civilised nations women have free or almost free
choice, which is not the case with barbarous races, yet their choice is
largely influenced by the social position and wealth of the men; and the
success of the latter in life largely depends on their intellectual powers and
energy, or on the fruits of these same powers in their forefathers.
There is, however, reason to believe that sexual selection has effected
something in certain civilised and semi-civilised nations. Many persons are
convinced, as it appears to me with justice, that the members of our

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aristocracy, including under this term all wealthy families in which
primogeniture has long prevailed, from having chosen during many
generations from all classes the more beautiful women as their wives, have
become handsomer, according to the European standard of beauty, than the
middle classes; yet the middle classes are placed under equally favourable
conditions of life for the perfect development of the body. Cook remarks
that the superiority in personal appearance “which is observable in the erees
or nobles in all the other islands (of the Pacific) is found in the Sandwich
islands;” but this may be chiefly due to their better food and manner of life.
The old traveller Chardin, in describing the Persians, says their “blood is
now highly refined by frequent intermixtures with the Georgians and
Circassians, two nations which surpass all the world in personal beauty.
There is hardly a man of rank in Persia who is not born of a Georgian or
Circassian mother.” He adds that they inherit their beauty, “not from their
ancestors, for without the above mixture, the men of rank in Persia, who are
descendants of the Tartars, would be extremely ugly.”448 Here is a more
curious case: the priestesses who attended the temple of Venus Erycina at
San-Giuliano in Sicily, were selected for their beauty out of the whole of
Greece; they were not vestal virgins, and Quatrefages,449 who makes this
statement, says that the women of San-Giuliano are famous at the present
day as the most beautiful in the island, and are sought by artists as models.
But it is obvious that the evidence in the above cases is doubtful.
The following case, though relating to savages, is well worth giving from
its curiosity. Mr. Winwood Reade informs me that the Jollofs, a tribe of
negroes on the west coast of Africa, “are remarkable for their uniformly fine
appearance.” A friend of his asked one of these men, “How is it that every
one whom I meet is so fine-looking, not only your men, but your women?”
The Jollof answered, “It is very easily explained: it has always been our
custom to pick out our worse-looking slaves and to sell them.” It need
hardly be added that with all savages female slaves serve as concubines.
That this negro should have attributed, whether rightly or wrongly, the fine
appearance of his tribe, to the long-continued elimination of the ugly
women, is not so surprising as it may at first appear; for I have elsewhere
shewn450 that negroes fully appreciate the importance of selection in the
breeding of their domestic animals, and I could give from Mr. Reade
additional evidence on this head.

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On the Causes which prevent or check the Action of Sexual Selection with
Savages.—The chief causes are, firstly, so-called communal marriages or
promiscuous intercourse; secondly, infanticide, especially of female infants;
thirdly, early betrothals; and lastly, the low estimation in which women are
held, as mere slaves. These four points must be considered in some detail.
It is obvious that as long as the pairing of man, or of any other animal, is
left to chance, with no choice exerted by either sex, there can be no sexual
selection; and no effect will be produced on the offspring by certain
individuals having had an advantage over others in their courtship. Now it
is asserted that there exist at the present day tribes which practise what Sir
J. Lubbock by courtesy calls communal marriages; that is, all the men and
women in the tribe are husbands and wives to each other. The licentiousness
of many savages is no doubt astonishingly great, but it seems to me that
more evidence is requisite before we fully admit that their existing
intercourse is absolutely promiscuous. Nevertheless all those who have
most closely studied the subject,451 and whose judgment is worth much
more than mine, believe that communal marriage was the original and
universal form throughout the world, including the intermarriage of
brothers and sisters. The indirect evidence in favour of this belief is
extremely strong, and rests chiefly on the terms of relationship which are
employed between the members of the same tribe, implying a connection
with the tribe alone, and not with either parent. But the subject is too large
and complex for even an abstract to be here given, and I will confine myself
to a few remarks. It is evident in the case of communal marriages, or where
the marriage-tie is very loose, that the relationship of the child to its father
cannot be known. But it seems almost incredible that the relationship of the
child to its mother should ever have been completely ignored, especially as
the women in most savage tribes nurse their infants for a long time.
Accordingly in many cases the lines of descent are traced through the
mother alone, to the exclusion of the father. But in many other cases the
terms employed express a connection with the tribe alone, to the exclusion
even of the mother. It seems possible that the connection between the
related members of the same barbarous tribe, exposed to all sorts of danger,
might be so much more important, owing to the need of mutual protection
and aid, than that between the mother and her child, as to lead to the sole

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use of terms expressive of the former relationships; but Mr. Morgan is
convinced that this view of the case is by no means sufficient.
The terms of relationship used in different parts of the world may be
divided, according to the author just quoted, into two great classes, the
classificatory and descriptive,—the latter being employed by us. It is the
classificatory system which so strongly leads to the belief that communal
and other extremely loose forms of marriage were originally universal. But
as far as I can see, there is no necessity on this ground for believing in
absolutely promiscuous intercourse. Men and women, like many of the
lower animals, might formerly have entered into strict though temporary
unions for each birth, and in this case nearly as much confusion would have
arisen in the terms of relationship as in the case of promiscuous intercourse.
As far as sexual selection is concerned, all that is required is that choice
should be exerted before the parents unite, and it signifies little whether the
unions last for life or only for a season.
Besides the evidence derived from the terms of relationship, other lines of
reasoning indicate the former wide prevalence of communal marriage. Sir J.
Lubbock ingeniously accounts452 for the strange and widely-extended habit
of exogamy,—that is, the men of one tribe always taking wives from a
distinct tribe,—by communism having been the original form of marriage;
so that a man never obtained a wife for himself unless he captured her from
a neighbouring and hostile tribe, and then she would naturally have become
his sole and valuable property. Thus the practice of capturing wives might
have arisen; and from the honour so gained might ultimately have become
the universal habit. We can also, according to Sir J. Lubbock,452 thus
understand “the necessity of expiation for marriage as an infringement of
tribal rites, since, according to old ideas, a man had no right to appropriate
to himself that which belonged to the whole tribe.” Sir J. Lubbock further
gives a most curious body of facts shewing that in old times high honour
was bestowed on women who were utterly licentious; and this, as he
explains, is intelligible, if we admit that promiscuous intercourse was the
aboriginal and therefore long revered custom of the tribe.453
Although the manner of development of the marriage-tie is an obscure
subject, as we may infer from the divergent opinions on several points
between the three authors who have studied it most closely, namely, Mr.

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Morgan, Mr. M’Lennan, and Sir J. Lubbock, yet from the foregoing and
several other lines of evidence it seems certain that the habit of marriage
has been gradually developed, and that almost promiscuous intercourse was
once extremely common throughout the world. Nevertheless from the
analogy of the lower animals, more particularly of those which come
nearest to man in the series, I cannot believe that this habit prevailed at an
extremely remote period, when man had hardly attained to his present rank
in the zoological scale. Man, as I have attempted to shew, is certainly
descended from some ape-like creature. With the existing Quadrumana, as
far as their habits are known, the males of some species are monogamous,
but live during only a part of the year with the females, as seems to be the
case with the Orang. Several kinds, as some of the Indian and American
monkeys, are strictly monogamous, and associate all the year round with
their wives. Others are polygamous, as the Gorilla and several American
species, and each family lives separate. Even when this occurs, the families
inhabiting the same district are probably to a certain extent social: the
Chimpanzee, for instance, is occasionally met with in large bands. Again,
other species are polygamous, but several males, each with their own
females, live associated in a body, as with several species of Baboons.454 We
may indeed conclude from what we know of the jealousy of all male
quadrupeds, armed, as many of them are, with special weapons for battling
with their rivals, that promiscuous intercourse in a state of nature is
extremely improbable. The pairing may not last for life, but only for each
birth; yet if the males which are the strongest and best able to defend or
otherwise assist their females and young offspring, were to select the more
attractive females, this would suffice for the work of sexual selection.
Therefore, if we look far enough back in the stream of time, it is extremely
improbable that primeval men and women lived promiscuously together.
Judging from the social habits of man as he now exists, and from most
savages being polygamists, the most probable view is that primeval man
aboriginally lived in small communities, each with as many wives as he
could support and obtain, whom he would have jealously guarded against
all other men. Or he may have lived with several wives by himself, like the
Gorilla; for all the natives “agree that but one adult male is seen in a band;
when the young male grows up, a contest takes place for mastery, and the
strongest, by killing and driving out the others, establishes himself as the

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head of the community.”455 The younger males, being thus expelled and
wandering about, would, when at last successful in finding a partner,
prevent too close interbreeding within the limits of the same family.
Although savages are now extremely licentious, and although communal
marriages may formerly have largely prevailed, yet many tribes practise
some form of marriage, but of a far more lax nature than with civilised
nations. Polygamy, as just stated, is almost universally followed by the
leading men in every tribe. Nevertheless there are tribes, standing almost at
the bottom of the scale, which are strictly monogamous. This is the case
with the Veddahs of Ceylon: they have a saying, according to Sir J.
Lubbock,456 “that death alone can separate husband and wife.” An
intelligent Kandyan chief, of course a polygamist, “was perfectly
scandalized at the utter barbarism of living with only one wife, and never
parting until separated by death.” It was, he said, “just like the Wanderoo
monkeys.” Whether savages who now enter into some form of marriage,
either polygamous or monogamous, have retained this habit from primeval
times, or whether they have returned to some form of marriage, after
passing through a stage of promiscuous intercourse, I will not pretend to
conjecture.

Infanticide.—This practice is now very common throughout the world, and
there is reason to believe that it prevailed much more extensively during
former times.457 Barbarians find it difficult to support themselves and their
children, and it is a simple plan to kill their infants. In South America some
tribes, as Azara states, formerly destroyed so many infants of both sexes,
that they were on the point of extinction. In the Polynesian Islands women
have been known to kill from four or five to even ten of their children; and
Ellis could not find a single woman who had not killed at least one.
Wherever infanticide prevails the struggle for existence will be in so far less
severe, and all the members of the tribe will have an almost equally good
chance of rearing their few surviving children. In most cases a larger
number of female than of male infants are destroyed, for it is obvious that
the latter are of most value to the tribe, as they will when grown up aid in
defending it, and can support themselves. But the trouble experienced by
the women in rearing children, their consequent loss of beauty, the higher
estimation set on them and their happier fate, when few in number, are

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assigned by the women themselves, and by various observers, as additional
motives for infanticide. In Australia, where female infanticide is still
common, Sir G. Grey estimated the proportion of native women to men as
one to three; but others say as two to three. In a village on the eastern
frontier of India, Colonel Macculloch found not a single female child.458
When, owing to female infanticide, the women of a tribe are few in number,
the habit of capturing wives from neighbouring tribes would naturally arise.
Sir J. Lubbock, however, as we have seen, attributes the practice in chief
part, to the former existence of communal marriage, and to the men having
consequently captured women from other tribes to hold as their sole
property. Additional causes might be assigned, such as the communities
being very small, in which case, marriageable women would often be
deficient. That the habit of capture was most extensively practised during
former times, even by the ancestors of civilised nations, is clearly shewn by
the preservation of many curious customs and ceremonies, of which Mr.
M’Lennan has given a most interesting account. In our own marriages the
“best man” seems originally to have been the chief abettor of the
bridegroom in the act of capture. Now as long as men habitually procured
their wives through violence and craft, it is not probable that they would
have selected the more attractive women; they would have been too glad to
have seized on any woman. But as soon as the practice of procuring wives
from a distinct tribe was effected through barter, as now occurs in many
places, the more attractive women would generally have been purchased.
The incessant crossing, however, between tribe and tribe, which necessarily
follows from any form of this habit would have tended to keep all the
people inhabiting the same country nearly uniform in character; and this
would have greatly interfered with the power of sexual selection in
differentiating the tribes.
The scarcity of women, consequent on female infanticide, leads, also, to
another practice, namely polyandry, which is still common in several parts
of the world, and which formerly, as Mr. M’Lennan believes, prevailed
almost universally; but this latter conclusion is doubted by Mr. Morgan and
Sir J. Lubbock.459 Whenever two or more men are compelled to marry one
woman, it is certain that all the women of the tribe will get married, and
there will be no selection by the men of the more attractive women. But
under these circumstances the women no doubt will have the power of

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choice, and will prefer the more attractive men. Azara, for instance,
describes how carefully a Guana woman bargains for all sorts of privileges,
before accepting some one or more husbands; and the men in consequence
take unusual care of their personal appearance.460 The very ugly men would
perhaps altogether fail in getting a wife, or get one later in life, but the
handsomer men, although the most successful in obtaining wives, would
not, as far as we can see, leave more offspring to inherit their beauty than
the less handsome husbands of the same women.

Early Betrothals and Slavery of Women.—With many savages it is the
custom to betroth the females whilst mere infants; and this would
effectually prevent preference being exerted on either side according to
personal appearance. But it would not prevent the more attractive women
from being afterwards stolen or taken by force from their husbands by the
more powerful men; and this often happens in Australia, America, and other
parts of the world. The same consequences with reference to sexual
selection would to a certain extent follow when women are valued almost
exclusively as slaves or beasts of burden, as is the case with most savages.
The men, however, at all times would prefer the handsomest slaves
according to their standard of beauty.
We thus see that several customs prevail with savages which would greatly
interfere with, or completely stop, the action of sexual selection. On the
other hand, the conditions of life to which savages are exposed, and some of
their habits, are favourable to natural selection; and this always comes into
play together with sexual selection. Savages are known to suffer severely
from recurrent famines; they do not increase their food by artificial means;
they rarely refrain from marriage,461 and generally marry young.
Consequently they must be subjected to occasional hard struggles for
existence, and the favoured individuals will alone survive.
Turning to primeval times when men had only doubtfully attained the rank
of manhood, they would probably have lived, as already stated, either as
polygamists or temporarily as monogamists. Their intercourse, judging
from analogy, would not then have been promiscuous. They would, no
doubt, have defended their females to the best of their power from enemies
of all kinds, and would probably have hunted for their subsistence, as well

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as for that of their offspring. The most powerful and able males would have
succeeded best in the struggle for life and in obtaining attractive females. At
this early period the progenitors of man, from having only feeble powers of
reason, would not have looked forward to distant contingencies. They
would have been governed more by their instincts and even less by their
reason than are savages at the present day. They would not at that period
have partially lost one of the strongest of all instincts, common to all the
lower animals, namely the love of their young offspring; and consequently
they would not have practised infanticide. There would have been no
artificial scarcity of women, and polyandry would not have been followed;
there would have been no early betrothals; women would not have been
valued as mere slaves; both sexes, if the females as well as the males were
permitted to exert any choice, would have chosen their partners, not for
mental charms, or property, or social position, but almost solely from
external appearance. All the adults would have married or paired, and all
the offspring, as far as that was possible, would have been reared; so that
the struggle for existence would have been periodically severe to an
extreme degree. Thus during these primordial times all the conditions for
sexual selection would have been much more favourable than at a later
period, when man had advanced in his intellectual powers, but had
retrograded in his instincts. Therefore, whatever influence sexual selection
may have had in producing the differences between the races of man, and
between man and the higher Quadrumana, this influence would have been
much more powerful at a very remote period than at the present day.

On the Manner of Action of Sexual Selection with mankind.—With primeval
men under the favourable conditions just stated, and with those savages
who at the present time enter into any marriage tie (but subject to greater or
less interference according as the habits of female infanticide, early
betrothals, &c., are more or less practised), sexual selection will probably
have acted in the following manner. The strongest and most vigorous men,
—those who could best defend and hunt for their families, and during later
times the chiefs or head-men,—those who were provided with the best
weapons and who possessed the most property, such as a larger number of
dogs or other animals, would have succeeded in rearing a greater average
number of offspring, than would the weaker, poorer and lower members of
the same tribes. There can, also, be no doubt that such men would generally

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have been able to select the more attractive women. At present the chiefs of
nearly every tribe throughout the world succeed in obtaining more than one
wife. Until recently, as I hear from Mr. Mantell, almost every girl in New
Zealand, who was pretty, or promised to be pretty, was tapu to some chief.
With the Kafirs, as Mr. C. Hamilton states,462 “the chiefs generally have the
pick of the women for many miles round, and are most persevering in
establishing or confirming their privilege.” We have seen that each race has
its own style of beauty, and we know that it is natural to man to admire each
characteristic point in his domestic animals, dress, ornaments, and personal
appearance, when carried a little beyond the common standard. If then the
several foregoing propositions be admitted, and I cannot see that they are
doubtful, it would be an inexplicable circumstance, if the selection of the
more attractive women by the more powerful men of each tribe, who would
rear on an average a greater number of children, did not after the lapse of
many generations modify to a certain extent the character of the tribe.
With our domestic animals, when a foreign breed is introduced into a new
country, or when a native breed is long and carefully attended to, either for
use or ornament, it is found after several generations to have undergone,
whenever the means of comparison exist, a greater or less amount of
change. This follows from unconscious selection during a long series of
generations—that is, the preservation of the most approved individuals—
without any wish or expectation of such a result on the part of the breeder.
So again, if two careful breeders rear during many years animals of the
same family, and do not compare them together or with a common standard,
the animals are found after a time to have become to the surprise of their
owners slightly different.463 Each breeder has impressed, as Von Nathusius
well expresses it, the character of his own mind—his own taste and
judgment—on his animals. What reason, then, can be assigned why similar
results should not follow from the long-continued selection of the most
admired women by those men of each tribe, who were able to rear to
maturity the greater number of children? This would be unconscious
selection, for an effect would be produced, independently of any wish or
expectation on the part of the men who preferred certain women to others.
Let us suppose the members of a tribe, in which some form of marriage was
practised, to spread over an unoccupied continent; they would soon split up
into distinct hordes, which would be separated from each other by various

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barriers, and still more effectually by the incessant wars between all
barbarous nations. The hordes would thus be exposed to slightly different
conditions and habits of life, and would sooner or later come to differ in
some small degree. As soon as this occurred, each isolated tribe would form
for itself a slightly different standard of beauty;464 and then unconscious
selection would come into action through the more powerful and leading
savages preferring certain women to others. Thus the differences between
the tribes, at first very slight, would gradually and inevitably be increased to
a greater and greater degree.

With animals in a state of nature, many characters proper to the males, such
as size, strength, special weapons, courage and pugnacity, have been
acquired through the law of battle. The semi-human progenitors of man,
like their allies the Quadrumana, will almost certainly have been thus
modified; and, as savages still fight for the possession of their women, a
similar process of selection has probably gone on in a greater or less degree
to the present day. Other characters proper to the males of the lower
animals, such as bright colours and various ornaments, have been acquired
by the more attractive males having been preferred by the females. There
are, however, exceptional cases in which the males, instead of having been
the selected, have been the selectors. We recognise such cases by the
females having been rendered more highly ornamented than the males,—
their ornamental characters having been transmitted exclusively or chiefly
to their female offspring. One such case has been described in the order to
which man belongs, namely, with the Rhesus monkey.
Man is more powerful in body and mind than woman, and in the savage
state he keeps her in a far more abject state of bondage than does the male
of any other animal; therefore it is not surprising that he should have gained
the power of selection. Women are everywhere conscious of the value of
their beauty; and when they have the means, they take more delight in
decorating themselves with all sorts of ornaments than do men. They
borrow the plumes of male birds, with which nature decked this sex in order
to charm the females. As women have long been selected for beauty, it is
not surprising that some of the successive variations should have been
transmitted in a limited manner; and consequently that women should have
transmitted their beauty in a somewhat higher degree to their female than to

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their male offspring. Hence women have become more beautiful, as most
persons will admit, than men. Women, however, certainly transmit most of
their characters, including beauty, to their offspring of both sexes; so that
the continued preference by the men of each race of the more attractive
women, according to their standard of taste, would tend to modify in the
same manner all the individuals of both sexes belonging to the race.
With respect to the other form of sexual selection (which with the lower
animals is much the most common), namely, when the females are the
selectors, and accept only those males which excite or charm them most, we
have reason to believe that it formerly acted on the progenitors of man. Man
in all probability owes his beard, and perhaps some other characters, to
inheritance from an ancient progenitor who gained in this manner his
ornaments. But this form of selection may have occasionally acted during
later times; for in utterly barbarous tribes the women have more power in
choosing, rejecting, and tempting their lovers, or of afterwards changing
their husbands, than might have been expected. As this is a point of some
importance, I will give in detail such evidence as I have been able to
collect.
Hearne describes how a woman in one of the tribes of Arctic America
repeatedly ran away from her husband and joined a beloved man; and with
the Charruas of S. America, as Azara states, the power of divorce is
perfectly free. With the Abipones, when a man chooses a wife he bargains
with the parents about the price. But “it frequently happens that the girl
rescinds what has been agreed upon between the parents and the
bridegroom, obstinately rejecting the very mention of marriage.” She often
runs away, hides herself, and thus eludes the bridegroom. In the Fiji Islands
the man seizes on the woman whom he wishes for his wife by actual or
pretended force; but “on reaching the home of her abductor, should she not
approve of the match, she runs to some one who can protect her; if,
however, she is satisfied, the matter is settled forthwith.” In Tierra del
Fuego a young man first obtains the consent of the parents by doing them
some service, and then he attempts to carry off the girl; “but if she is
unwilling, she hides herself in the woods until her admirer is heartily tired
of looking for her, and gives up the pursuit; but this seldom happens.” With
the Kalmucks there is a regular race between the bride and bridegroom, the
former having a fair start; and Clarke “was assured that no instance occurs

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of a girl being caught, unless she has a partiality to the pursuer.” So with the
wild tribes of the Malay archipelago there is a similar racing match; and it
appears from M. Bourien’s account, as Sir J. Lubbock remarks, that “the
race ‘is not to the swift, nor the battle to the strong,’ but to the young man
who has the good fortune to please his intended bride.”
Turning to Africa: the Kafirs buy their wives, and girls are severely beaten
by their fathers if they will not accept a chosen husband; yet it is manifest
from many facts given by the Rev. Mr. Shooter, that they have considerable
power of choice. Thus very ugly, though rich men, have been known to fail
in getting wives. The girls, before consenting to be betrothed, compel the
men to shew themselves off, first in front and then behind, and “exhibit
their paces.” They have been known to propose to a man, and they not
rarely run away with a favoured lover. With the degraded bushwomen of S.
Africa, “when a girl has grown up to womanhood without having been
betrothed, which, however, does not often happen, her lover must gain her
approbation, as well as that of the parents.”465 Mr. Winwood Reade made
inquiries for me with respect to the negroes of Western Africa, and he
informs me that “the women, at least among the more intelligent Pagan
tribes, have no difficulty in getting the husbands whom they may desire,
although it is considered unwomanly to ask a man to marry them. They are
quite capable of falling in love, and of forming tender, passionate, and
faithful attachments.”
We thus see that with savages the women are not in quite so abject a state in
relation to marriage as has often been supposed. They can tempt the men
whom they prefer, and can sometimes reject those whom they dislike, either
before or after marriage. Preference on the part of the women, steadily
acting in any one direction, would ultimately affect the character of the
tribe; for the women would generally choose not merely the handsomer
men, according to their standard of taste, but those who were at the same
time best able to defend and support them. Such well-endowed pairs would
commonly rear a larger number of offspring than the less well endowed.
The same result would obviously follow in a still more marked manner if
there was selection on both sides; that is if the more attractive, and at the
same time more powerful men were to prefer, and were preferred by, the
more attractive women. And these two forms of selection seem actually to

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have occurred, whether or not simultaneously, with mankind, especially
during the earlier periods of our long history.
We will now consider in a little more detail, relatively to sexual selection,
some of the characters which distinguish the several races of man from each
other and from the lower animals, namely, the more or less complete
absence of hair from the body and the colour of the skin. We need say
nothing about the great diversity in the shape of the features and of the skull
between the different races, as we have seen in the last chapter how
different is the standard of beauty in these respects. These characters will
therefore probably have been acted on through sexual selection; but we
have no means of judging, as far as I can see, whether they have been acted
on chiefly through the male or female side. The musical faculties of man
have likewise been already discussed.

Absence of Hair on the Body, and its Development on the Face and Head.
—From the presence of the woolly hair or lanugo on the human fœtus, and
of rudimentary hairs scattered over the body during maturity, we may infer
that man is descended from some animal which was born hairy and
remained so during life. The loss of hair is an inconvenience and probably
an injury to man even under a hot climate, for he is thus exposed to sudden
chills, especially during wet weather. As Mr. Wallace remarks, the natives in
all countries are glad to protect their naked backs and shoulders with some
slight covering. No one supposes that the nakedness of the skin is any direct
advantage to man, so that his body cannot have been divested of hair
through natural selection.466 Nor have we any grounds for believing, as
shewn in a former chapter, that this can be due to the direct action of the
conditions to which man has long been exposed, or that it is the result of
correlated development.
The absence of hair on the body is to a certain extent a secondary sexual
character; for in all parts of the world women are less hairy than men.
Therefore we may reasonably suspect that this is a character which has been
gained through sexual selection. We know that the faces of several species
of monkeys, and large surfaces at the posterior end of the body in other
species, have been denuded of hair; and this we may safely attribute to
sexual selection, for these surfaces are not only vividly coloured, but

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sometimes, as with the male mandrill and female rhesus, much more vividly
in the one sex than in the other. As these animals gradually reach maturity
the naked surfaces, as I am informed by Mr. Bartlett, grow larger, relatively
to the size of their bodies. The hair, however, appears to have been removed
in these cases, not for the sake of nudity, but that the colour of the skin
should be more fully displayed. So again with many birds the head and
neck have been divested of feathers through sexual selection, for the sake of
exhibiting the brightly-coloured skin.
As woman has a less hairy body than man, and as this character is common
to all races, we may conclude that our female semi-human progenitors were
probably first partially divested of hair; and that this occurred at an
extremely remote period before the several races had diverged from a
common stock. As our female progenitors gradually acquired this new
character of nudity, they must have transmitted it in an almost equal degree
to their young offspring of both sexes; so that its transmission, as in the case
of many ornaments with mammals and birds, has not been limited either by
age or sex. There is nothing surprising in a partial loss of hair having been
esteemed as ornamental by the ape-like progenitors of man, for we have
seen that with animals of all kinds innumerable strange characters have
been thus esteemed, and have consequently been modified through sexual
selection. Nor is it surprising that a character in a slight degree injurious
should have been thus acquired; for we know that this is the case with the
plumes of some birds, and with the horns of some stags.
The females of certain anthropoid apes, as stated in a former chapter, are
somewhat less hairy on the under surface than are the males; and here we
have what might have afforded a commencement for the process of
denudation. With respect to the completion of the process through sexual
selection, it is well to bear in mind the New Zealand proverb, “there is no
woman for a hairy man.” All who have seen photographs of the Siamese
hairy family will admit how ludicrously hideous is the opposite extreme of
excessive hairiness. Hence the king of Siam had to bribe a man to marry the
first hairy woman in the family, who transmitted this character to her young
offspring of both sexes.467
Some races are much more hairy than others, especially on the male side;
but it must not be assumed that the more hairy races, for instance

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Europeans, have retained a primordial condition more completely than have
the naked races, such as the Kalmucks or Americans. It is a more probable
view that the hairiness of the former is due to partial reversion, for
characters which have long been inherited are always apt to return. It does
not appear that a cold climate has been influential in leading to this kind of
reversion; excepting perhaps with the negroes, who have been reared during
several generations, in the United States,468 and possibly with the Ainos,
who inhabit the northern islands of the Japan archipelago. But the laws of
inheritance are so complex than we can seldom understand their action. If
the greater hairiness of certain races be the result of reversion, unchecked
by any form of selection, the extreme variability of this character, even
within the limits of the same race, ceases to be remarkable.
With respect to the beard, if we turn to our best guide, namely the
Quadrumana, we find beards equally well developed in both sexes of many
species, but in others, either confined to the males, or more developed in
them than in the females. From this fact, and from the curious arrangement,
as well as the bright colours, of the hair about the heads of many monkeys,
it is highly probable, as before explained, that the males first acquired their
beards as an ornament through sexual selection, transmitting them in most
cases, in an equal or nearly equal degree, to their offspring of both sexes.
We know from Eschricht469 that with mankind, the female as well as the
male fœtus is furnished with much hair on the face, especially round the
mouth; and this indicates that we are descended from a progenitor, of which
both sexes were bearded. It appears therefore at first sight probable that
man has retained his beard from a very early period, whilst woman lost her
beard at the same time when her body became almost completely divested
of hair. Even the colour of the beard with mankind seems to have been
inherited from an ape-like progenitor; for when there is any difference in
tint between the hair of the head and the beard, the latter is lighter coloured
in all monkeys and in man. There is less improbability in the men of the
bearded races having retained their beards from primordial times, than in
the case of the hair on the body; for with those Quadrumana, in which the
male has a larger beard than that of the female, it is fully developed only at
maturity, and the later stages of development may have been exclusively
transmitted to mankind. We should then see what is actually the case,
namely, our male children, before they arrive at maturity, as destitute of

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beards as are our female children. On the other hand the great variability of
the beard within the limits of the same race and in different races indicates
that reversion has come into action. However this may be, we must not
overlook the part which sexual selection may have played even during later
times; for we know that with savages, the men of the beardless races take
infinite pains in eradicating every hair from their faces, as something
odious, whilst the men of the bearded races feel the greatest pride in their
beards. The women, no doubt, participate in these feelings, and if so sexual
selection can hardly have failed to have effected something in the course of
later times.470
It is rather difficult to form a judgment how the long hair on our heads
became developed. Eschricht 471 states that in the human fœtus the hair on
the face during the fifth month is longer than that on the head; and this
indicates that our semi-human progenitors were not furnished with long
tresses, which consequently must have been a late acquisition. This is
likewise indicated by the extraordinary difference in the length of the hair in
the different races; in the negro the hair forms a mere curly mat; with us it is
of great length, and with the American natives it not rarely reaches to the
ground. Some species of Semnopithecus have their heads covered with
moderately long hair, and this probably serves as an ornament and was
acquired through sexual selection. The same view may be extended to
mankind, for we know that long tresses are now and were formerly much
admired, as may be observed in the works of almost every poet; St. Paul
says, “if a woman have long hair, it is a glory to her;” and we have seen that
in North America a chief was elected solely from the length of his hair.

Colour of the Skin.—The best kind of evidence that the colour of the skin
has been modified through sexual selection is wanting in the case of
mankind; for the sexes do not differ in this respect, or only slightly and
doubtfully. On the other hand we know from many facts already given that
the colour of the skin is regarded by the men of all races as a highly
important element in their beauty; so that it is a character which would be
likely to be modified through selection, as has occurred in innumerable
instances with the lower animals. It seems at first sight a monstrous
supposition that the jet blackness of the negro has been gained through
sexual selection; but this view is supported by various analogies, and we

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know that negroes admire their own blackness. With mammals, when the
sexes differ in colour, the male is often black or much darker than the
female; and it depends merely on the form of inheritance whether this or
any other tint shall be transmitted to both sexes or to one alone. The
resemblance of Pithecia satanas with his jet black skin, white rolling
eyeballs, and hair parted on the top of the head, to a negro in miniature, is
almost ludicrous.
The colour of the face differs much more widely in the various kinds of
monkeys than it does in the races of man; and we have good reason to
believe that the red, blue, orange, almost white and black tints of their skin,
even when common to both sexes, and the bright colours of their fur, as
well as the ornamental tufts of hair about the head, have all been acquired
through sexual selection. As the newly-born infants of the most distinct
races do not differ nearly as much in colour as do the adults, although their
bodies are completely destitute of hair, we have some slight indication that
the tints of the different races were acquired subsequently to the removal of
the hair, which, as before stated, must have occurred at a very early period.

Summary.—We may conclude that the greater size, strength, courage,
pugnacity, and even energy of man, in comparison with the same qualities
in woman, were acquired during primeval times, and have subsequently
been augmented, chiefly through the contests of rival males for the
possession of the females. The greater intellectual vigour and power of
invention in man is probably due to natural selection combined with the
inherited effects of habit, for the most able men will have succeeded best in
defending and providing for themselves, their wives and offspring. As far as
the extreme intricacy of the subject permits us to judge, it appears that our
male ape-like progenitors acquired their beards as an ornament to charm or
excite the opposite sex, and transmitted them to man as he now exists. The
females apparently were first denuded of hair in like manner as a sexual
ornament; but they transmitted this character almost equally to both sexes.
It is not improbable that the females were modified in other respects for the
same purpose and through the same means; so that women have acquired
sweeter voices and become more beautiful than men.

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It deserves particular attention that with mankind all the conditions for
sexual selection were much more favourable, during a very early period,
when man had only just attained to the rank of manhood, than during later
times. For he would then, as we may safely conclude, have been guided
more by his instinctive passions, and less by foresight or reason. He would
not then have been so utterly licentious as many savages now are; and each
male would have jealously guarded his wife or wives. He would not then
have practised infanticide; nor valued his wives merely as useful slaves; nor
have been betrothed to them during infancy. Hence we may infer that the
races of men were differentiated, as far as sexual selection is concerned, in
chief part during a very remote epoch; and this conclusion throws light on
the remarkable fact that at the most ancient period, of which we have as yet
obtained any record, the races of man had already come to differ nearly or
quite as much as they do at the present day.
The views here advanced, on the part which sexual selection has played in
the history of man, want scientific precision. He who does not admit this
agency in the case of the lower animals, will properly disregard all that I
have written in the later chapters on man. We cannot positively say that this
character, but not that, has been thus modified; it has, however, been shewn
that the races of man differ from each other and from their nearest allies
amongst the lower animals, in certain characters which are of no service to
them in their ordinary habits of life, and which it is extremely probable
would have been modified through sexual selection. We have seen that with
the lowest savages the people of each tribe admire their own characteristic
qualities,—the shape of the head and face, the squareness of the cheek-
bones, the prominence or depression of the nose, the colour of the skin, the
length of the hair on the head, the absence of hair on the face and body, or
the presence of a great beard, and so forth. Hence these and other such
points could hardly fail to have been slowly and gradually exaggerated from
the more powerful and able men in each tribe, who would succeed in
rearing the largest number of offspring, having selected during many
generations as their wives the most strongly characterised and therefore
most attractive women. For my own part I conclude that of all the causes
which have led to the differences in external appearance between the races
of man, and to a certain extent between man and the lower animals, sexual
selection has been by far the most efficient.

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CHAPTER XXI.

General Summary and Conclusion.
Main conclusion that man is descended from some lower form—Manner of development—
Genealogy of man—Intellectual and moral faculties—Sexual selection—Concluding remarks.

A brief summary will here be sufficient to recall to the reader’s mind the
more salient points in this work. Many of the views which have been
advanced are highly speculative, and some no doubt will prove erroneous;
but I have in every case given the reasons which have led me to one view
rather than to another. It seemed worth while to try how far the principle of
evolution would throw light on some of the more complex problems in the
natural history of man. False facts are highly injurious to the progress of
science, for they often long endure; but false views, if supported by some
evidence, do little harm, as every one takes a salutary pleasure in proving
their falseness; and when this is done, one path towards error is closed and
the road to truth is often at the same time opened.
The main conclusion arrived at in this work, and now held by many
naturalists who are well competent to form a sound judgment, is that man is
descended from some less highly organised form. The grounds upon which
this conclusion rests will never be shaken, for the close similarity between
man and the lower animals in embryonic development, as well as in
innumerable points of structure and constitution, both of high and of the
most trifling importance,—the rudiments which he retains, and the
abnormal reversions to which he is occasionally liable,—are facts which
cannot be disputed. They have long been known, but until recently they told
us nothing with respect to the origin of man. Now when viewed by the light
of our knowledge of the whole organic world, their meaning is
unmistakeable. The great principle of evolution stands up clear and firm,
when these groups of facts are considered in connection with others, such as
the mutual affinities of the members of the same group, their geographical
distribution in past and present times, and their geological succession. It is
incredible that all these facts should speak falsely. He who is not content to
look, like a savage, at the phenomena of nature as disconnected, cannot any

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longer believe that man is the work of a separate act of creation. He will be
forced to admit that the close resemblance of the embryo of man to that, for
instance, of a dog—the construction of his skull, limbs, and whole frame,
independently of the uses to which the parts may be put, on the same plan
with that of other mammals—the occasional reappearance of various
structures, for instance of several distinct muscles, which man does not
normally possess, but which are common to the Quadrumana—and a crowd
of analogous facts—all point in the plainest manner to the conclusion that
man is the co-descendant with other mammals of a common progenitor.
We have seen that man incessantly presents individual differences in all
parts of his body and in his mental faculties. These differences or variations
seem to be induced by the same general causes, and to obey the same laws
as with the lower animals. In both cases similar laws of inheritance prevail.
Man tends to increase at a greater rate than his means of subsistence;
consequently he is occasionally subjected to a severe struggle for existence,
and natural selection will have effected whatever lies within its scope. A
succession of strongly-marked variations of a similar nature are by no
means requisite; slight fluctuating differences in the individual suffice for
the work of natural selection. We may feel assured that the inherited effects
of the long-continued use or disuse of parts will have done much in the
same direction with natural selection. Modifications formerly of
importance, though no longer of any special use, will be long inherited.
When one part is modified, other parts will change through the principle of
correlation, of which we have instances in many curious cases of correlated
monstrosities. Something may be attributed to the direct and definite action
of the surrounding conditions of life, such as abundant food, heat, or
moisture; and lastly, many characters of slight physiological importance,
some indeed of considerable importance, have been gained through sexual
selection.
No doubt man, as well as every other animal, presents structures, which as
far as we can judge with our little knowledge, are not now of any service to
him, nor have been so during any former period of his existence, either in
relation to his general conditions of life, or of one sex to the other. Such
structures cannot be accounted for by any form of selection, or by the
inherited effects of the use and disuse of parts. We know, however, that
many strange and strongly-marked peculiarities of structure occasionally

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appear in our domesticated productions, and if the unknown causes which
produce them were to act more uniformly, they would probably become
common to all the individuals of the species. We may hope hereafter to
understand something about the causes of such occasional modifications,
especially through the study of monstrosities: hence the labours of
experimentalists, such as those of M. Camille Dareste, are full of promise
for the future. In the greater number of cases we can only say that the cause
of each slight variation and of each monstrosity lies much more in the
nature or constitution of the organism, than in the nature of the surrounding
conditions; though new and changed conditions certainly play an important
part in exciting organic changes of all kinds.
Through the means just specified, aided perhaps by others as yet
undiscovered, man has been raised to his present state. But since he attained
to the rank of manhood, he has diverged into distinct races, or as they may
be more appropriately called sub-species. Some of these, for instance the
Negro and European, are so distinct that, if specimens had been brought to a
naturalist without any further information, they would undoubtedly have
been considered by him as good and true species. Nevertheless all the races
agree in so many unimportant details of structure and in so many mental
peculiarities, that these can be accounted for only through inheritance from
a common progenitor; and a progenitor thus characterised would probably
have deserved to rank as man.
It must not be supposed that the divergence of each race from the other
races, and of all the races from a common stock, can be traced back to any
one pair of progenitors. On the contrary, at every stage in the process of
modification, all the individuals which were in any way best fitted for their
conditions of life, though in different degrees, would have survived in
greater numbers than the less well fitted. The process would have been like
that followed by man, when he does not intentionally select particular
individuals, but breeds from all the superior and neglects all the inferior
individuals. He thus slowly but surely modifies his stock, and
unconsciously forms a new strain. So with respect to modifications,
acquired independently of selection, and due to variations arising from the
nature of the organism and the action of the surrounding conditions, or from
changed habits of life, no single pair will have been modified in a much

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greater degree than the other pairs which inhabit the same country, for all
will have been continually blended through free intercrossing.
By considering the embryological structure of man,—the homologies which
he presents with the lower animals,—the rudiments which he retains,—and
the reversions to which he is liable, we can partly recall in imagination the
former condition of our early progenitors; and can approximately place
them in their proper position in the zoological series. We thus learn that
man is descended from a hairy quadruped, furnished with a tail and pointed
ears, probably arboreal in its habits, and an inhabitant of the Old World.
This creature, if its whole structure had been examined by a naturalist,
would have been classed amongst the Quadrumana, as surely as would the
common and still more ancient progenitor of the Old and New World
monkeys. The Quadrumana and all the higher mammals are probably
derived from an ancient marsupial animal, and this through a long line of
diversified forms, either from some reptile-like or some amphibian-like
creature, and this again from some fish-like animal. In the dim obscurity of
the past we can see that the early progenitor of all the Vertebrata must have
been an aquatic animal, provided with branchiæ, with the two sexes united
in the same individual, and with the most important organs of the body
(such as the brain and heart) imperfectly developed. This animal seems to
have been more like the larvæ of our existing marine Ascidians than any
other known form.

The greatest difficulty which presents itself, when we are driven to the
above conclusion on the origin of man, is the high standard of intellectual
power and of moral disposition which he has attained. But every one who
admits the general principle of evolution, must see that the mental powers
of the higher animals, which are the same in kind with those of mankind,
though so different in degree, are capable of advancement. Thus the interval
between the mental powers of one of the higher apes and of a fish, or
between those of an ant and scale-insect, is immense. The development of
these powers in animals does not offer any special difficulty; for with our
domesticated animals, the mental faculties are certainly variable, and the
variations are inherited. No one doubts that these faculties are of the utmost
importance to animals in a state of nature. Therefore the conditions are
favourable for their development through natural selection. The same

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conclusion may be extended to man; the intellect must have been all-
important to him, even at a very remote period, enabling him to use
language, to invent and make weapons, tools, traps, &c.; by which means,
in combination with his social habits, he long ago became the most
dominant of all living creatures.
A great stride in the development of the intellect will have followed, as
soon as, through a previous considerable advance, the half-art and half-
instinct of language came into use; for the continued use of language will
have reacted on the brain, and produced an inherited effect; and this again
will have reacted on the improvement of language. The large size of the
brain in man, in comparison with that of the lower animals, relatively to the
size of their bodies, may be attributed in chief part, as Mr. Chauncey Wright
has well remarked,472 to the early use of some simple form of language,—
that wonderful engine which affixes signs to all sorts of objects and
qualities, and excites trains of thought which would never arise from the
mere impression of the senses, and if they did arise could not be followed
out. The higher intellectual powers of man, such as those of ratiocination,
abstraction, self-consciousness, &c., will have followed from the continued
improvement of other mental faculties; but without considerable culture of
the mind, both in the race and in the individual, it is doubtful whether these
high powers would be exercised, and thus fully attained.
The development of the moral qualities is a more interesting and difficult
problem. Their foundation lies in the social instincts, including in this term
the family ties. These instincts are of a highly complex nature, and in the
case of the lower animals give special tendencies towards certain definite
actions; but the more important elements for us are love, and the distinct
emotion of sympathy. Animals endowed with the social instincts take
pleasure in each other’s company, warn each other of danger, defend and
aid each other in many ways. These instincts are not extended to all the
individuals of the species, but only to those of the same community. As they
are highly beneficial to the species, they have in all probability been
acquired through natural selection.
A moral being is one who is capable of comparing his past and future
actions and motives,—of approving of some and disapproving of others;
and the fact that man is the one being who with certainty can be thus

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designated makes the greatest of all distinctions between him and the lower
animals. But in our third chapter I have endeavoured to shew that the moral
sense follows, firstly, from the enduring and always present nature of the
social instincts, in which respect man agrees with the lower animals; and
secondly, from his mental faculties being highly active and his impressions
of past events extremely vivid, in which respects he differs from the lower
animals. Owing to this condition of mind, man cannot avoid looking
backwards and comparing the impressions of past events and actions. He
also continually looks forward. Hence after some temporary desire or
passion has mastered his social instincts, he will reflect and compare the
now weakened impression of such past impulses, with the ever present
social instinct; and he will then feel that sense of dissatisfaction which all
unsatisfied instincts leave behind them. Consequently he resolves to act
differently for the future—and this is conscience. Any instinct which is
permanently stronger or more enduring than another, gives rise to a feeling
which we express by saying that it ought to be obeyed. A pointer dog, if
able to reflect on his past conduct, would say to himself, I ought (as indeed
we say of him) to have pointed at that hare and not have yielded to the
passing temptation of hunting it.
Social animals are partly impelled by a wish to aid the members of the same
community in a general manner, but more commonly to perform certain
definite actions. Man is impelled by the same general wish to aid his
fellows, but has few or no special instincts. He differs also from the lower
animals in being able to express his desires by words, which thus become
the guide to the aid required and bestowed. The motive to give aid is
likewise somewhat modified in man: it no longer consists solely of a blind
instinctive impulse, but is largely influenced by the praise or blame of his
fellow men. Both the appreciation and the bestowal of praise and blame rest
on sympathy; and this emotion, as we have seen, is one of the most
important elements of the social instincts. Sympathy, though gained as an
instinct, is also much strengthened by exercise or habit. As all men desire
their own happiness, praise or blame is bestowed on actions and motives,
according as they lead to this end; and as happiness is an essential part of
the general good, the greatest-happiness principle indirectly serves as a
nearly safe standard of right and wrong. As the reasoning powers advance
and experience is gained, the more remote effects of certain lines of conduct

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on the character of the individual, and on the general good, are perceived;
and then the self-regarding virtues, from coming within the scope of public
opinion, receive praise, and their opposites receive blame. But with the less
civilised nations reason often errs, and many bad customs and base
superstitions come within the same scope, and consequently are esteemed
as high virtues, and their breach as heavy crimes.
The moral faculties are generally esteemed, and with justice, as of higher
value than the intellectual powers. But we should always bear in mind that
the activity of the mind in vividly recalling past impressions is one of the
fundamental though secondary bases of conscience. This fact affords the
strongest argument for educating and stimulating in all possible ways the
intellectual faculties of every human being. No doubt a man with a torpid
mind, if his social affections and sympathies are well developed, will be led
to good actions, and may have a fairly sensitive conscience. But whatever
renders the imagination of men more vivid and strengthens the habit of
recalling and comparing past impressions, will make the conscience more
sensitive, and may even compensate to a certain extent for weak social
affections and sympathies.
The moral nature of man has reached the highest standard as yet attained,
partly through the advancement of the reasoning powers and consequently
of a just public opinion, but especially through the sympathies being
rendered more tender and widely diffused through the effects of habit,
example, instruction, and reflection. It is not improbable that virtuous
tendencies may through long practice be inherited. With the more civilised
races, the conviction of the existence of an all-seeing Deity has had a potent
influence on the advancement of morality. Ultimately man no longer
accepts the praise or blame of his fellows as his chief guide, though few
escape this influence, but his habitual convictions controlled by reason
afford him the safest rule. His conscience then becomes his supreme judge
and monitor. Nevertheless the first foundation or origin of the moral sense
lies in the social instincts, including sympathy; and these instincts no doubt
were primarily gained, as in the case of the lower animals, through natural
selection.

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The belief in God has often been advanced as not only the greatest, but the
most complete of all the distinctions between man and the lower animals. It
is however impossible, as we have seen, to maintain that this belief is innate
or instinctive in man. On the other hand a belief in all-pervading spiritual
agencies seems to be universal; and apparently follows from a considerable
advance in the reasoning powers of man, and from a still greater advance in
his faculties of imagination, curiosity and wonder. I am aware that the
assumed instinctive belief in God has been used by many persons as an
argument for His existence. But this is a rash argument, as we should thus
be compelled to believe in the existence of many cruel and malignant
spirits, possessing only a little more power than man; for the belief in them
is far more general than of a beneficent Deity. The idea of a universal and
beneficent Creator of the universe does not seem to arise in the mind of
man, until he has been elevated by long-continued culture.
He who believes in the advancement of man from some lowly-organised
form, will naturally ask how does this bear on the belief in the immortality
of the soul. The barbarous races of man, as Sir J. Lubbock has shewn,
possess no clear belief of this kind; but arguments derived from the
primeval beliefs of savages are, as we have just seen, of little or no avail.
Few persons feel any anxiety from the impossibility of determining at what
precise period in the development of the individual, from the first trace of
the minute germinal vesicle to the child either before or after birth, man
becomes an immortal being; and there is no greater cause for anxiety
because the period in the gradually ascending organic scale cannot possibly
be determined.473
I am aware that the conclusions arrived at in this work will be denounced
by some as highly irreligious; but he who thus denounces them is bound to
shew why it is more irreligious to explain the origin of man as a distinct
species by descent from some lower form, through the laws of variation and
natural selection, than to explain the birth of the individual through the laws
of ordinary reproduction. The birth both of the species and of the individual
are equally parts of that grand sequence of events, which our minds refuse
to accept as the result of blind chance. The understanding revolts at such a
conclusion, whether or not we are able to believe that every slight variation
of structure,—the union of each pair in marriage,—the dissemination of

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each seed,—and other such events, have all been ordained for some special
purpose.

Sexual selection has been treated at great length in these volumes; for, as I
have attempted to shew, it has played an important part in the history of the
organic world. As summaries have been given to each chapter, it would be
superfluous here to add a detailed summary. I am aware that much remains
doubtful, but I have endeavoured to give a fair view of the whole case. In
the lower divisions of the animal kingdom, sexual selection seems to have
done nothing: such animals are often affixed for life to the same spot, or
have the two sexes combined in the same individual, or what is still more
important, their perceptive and intellectual faculties are not sufficiently
advanced to allow of the feelings of love and jealousy, or of the exertion of
choice. When, however, we come to the Arthropoda and Vertebrata, even to
the lowest classes in these two great Sub-Kingdoms, sexual selection has
effected much; and it deserves notice that we here find the intellectual
faculties developed, but in two very distinct lines, to the highest standard,
namely in the Hymenoptera (ants, bees, &c.) amongst the Arthropoda, and
in the Mammalia, including man, amongst the Vertebrata.
In the most distinct classes of the animal kingdom, with mammals, birds,
reptiles, fishes, insects, and even crustaceans, the differences between the
sexes follow almost exactly the same rules. The males are almost always
the wooers; and they alone are armed with special weapons for fighting
with their rivals. They are generally stronger and larger than the females,
and are endowed with the requisite qualities of courage and pugnacity. They
are provided, either exclusively or in a much higher degree than the
females, with organs for producing vocal or instrumental music, and with
odoriferous glands. They are ornamented with infinitely diversified
appendages, and with the most brilliant or conspicuous colours, often
arranged in elegant patterns, whilst the females are left unadorned. When
the sexes differ in more important structures, it is the male which is
provided with special sense-organs for discovering the female, with
locomotive organs for reaching her, and often with prehensile organs for
holding her. These various structures for securing or charming the female
are often developed in the male during only part of the year, namely the
breeding season. They have in many cases been transferred in a greater or

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less degree to the females; and in the latter case they appear in her as mere
rudiments. They are lost by the males after emasculation. Generally they are
not developed in the male during early youth, but appear a short time before
the age for reproduction. Hence in most cases the young of both sexes
resemble each other; and the female resembles her young offspring
throughout life. In almost every great class a few anomalous cases occur in
which there has been an almost complete transposition of the characters
proper to the two sexes; the females assuming characters which properly
belong to the males. This surprising uniformity in the laws regulating the
differences between the sexes in so many and such widely separated
classes, is intelligible if we admit the action throughout all the higher
divisions of the animal kingdom of one common cause, namely sexual
selection.
Sexual selection depends on the success of certain individuals over others
of the same sex in relation to the propagation of the species; whilst natural
selection depends on the success of both sexes, at all ages, in relation to the
general conditions of life. The sexual struggle is of two kinds; in the one it
is between the individuals of the same sex, generally the male sex, in order
to drive away or kill their rivals, the females remaining passive; whilst in
the other, the struggle is likewise between the individuals of the same sex,
in order to excite or charm those of the opposite sex, generally the females,
which no longer remain passive, but select the more agreeable partners.
This latter kind of selection is closely analogous to that which man
unintentionally, yet effectually, brings to bear on his domesticated
productions, when he continues for a long time choosing the most pleasing
or useful individuals, without any wish to modify the breed.
The laws of inheritance determine whether characters gained through sexual
selection by either sex shall be transmitted to the same sex, or to both sexes;
as well as the age at which they shall be developed. It appears that
variations which arise late in life are commonly transmitted to one and the
same sex. Variability is the necessary basis for the action of selection, and is
wholly independent of it. It follows from this, that variations of the same
general nature have often been taken advantage of and accumulated through
sexual selection in relation to the propagation of the species, and through
natural selection in relation to the general purposes of life. Hence secondary
sexual characters, when equally transmitted to both sexes can be

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distinguished from ordinary specific characters only by the light of analogy.
The modifications acquired through sexual selection are often so strongly
pronounced that the two sexes have frequently been ranked as distinct
species, or even as distinct genera. Such strongly-marked differences must
be in some manner highly important; and we know that they have been
acquired in some instances at the cost not only of inconvenience, but of
exposure to actual danger.
The belief in the power of sexual selection rests chiefly on the following
considerations. The characters which we have the best reason for supposing
to have been thus acquired are confined to one sex; and this alone renders it
probable that they are in some way connected with the act of reproduction.
These characters in innumerable instances are fully developed only at
maturity; and often during only a part of the year, which is always the
breeding-season. The males (passing over a few exceptional cases) are the
most active in courtship; they are the best armed, and are rendered the most
attractive in various ways. It is to be especially observed that the males
display their attractions with elaborate care in the presence of the females;
and that they rarely or never display them excepting during the season of
love. It is incredible that all this display should be purposeless. Lastly we
have distinct evidence with some quadrupeds and birds that the individuals
of the one sex are capable of feeling a strong antipathy or preference for
certain individuals of the opposite sex.
Bearing these facts in mind, and not forgetting the marked results of man’s
unconscious selection, it seems to me almost certain that if the individuals
of one sex were during a long series of generations to prefer pairing with
certain individuals of the other sex, characterised in some peculiar manner,
the offspring would slowly but surely become modified in this same
manner. I have not attempted to conceal that, excepting when the males are
more numerous than the females, or when polygamy prevails, it is doubtful
how the more attractive males succeed in leaving a larger number of
offspring to inherit their superiority in ornaments or other charms than the
less attractive males; but I have shewn that this would probably follow from
the females,—especially the more vigorous females which would be the
first to breed, preferring not only the more attractive but at the same time
the more vigorous and victorious males.

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Although we have some positive evidence that birds appreciate bright and
beautiful objects, as with the Bower-birds of Australia, and although they
certainly appreciate the power of song, yet I fully admit that it is an
astonishing fact that the females of many birds and some mammals should
be endowed with sufficient taste for what has apparently been effected
through sexual selection; and this is even more astonishing in the case of
reptiles, fish, and insects. But we really know very little about the minds of
the lower animals. It cannot be supposed that male Birds of Paradise or
Peacocks, for instance, should take so much pains in erecting, spreading,
and vibrating their beautiful plumes before the females for no purpose. We
should remember the fact given on excellent authority in a former chapter,
namely that several peahens, when debarred from an admired male,
remained widows during a whole season rather than pair with another bird.
Nevertheless I know of no fact in natural history more wonderful than that
the female Argus pheasant should be able to appreciate the exquisite
shading of the ball-and-socket ornaments and the elegant patterns on the
wing-feathers of the male. He who thinks that the male was created as he
now exists must admit that the great plumes, which prevent the wings from
being used for flight, and which, as well as the primary feathers, are
displayed in a manner quite peculiar to this one species during the act of
courtship, and at no other time, were given to him as an ornament. If so, he
must likewise admit that the female was created and endowed with the
capacity of appreciating such ornaments. I differ only in the conviction that
the male Argus pheasant acquired his beauty gradually, through the females
having preferred during many generations the more highly ornamented
males; the æsthetic capacity of the females having been advanced through
exercise or habit in the same manner as our own taste is gradually
improved. In the male, through the fortunate chance of a few feathers not
having been modified, we can distinctly see how simple spots with a little
fulvous shading on one side might have been developed by small and
graduated steps into the wonderful ball-and-socket ornaments; and it is
probable that they were actually thus developed.
Everyone who admits the principle of evolution, and yet feels great
difficulty in admitting that female mammals, birds, reptiles, and fish, could
have acquired the high standard of taste which is implied by the beauty of
the males, and which generally coincides with our own standard, should

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reflect that in each member of the vertebrate series the nerve-cells of the
brain are the direct offshoots of those possessed by the common progenitor
of the whole group. It thus becomes intelligible that the brain and mental
faculties should be capable under similar conditions of nearly the same
course of development, and consequently of performing nearly the same
functions.
The reader who has taken the trouble to go through the several chapters
devoted to sexual selection, will be able to judge how far the conclusions at
which I have arrived are supported by sufficient evidence. If he accepts
these conclusions, he may, I think, safely extend them to mankind; but it
would be superfluous here to repeat what I have so lately said on the
manner in which sexual selection has apparently acted on both the male and
female side, causing the two sexes of man to differ in body and mind, and
the several races to differ from each other in various characters, as well as
from their ancient and lowly-organised progenitors.
He who admits the principle of sexual selection will be led to the
remarkable conclusion that the cerebral system not only regulates most of
the existing functions of the body, but has indirectly influenced the
progressive development of various bodily structures and of certain mental
qualities. Courage, pugnacity, perseverance, strength and size of body,
weapons of all kinds, musical organs, both vocal and instrumental, bright
colours, stripes and marks, and ornamental appendages, have all been
indirectly gained by the one sex or the other, through the influence of love
and jealousy, through the appreciation of the beautiful in sound, colour or
form, and through the exertion of a choice; and these powers of the mind
manifestly depend on the development of the cerebral system.

Man scans with scrupulous care the character and pedigree of his horses,
cattle, and dogs before he matches them; but when he comes to his own
marriage he rarely, or never, takes any such care. He is impelled by nearly
the same motives as are the lower animals when left to their own free
choice, though he is in so far superior to them that he highly values mental
charms and virtues. On the other hand he is strongly attracted by mere
wealth or rank. Yet he might by selection do something not only for the
bodily constitution and frame of his offspring, but for their intellectual and

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moral qualities. Both sexes ought to refrain from marriage if in any marked
degree inferior in body or mind; but such hopes are Utopian and will never
be even partially realised until the laws of inheritance are thoroughly
known. All do good service who aid towards this end. When the principles
of breeding and of inheritance are better understood, we shall not hear
ignorant members of our legislature rejecting with scorn a plan for
ascertaining by an easy method whether or not consanguineous marriages
are injurious to man.
The advancement of the welfare of mankind is a most intricate problem: all
ought to refrain from marriage who cannot avoid abject poverty for their
children; for poverty is not only a great evil, but tends to its own increase
by leading to recklessness in marriage. On the other hand, as Mr. Galton has
remarked, if the prudent avoid marriage, whilst the reckless marry, the
inferior members will tend to supplant the better members of society. Man,
like every other animal, has no doubt advanced to his present high condition
through a struggle for existence consequent on his rapid multiplication; and
if he is to advance still higher he must remain subject to a severe struggle.
Otherwise he would soon sink into indolence, and the more highly-gifted
men would not be more successful in the battle of life than the less gifted.
Hence our natural rate of increase, though leading to many and obvious
evils, must not be greatly diminished by any means. There should be open
competition for all men; and the most able should not be prevented by laws
or customs from succeeding best and rearing the largest number of
offspring. Important as the struggle for existence has been and even still is,
yet as far as the highest part of man’s nature is concerned there are other
agencies more important. For the moral qualities are advanced, either
directly or indirectly, much more through the effects of habit, the reasoning
powers, instruction, religion, &c., than through natural selection; though to
this latter agency the social instincts, which afforded the basis for the
development of the moral sense, may be safely attributed.

The main conclusion arrived at in this work, namely that man is descended
from some lowly-organised form, will, I regret to think, be highly
distasteful to many persons. But there can hardly be a doubt that we are
descended from barbarians. The astonishment which I felt on first seeing a
party of Fuegians on a wild and broken shore will never be forgotten by me,

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for the reflection at once rushed into my mind—such were our ancestors.
These men were absolutely naked and bedaubed with paint, their long hair
was tangled, their mouths frothed with excitement, and their expression was
wild, startled, and distrustful. They possessed hardly any arts, and like wild
animals lived on what they could catch; they had no government, and were
merciless to every one not of their own small tribe. He who has seen a
savage in his native land will not feel much shame, if forced to
acknowledge that the blood of some more humble creature flows in his
veins. For my own part I would as soon be descended from that heroic little
monkey, who braved his dreaded enemy in order to save the life of his
keeper; or from that old baboon, who, descending from the mountains,
carried away in triumph his young comrade from a crowd of astonished
dogs—as from a savage who delights to torture his enemies, offers up
bloody sacrifices, practises infanticide without remorse, treats his wives like
slaves, knows no decency, and is haunted by the grossest superstitions.
Man may be excused for feeling some pride at having risen, though not
through his own exertions, to the very summit of the organic scale; and the
fact of his having thus risen, instead of having been aboriginally placed
there, may give him hopes for a still higher destiny in the distant future. But
we are not here concerned with hopes or fears, only with the truth as far as
our reason allows us to discover it. I have given the evidence to the best of
my ability; and we must acknowledge, as it seems to me, that man with all
his noble qualities, with sympathy which feels for the most debased, with
benevolence which extends not only to other men but to the humblest living
creature, with his god-like intellect which has penetrated into the
movements and constitution of the solar system—with all these exalted
powers—Man still bears in his bodily frame the indelible stamp of his
lowly origin.

1 Yarrell’s ‘Hist. of British Fishes,’ vol. ii. 1836, p. 417, 425, 436. Dr. Günther
informs me that the spines in R. clavata are peculiar to the female.
2 See Mr. R. Warington’s interesting articles in ‘Annals and Mag. of Nat. Hist.’
Oct. 1852 and Nov. 1855.
3 Noel Humphreys, ‘River Gardens,’ 1857.

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4 Loudon’s ‘Mag. of Natural History,’ vol. iii. 1830, p. 331.
5 ‘The Field,’ June 29th, 1867. For Mr. Shaw’s statement, see 'Edinburgh
Review,’ 1843. Another experienced observer (Scrope’s 'Days of Salmon
Fishing,’ p. 60) remarks that the male would, if he could, keep, like the stag, all
other males away.
6 Yarrell, ‘History of British Fishes,’ vol. ii. 1836, p. 10.
7 ‘The Naturalist in Vancouver’s Island,’ vol. i. 1866, p. 54.
8 ‘Scandinavian Adventures,’ vol. i. 1854, p. 100, 104.
9 See Yarrell’s account of the Rays in his ‘Hist. of British Fishes,’ vol. ii. 1836,
p. 416, with an excellent figure, and p. 422, 432.
10 As quoted in ‘The Farmer,’ 1868, p. 369.
11 I have drawn up this description from Yarrell’s ‘British Fishes,’ vol. i. 1836, p.
261 and 266.
12 ‘Catalogue of Acanth. Fishes in the British Museum,’ by Dr. Günther, 1861,
p. 138-151.
13 ‘Game Birds of Sweden,’ &c., 1867, p. 466.
14 With respect to this and the following species I am indebted to Dr. Günther
for information: see also his paper on the Fishes of Central America, in
‘Transact. Zoolog. Soc.’ vol. vi. 1868, p. 485.
15 Dr. Günther makes this remark; ‘Catalogue of Fishes in the British Museum,’
vol. iii. 1861, p. 141.
16 See Dr. Günther on this genus, in ‘Proc. Zoolog. Soc.’ 1868, p. 232.
17 F. Buckland, in ‘Land and Water,’ July, 1868, p. 377, with a figure.
18 Dr. Günther, ‘Catalogue of Fishes,’ vol. iii. p. 221 and 240.
19 See also ‘A Journey in Brazil,’ by Prof. and Mrs. Agassiz, 1868, p. 220.
20 Yarrell, ‘British Fishes,’ vol. ii. 1836, p. 10, 12, 35.
21 W. Thompson, in ‘Annals and Mag. of Nat. History,’ vol. vi. 1841, p. 440.
22 ‘The American Agriculturist,’ 1868, p. 100.
23 ‘Annals and Mag. of Nat. Hist.’ Oct. 1852.
24 Loudon’s ‘Mag. of Nat. Hist.’ vol. v. 1832, p. 681.
25 Bory de Saint Vincent, in ‘Dict. Class. d’Hist. Nat.’ tom. ix. 1826, p. 151.
26 Owing to some remarks on this subject, made in my work ‘On the Variation
of Animals under Domestication,’ Mr. W. F. Mayers (‘Chinese Notes and
Queries,’ Aug. 1868, p. 123) has searched the ancient Chinese encyclopedias. He
finds that goldfish were first reared in confinement during the Sung Dynasty,
which commenced a.d. 960. In the year 1129 these fishes abounded. In another

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place it is said that since the year 1548 there has been “produced at Hang-chow a
variety called the fire-fish, from its intensely red colour. It is universally
admired, and there is not a household where it is not cultivated, in rivalry as to
its colour, and as a source of profit.”
27 ‘Westminster Review,’ July, 1867, p. 7.
28 “Indian Cyprinidæ,” by Mr. J. M’Clelland, ‘Asiatic Researches,’ vol. xix. part
ii. 1839, p. 230.
29 ‘Proc. Zoolog. Soc.’ 1865, p. 327, pl. xiv. and xv.
30 Yarrell, ‘British Fishes,’ vol. ii. p. 11.
31 According to the observations of M. Gerbe; see Günther’s ‘Record of Zoolog.
Literature,’ 1865, p. 194.
32 Cuvier, ‘Règne Animal,’ vol. ii. 1829, p. 242.
33 See Mr. Warington’s most interesting description of the habits of the
Gasterosteus leiurus, in ‘Annals and Mag. of Nat. Hist.’ November, 1855.
34 Prof. Wyman, in ‘Proc. Boston Soc. of Nat. Hist.’ Sept. 15, 1857. Also W.
Turner, in ‘Journal of Anatomy and Phys.’ Nov. 1, 1866, p. 78. Dr. Günther has
likewise described other cases.
35 Yarrell, ‘Hist. of British Fishes,’ vol. ii. 1836, p. 329, 338.
36 Dr. Günther, since publishing an account of this species in ‘The Fishes of
Zanzibar,’ by Col. Playfair, 1866, p. 137, has re-examined the specimens, and
has given me the above information.
37 The Rev. C. Kingsley, in ‘Nature,’ May, 1870, p. 40.
38 Bell, ‘History of British Reptiles,’ 2nd edit. 1849, p. 156-159.
39 Bell, ibid. p. 146, 151.
40 ‘Zoology of the Voyage of the “Beagle,”’ 1843. “Reptiles,” by Mr. Bell, p. 49.
41 ‘The Reptiles of India,’ by Dr. A. Günther, Ray Soc. 1864, p. 413.
42 Bell, ‘History of British Reptiles,’ 1849, p. 93.
43 J. Bishop, in ‘Todd’s Cyclop. of Anat. and Phys.’ vol. iv. p. 1503.
44 Bell, ibid. p. 112-114.
45 Mr. C. J. Maynard, ‘The American Naturalist,’ Dec. 1869, p. 555.
46 See my ‘Journal of Researches during the Voyage of the “Beagle,”’ 1845, p.
384.
47 ‘Travels through Carolina,’ &c., 1791, p. 128.
48 Owen, ‘Anatomy of Vertebrates,’ vol. i. 1866, p. 615.
49 Sir Andrew Smith, ‘Zoolog. of S. Africa: Reptilia,’ 1849, pl. x.

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50 Dr. A. Günther, ‘Reptiles of British India,’ Ray Soc. 1864, p. 304, 308.
51 Owen, ‘Anatomy of Vertebrates,’ vol. i. 1866, p. 615.
52 The celebrated botanist Schleiden incidently remarks (‘Ueber den
Darwinismus: Unsere Zeit,’ 1869, s. 269), that Rattle-snakes use their rattles as a
sexual call, by which the two sexes find each other. I do not know whether this
suggestion rests on any direct observations. These snakes pair in the Zoological
Gardens, but the keepers have never observed that they use their rattles at this
season more than at any other.
53 “Rambles in Ceylon,” ‘Annals and Mag. of Nat. Hist.’ 2nd series, vol. ix.
1852, p. 333.
54 ‘Westminster Review,’ July 1st, 1867, p. 32.
55 Mr. N. L. Austen kept these animals alive for a considerable time, see ‘Land
and Water,’ July, 1867, p. 9.
56 All these statements and quotations, in regard to Cophotis, Sitana and Draco,
as well as the following facts in regard to Ceratophora, are taken from Dr.
Günther’s magnificent work on the ‘Reptiles of British India,’ Ray Soc. 1864, p.
122, 130, 135.
57 Bell, ‘History of British Reptiles,’ 2nd edit. 1849, p. 40.
58 For Proctotretus see ‘Zoology of the Voyage of the “Beagle:” Reptiles,’ by
Mr. Bell, p. 8. For the Lizards of S. Africa, see ‘Zoology of S. Africa: Reptiles,’
by Sir Andrew Smith, pl. 25 and 39. For the Indian Calotes, see ‘Reptiles of
British India,’ by Dr. Günther, p. 143.
59 ‘Ibis,’ vol. iii. (new series) 1867, p. 414.
60 Gould, ‘Handbook to the Birds of Australia,’ 1865, vol. ii. p. 383.
61 Quoted by Mr. Gould, ‘Introduction to the Trochilidæ,’ 1861, p. 29.
62 Gould, ibid. p. 52.
63 W. Thompson, ‘Nat. Hist. of Ireland: Birds,’ vol. ii. 1850, p. 327.
64 Jerdon, ‘Birds of India,’ 1863, vol. ii. p. 96.
65 Macgillivray, ‘Hist. Brit. Birds,’ vol. iv. 1852, p. 177-181.
66 Sir R. Schomburgk, in ‘Journal of R. Geograph. Soc.’ vol. xiii. 1843, p. 31.
67 ‘Ornithological Biography,’ vol. i. p. 191. For pelicans and snipes, see vol. iii.
p. 381, 477.
68 Gould, ‘Handbook of Birds of Australia,’ vol. i. p. 395; vol. ii. p. 383.
69 Mr. Hewitt in the ‘Poultry Book by Tegetmeier,’ 1866, p. 137.
70 Layard, ‘Annals and Mag. of Nat. Hist.’ vol. xiv. 1854, p. 63.
71 Jerdon, ‘Birds of India,’ vol. iii. p. 574.

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72 Brehm, ‘Illust. Thierleben,’ 1867, B. iv. s. 351. Some of the foregoing
statements are taken from L. Lloyd, ‘The Game Birds of Sweden,’ &c., 1867, p.
79.
73 Jerdon, ‘Birds of India:’ on Ithaginis, vol. iii. p. 523; on Galloperdix, p. 541.
74 For the Egyptian goose, see Macgillivray, ‘British Birds,’ vol. iv. p. 639. For
Plectropterus, ‘Livingstone’s Travels,’ p. 254. For Palamedea, Brehm’s
‘Thierleben,’ B. iv. s. 740. See also on this bird Azara, 'Voyages dans l’Amérique
mérid.’ tom. iv. 1809, p. 179, 253.
75 See, on our peewit, Mr. R. Carr in ‘Land and Water,’ Aug. 8th, 1868, p. 46. In
regard to Lobivanellus, see Jerdon’s ‘Birds of India,’ vol. iii. p. 647, and Gould’s
‘Handbook of Birds of Australia,’ vol. ii. p. 220. For the Hoplopterus, see Mr.
Allen in the ‘Ibis,’ vol. v. 1863, p. 156.
76 Audubon, ‘Ornith. Biography,’ vol. ii. p. 492; vol. i. p. 4-13.
77 Mr. Blyth, ‘Land and Water,’ 1867, p. 212.
78 Richardson, on Tetrao umbellus, ‘Fauna Bor. Amer.: Birds,’ 1831, p. 343. L.
Lloyd, ‘Game Birds of Sweden,’ 1867, p. 22, 79, on the capercailzie and black-
cock. Brehm, however, asserts (‘Thierleben,’ &c., B. iv. s. 352) that in Germany
the grey-hens do not generally attend the Balzen of the black-cocks, but this is an
exception to the common rule; possibly the hens may lie hidden in the
surrounding bushes, as is known to be the case with the grey-hens in
Scandinavia, and with other species in N. America.
79 ‘Ornithological Biography,’ vol. ii. p. 275.
80 Brehm, ‘Thierleben,’ &c., B. iv. 1867, p. 990. Audubon, ‘Ornith. Biography,’
vol. ii. p. 492.
81 ‘Land and Water,’ July 25th, 1868, p. 14.
82 Audubon’s ‘Ornitholog. Biography;’ on Tetrao cupido, vol. ii. p. 492; on the
Sturnus, vol. ii. p. 219.
83 ‘Ornithological Biograph.’ vol. v. p. 601.
84 The Hon. Daines Barrington, ‘Philosoph. Transact.’ 1773, p. 252.
85 ‘Ornithological Dictionary,’ 1833, p. 475.
86 ‘Naturgeschichte der Stubenvögel,’ 1840, s. 4. Mr. Harrison Weir likewise
writes to me:—“I am informed that the best singing males generally get a mate
first when they are bred in the same room.”
87 ‘Philosophical Transactions,’ 1773, p. 263. White’s ‘Natural History of
Selborne,’ vol. i. 1825, p. 246.
88 ‘Naturges. der Stubenvögel,’ 1840, s. 252.
89 Mr. Bold, ‘Zoologist,’ 1843-44, p. 659.

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90 D. Barrington, ‘Phil. Transact.’ 1773, p. 262. Bechstein, ‘Stubenvögel,’ 1840,
s. 4.
91 This is likewise the case with the water-ouzel, see Mr. Hepburn in the
‘Zoologist,’ 1845-1846, p. 1068.
92 L. Lloyd, ‘Game Birds of Sweden,’ 1867, p. 25.
93 Barrington, ibid. p. 264. Bechstein, ibid. s. 5.
94 Dureau de la Malle gives a curious instance (‘Annales des Sc. Nat.’ 3rd series,
Zoolog. tom. x. p. 118) of some wild blackbirds in his garden in Paris which
naturally learnt from a caged bird a republican air.
95 Bishop, in ‘Todd’s Cyclop. of Anat. and Phys.’ vol. iv. p. 1496.
96 As stated by Barrington in ‘Philosoph. Transact.’ 1773, p. 262.
97 Gould, ‘Handbook to the Birds of Australia,’ vol. i. 1865, p. 308-310. See also
Mr. T. W. Wood in the ‘Student,’ April, 1870, p. 125.
98 See remarks to this effect in Gould’s ‘Introduction to the Trochilidæ,’ 1861, p.
22.
99 ‘The Sportsman and Naturalist in Canada,’ by Major W. Ross King, 1866, p.
144-146. Mr. T. W. Wood gives in the ‘Student’ (April, 1870, p. 116) an excellent
account of the attitude and habits of this bird during its courtship. He states that
the ear-tufts or neck-plumes are erected, so that they meet over the crown of the
head.
100 Richardson, ‘Fauna Bor. Americana: Birds,’ 1831, p. 359. Audubon, ibid.
vol. iv. p. 507.
101 The following papers have been lately written on this subject:—Prof. A.
Newton, in the ‘Ibis,’ 1862, p. 107; Dr. Cullen, ibid. 1865, p. 145; Mr. Flower, in
‘Proc. Zool. Soc.’ 1865, p. 747; and Dr. Murie, in ‘Proc. Zool. Soc.’ 1868, p. 471.
In this latter paper an excellent figure is given of the male Australian Bustard in
full display with the sack distended.
102 Bates, ‘The Naturalist on the Amazons,’ 1863, vol. ii. p. 284; Wallace, in
‘Proc. Zool. Soc.’ 1850, p. 206. A new species, with a still larger neck-appendage
(C. penduliger), has lately been discovered, see 'Ibis,’ vol. i. p. 457.
103 Bishop, in Todd’s ‘Cyclop. of Anat. and Phys.’ vol. iv. p. 1499.
104 The spoonbill (Platalea) has its trachea convoluted into a figure of eight,
and yet this bird (Jerdon, ‘Birds of India,’ vol. iii. p. 763) is mute; but Mr. Blyth
informs me that the convolutions are not constantly present, so that perhaps they
are now tending towards abortion.
105 ‘Elements of Comp. Anat.’ by R. Wagner, Eng. translat. 1845, p. 111. With
respect to the swan, as given above, Yarrell’s ‘Hist. of British Birds,’ 2nd edit.
1845, vol. iii. p. 193.
106 C. L. Bonaparte, quoted in the ‘Naturalist Library: Birds,’ vol. xiv. p. 126.

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107 L. Lloyd, ‘The Game Birds of Sweden,’ &c., 1867, p. 22, 81.
108 Jenner, ‘Philosoph. Transactions,’ 1824, p. 20.
109 For the foregoing several facts see, on Birds of Paradise, Brehm,
'Thierleben,’ Band iii. s. 325. On Grouse, Richardson, ‘Fauna Bor. Americ.:
Birds,’ p. 343 and 359; Major W. Ross King, ‘The Sportsman in Canada,’ 1866,
p. 156; Audubon, ‘American Ornitholog. Biograph.’ vol. i. p. 216. On the Kalij
pheasant, Jerdon, ‘Birds of India,’ vol. iii. p. 533. On the Weavers, ‘Livingstone’s
Expedition to the Zambesi,’ 1865, p. 425. On Woodpeckers, Macgillivray, ‘Hist.
of British Birds,’ vol. iii. 1840, p. 84, 88, 89, and 95. On the Hoopoe, Mr.
Swinhoe, in ‘Proc. Zoolog. Soc.’ June 23, 1863. On the Night-Jar, Audubon,
ibid. vol. ii. p. 255. The English Night-Jar likewise makes in the spring a curious
noise during its rapid flight.
110 See M. Meves’ interesting paper in ‘Proc. Zool. Soc.’ 1858, p. 199. For the
habits of the snipe, Macgillivray, ‘Hist. British Birds,’ vol. iv. p. 371. For the
American snipe, Capt. Blakiston, ‘Ibis,’ vol. v. 1863, p. 131.
111 Mr. Salvin, in ‘Proc. Zool. Soc.’ 1867, p. 160. I am much indebted to this
distinguished ornithologist for sketches of the feathers of the Chamæpetes, and
for other information.
112 Jerdon, ‘Birds of India,’ vol. iii. p. 618, 621.
113 Gould, ‘Introduction to the Trochilidæ,’ 1861, p. 49. Salvin, 'Proc. Zoolog.
Soc.’ 1867, p. 160.
114 Sclater, in ‘Proc. Zool. Soc.’ 1860, p. 90, and in ‘Ibis,’ vol. iv. 1862, p. 175.
Also Salvin, in ‘Ibis,’ 1860, p. 37.
115 ‘The Nile Tributaries of Abyssinia,’ 1867, p. 203.
116 For Tetrao phasianellus, see Richardson, ‘Fauna Bor. America,’ p. 361, and
for further particulars Capt. Blakiston, ‘Ibis,’ 1863, p. 125. For the Cathartes and
Ardea, Audubon, ‘Ornith. Biography,’ vol. ii. p. 51, and vol. iii. p. 89. On the
White-throat, Macgillivray, ‘Hist. British Birds,’ vol. ii. p. 354. On the Indian
Bustard, Jerdon, ‘Birds of India,’ vol. iii. p. 618.
117 Gould, ‘Handbook to the Birds of Australia,’ vol. i. p. 444, 449, 455. The
bower of the Satin Bower-bird may always be seen in the Zoological Society’s
Gardens, Regent’s Park.
118 See remarks to this effect, on the “Feeling of Beauty among Animals,” by
Mr. J. Shaw, in the ‘Athenæum,’ Nov. 24th, 1866, p. 681.
119 Mr. Monteiro, ‘Ibis,’ vol. iv. 1862, p. 339.
120 ‘Land and Water,’ 1868, p. 217.
121 Jardine’s ‘Naturalist Library: Birds,’ vol. xiv. p. 166.
122 Sclater, in the ‘Ibis,’ vol. vi. 1864, p. 114. Livingstone, ‘Expedition to the
Zambesi,’ 1865, p. 66.
123 Jerdon, ‘Birds of India,’ vol. iii. p. 620.

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124 Wallace, in ‘Annals and Mag. of Nat. Hist.’ vol. xx. 1857, p. 416; and in his
‘Malay Archipelago,’ vol. ii. 1869, p. 390.
125 See my work on ‘The Variation of Animals and Plants under Domestication,’
vol. i. p. 289, 293.
126 Quoted from M. de Lafresnaye, in ‘Annals and Mag. of Nat. Hist.’ vol. xiii.
1854, p. 157: see also Mr. Wallace’s much fuller account in vol. xx. 1857, p. 412,
and in his Malay Archipelago.
127 Wallace, ‘The Malay Archipelago,’ vol. ii. 1869, p. 405.
128 Mr. Sclater, ‘Intellectual Observer,’ Jan. 1867. ‘Waterton’s Wanderings,’ p.
118. See also Mr. Salvin’s interesting paper, with a plate, in the ‘Ibis,’ 1865, p.
90.
129 ‘Land and Water,’ 1867, p. 394.
130 Mr. D. G. Elliot, in ‘Proc. Zool. Soc.’ 1869, p. 589.
131 ‘Nitzsch’s Pterylography,’ edited by P. L. Sclater. Ray Soc. 1867, p. 14.
132 The brown mottled summer plumage of the ptarmigan is of as much
importance to it, as a protection, as the white winter plumage; for in Scandinavia,
during the spring, when the snow has disappeared, this bird is known to suffer
greatly from birds of prey, before it has acquired its summer dress: see Wilhelm
von Wright, in Lloyd, ‘Game Birds of Sweden,’ 1867, p. 125.
133 In regard to the previous statements on moulting, see, on snipes, &c.,
Macgillivray, ‘Hist. Brit. Birds,’ vol. iv. p. 371; on Glareolæ, curlews, and
bustards, Jerdon, ‘Birds of India,’ vol. iii. p. 615, 630, 683; on Totanus, ibid, p.
700; on the plumes of herons, ibid, p. 738, and Macgillivray, vol. iv. p. 435 and
444, and Mr. Stafford Allen, in the ‘Ibis,’ vol. v. 1863, p. 33.
134 On the moulting of the ptarmigan, see Gould’s ‘Birds of Great Britain.’ On
the honey-suckers, Jerdon, ‘Birds of India,’ vol. i. p. 359, 365, 369. On the
moulting of Anthus, see Blyth, in ‘Ibis,’ 1867, p. 32.
135 For the foregoing statements in regard to partial moults, and on old males
retaining their nuptial plumage, see Jerdon, on bustards and plovers, in ‘Birds of
India,’ vol. iii. p. 617, 637, 709, 711. Also Blyth in ‘Land and Water,’ 1867, p.
84. On the Vidua, ‘Ibis,’ vol. iii. 1861, p. 133. On the Drongo shrikes, Jerdon,
ibid. vol. i. p. 435. On the vernal moult of the Herodias bubulcus, Mr. S. S.
Allen, in ‘Ibis,’ 1863, p. 33. On Gallus bankiva, Blyth, in ‘Annals and Mag. of
Nat. Hist.’ vol. i. 1848, p. 455; see, also, on this subject, my ‘Variation of
Animals under Domestication,’ vol. i. p. 236.
136 See Macgillivray, ‘Hist. British Birds’ (vol. v. p. 34, 70, and 223), on the
moulting of the Anatidæ, with quotations from Waterton and Montagu. Also
Yarrell, ‘Hist. of British Birds,’ vol. iii. p. 243.
137 On the pelican, see Sclater, in ‘Proc. Zool. Soc.’ 1868, p. 265. On the
American finches, see Audubon, ‘Ornith. Biography,’ vol. i. p. 174, 221, and
Jerdon, ‘Birds of India,’ vol. ii. p. 383. On the Fringilla cannabina of Madeira,
Mr. E. Vernon Harcourt, ‘Ibis,’ vol. v., 1863, p. 230.

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138 See also ‘Ornamental Poultry,’ by Rev. E. S. Dixon, 1848, p. 8.
139 ‘Birds of India,’ introduct. vol. i. p. xxiv.; on the peacock, vol. iii. p. 507.
See Gould’s ‘Introduction to the Trochilidæ,’ 1861, p. 15 and 111.
140 ‘Journal of R. Geograph. Soc.’ vol. x. 1840, p. 236.
141 ‘Annals and Mag. of Nat. Hist.’ vol. xiii. 1854, p. 157; also Wallace, ibid.
vol. xx. 1857, p. 412, and ‘The Malay Archipelago,’ vol. ii. 1869, p. 252. Also
Dr. Bennett, as quoted by Brehm, ‘Thierleben,’ B. iii. s. 326.
142 Mr. T. W. Wood has given (‘The Student,’ April, 1870, p. 115) a full account
of this manner of display, which he calls the lateral or one-sided, by the gold
pheasant and by the Japanese pheasant, Ph. versicolor.
143 ‘The Reign of Law,’ 1867, p. 263.
144 For the description of these birds, see Gould’s ‘Handbook to the Birds of
Australia,’ vol. i. 1865, p. 417.
145 ‘Birds of India,’ vol. ii. p. 96.
146 On the Cosmetornis, see Livingstone’s ‘Expedition to the Zambesi,’ 1865, p.
66. On the Argus pheasant, Jardine’s ‘Nat. Hist. Lib.: Birds,’ vol. xiv. p. 167. On
Birds of Paradise, Lesson, quoted by Brehm, 'Thierleben,’ B. iii. s. 325. On the
widow-bird, Barrow’s ‘Travels in Africa,’ vol. i. p. 243, and ‘Ibis,’ vol. iii. 1861,
p. 133. Mr. Gould, on the shyness of male birds, ‘Handbook to Birds of
Australia,’ vol. i. 1865, p. 210, 457.
147 Tegetmeier, ‘The Poultry Book,’ 1866, p. 139.
148 Nordmann describes (‘Bull. Soc. Imp. des Nat. Moscow,’ 1861, tom. xxxiv.
p. 264) the balzen of Tetrao urogalloides in Amur Land. He estimated the
number of assembled males at above a hundred, the females, which lie hid in the
surrounding bushes, not being counted. The noises uttered differ from those of
the T. urogallus or the capercailzie.
149 With respect to the assemblages of the above named grouse see Brehm,
‘Thierleben,’ B. iv. s. 350; also L. Lloyd, ‘Game Birds of Sweden,’ 1867, p. 19,
78. Richardson, ‘Fauna Bor. Americana,’ Birds, p. 362. References in regard to
the assemblages of other birds have previously been given. On Paradisea see
Wallace, in ‘Annals and Mag. of Nat. Hist.’ vol. xx. 1857, p. 412. On the snipe,
Lloyd, ibid. p. 221.

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150 Quoted by Mr. T. W. Wood in the ‘Student,’ April, 1870, p. 125.
151 Gould, ‘Handbook of Birds of Australia,’ vol. i. p. 300, 308, 448, 451. On
the ptarmigan, above alluded to, see Lloyd, ibid. p. 129.
152 On magpies, Jenner, in ‘Phil. Transact.’ 1824, p. 21. Macgillivray, 'Hist.
British Birds,’ vol. i. p. 570. Thompson, in ‘Annals and Mag. of Nat. Hist.’ vol.
viii. 1842, p. 494.
153 On the peregrine falcon see Thompson, ‘Nat. Hist. of Ireland: Birds,’ vol. i.
1849, p. 39. On owls, sparrows, and partridges, see White, 'Nat. Hist. of
Selborne,’ edit. of 1825, vol. i. p. 139. On the Phœnicura, see Loudon’s ‘Mag. of
Nat. Hist.’ vol. vii. 1834, p. 245. Brehm, 'Thierleben,’ B. iv. s. 991) also alludes
to cases of birds thrice mated during same day.
154 See White (‘Nat. Hist. of Selborne,’ 1825, vol. i. p. 140) on the existence,
early in the season, of small coveys of male partridges, of which fact I have
heard other instances. See Jenner, on the retarded state of the generative organs
in certain birds, in ‘Phil. Transact.’ 1824. In regard to birds living in triplets, I
owe to Mr. Jenner Weir the cases of the starling and parrots, and to Mr. Fox, of
partridges; on carrion-crows, see the ‘Field,’ 1868, p. 415. On various male birds
singing after the proper period, see Rev. L. Jenyns, ‘Observations in Natural
History,’ 1846, p. 87.
155 The following case has been given (‘The Times,’ Aug. 6th, 1868) by the Rev.
F. O. Morris, on the authority of the Hon. and Rev. O. W. Forester. “The
gamekeeper here found a hawk’s nest this year, with five young ones in it. He
took four and killed them, but left one with its wings clipped as a decoy to
destroy the old ones by. They were both shot next day, in the act of feeding the
young one, and the keeper thought it was done with. The next day he came again
and found two other charitable hawks, who had come with an adopted feeling to
succour the orphan. These two he killed, and then left the nest. On returning
afterwards he found two more charitable individuals on the same errand of
mercy. One of these he killed; the other he also shot, but could not find. No more
came on the like fruitless errand.”
156 For instance, Mr. Yarrell states (‘Hist. British Birds,’ vol. iii. 1845, p. 585)
that a gull was not able to swallow a small bird which had been given to it. The
gull “paused for a moment, and then, as if suddenly recollecting himself, ran off
at full speed to a pan of water, shook the bird about in it until well soaked, and
immediately gulped it down. Since that time he invariably has had recourse to
the same expedient in similar cases.”
157 ‘A Tour in Sutherlandshire,’ vol. i. 1849, p. 185.
158 ‘Acclimatization of Parrots,’ by C. Buxton, M.P. ‘Annals and Mag. of Nat.
Hist.’ Nov. 1868, p. 381.
159 ‘The Zoologist,’ 1847-1848, p. 1602.
160 Hewitt on wild ducks, ‘Journal of Horticulture,’ Jan. 13, 1863, p. 39.
Audubon on the wild turkey, ‘Ornith. Biography,’ vol. i. p. 14. On the mocking
thrush, ibid. vol. i. p. 110.

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161 The ‘Ibis,’ vol. ii. 1860, p. 344.
162 On the ornamented nests of humming-birds, Gould, ‘Introduction to the
Trochilidæ,’ 1861, p. 19. On the bower-birds, Gould, 'Handbook to the Birds of
Australia,’ 1865, vol. i. p. 444-461. Mr. Ramsay in the ‘Ibis,’ 1867, p. 456.
163 ‘Hist. of British Birds,’ vol. ii. p. 92.
164 ‘Zoologist,’ 1853-1854, p. 3946.
165 Waterton, ‘Essays on Nat. Hist.’ 2nd series, p. 42, 117. For the following
statements, see on the wigeon, Loudon’s ‘Mag. of Nat. Hist.’ vol. ix. p. 616; L.
Lloyd, ‘Scandinavian Adventures,’ vol. i. 1854, p. 452; Dixon, ‘Ornamental and
Domestic Poultry,’ p. 137; Hewitt, in ‘Journal of Horticulture,’ Jan. 13, 1863, p.
40; Bechstein, ‘Stubenvögel,’ 1840, s. 230.
166 Audubon, ‘Ornitholog. Biography,’ vol. i. p. 191, 349; vol. ii. p. 42, 275; vol.
iii. p. 2.
167 ‘Rare and Prize Poultry,’ 1854, p. 27.
168 ‘The Variation of Animals and Plants under Domestication,’ vol. ii. p. 103.
169 Boitard and Corbié, ‘Les Pigeons,’ 1824, p. 12. Prosper Lucas (‘Traité de
l’Héréd. Nat.’ tom. ii. 1850, p. 296) has himself observed nearly similar facts
with pigeons.
170 ‘Die Taubenzucht,’ 1824, s. 86.
171 ‘Ornithological Biography,’ vol. i. p. 13.
172 ‘Proc. Zool. Soc.’ 1835, p. 54. The japanned peacock is considered by Mr.
Sclater as a distinct species, and has been named Pavo nigripennis.
173 Rudolphi, ‘Beyträge zur Anthropologie,’ 1812, s. 184.
174 ‘Die Darwin’sche Theorie, und ihre Stellung zu Moral und Religion,’ 1869,
s. 59.
175 In regard to peafowl, see Sir R. Heron, ‘Proc. Zoolog. Soc.’ 1835, p. 54, and
the Rev. E. S. Dixon, ‘Ornamental Poultry,’ 1848, p. 8. For the turkey, Audubon,
ibid. p. 4. For the capercailzie, Lloyd, 'Game Birds of Sweden,’ 1867, p. 23.
176 Mr. Hewitt, quoted in ‘Tegetmeier’s Poultry Book,’ 1866, p. 165.
177 Quoted in Lloyd’s ‘Game Birds of Sweden,’ p. 345.
178 According to Dr. Blasius (‘Ibis,’ vol. ii. 1860, p. 297), there are 425
indubitable species of birds which breed in Europe, besides 60 forms, which are
frequently regarded as distinct species. Of the latter, Blasius thinks that only ten
are really doubtful, and that the other fifty ought to be united with their nearest
allies; but this shews that there must be a considerable amount of variation with
some of our European birds. It is also an unsettled point with naturalists, whether
several North American birds ought to be ranked as specifically distinct from the
corresponding European species.

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179 ‘Origin of Species,’ fifth edit. 1869, p. 104. I had always perceived, that rare
and strongly-marked deviations of structure, deserving to be called
monstrosities, could seldom be preserved through natural selection, and that the
preservation of even highly-beneficial variations would depend to a certain
extent on chance. I had also fully appreciated the importance of mere individual
differences, and this led me to insist so strongly on the importance of that
unconscious form of selection by man, which follows from the preservation of
the most valued individuals of each breed, without any intention on his part to
modify the characters of the breed. But until I read an able article in the ‘North
British Review’ (March, 1867, p. 289, et seq.), which has been of more use to me
than any other Review, I did not see how great the chances were against the
preservation of variations, whether slight or strongly pronounced, occurring only
in single individuals.
180 ‘Introduct. to the Trochilidæ,’ p. 102.
181 Gould, ‘Handbook of Birds of Australia,’ vol. ii. p. 32 and 68.
182 Audubon, ‘Ornitholog. Biography,’ 1838, vol. iv. p. 389.
183 Jerdon, ‘Birds of India,’ vol. i. p. 108; and Mr. Blyth, in ‘Land and Water,’
1868, p. 381.
184 Graba, ‘Tagebuch, Reise nach Färo,’ 1830, s. 51-54. Macgillivray, 'Hist.
British Birds,’ vol. iii. p. 745. ‘Ibis,’ vol. v. 1863, p. 469.
185 Graba, ibid. s. 54. Macgillivray, ibid. vol. v. p. 327.
186 ‘Variation of Animals and Plants under Domestication,’ vol. ii. p. 92.
187 On these points see also ‘Variation of Animals and Plants under
Domestication,’ vol. i. p. 253; vol. ii. p. 73, 75.
188 See, for instance, on the irides of a Podica and Gallicrex in ‘Ibis,’ vol. ii.
1860, p. 206; and vol. v. 1863, p. 426.
189 See also Jerdon, ‘Birds of India,’ vol. i. p. 243-245.
190 ‘Zoology of the Voyage of H.M.S. Beagle,’ 1841, p. 6.
191 Bechstein, ‘Naturgeschichte Deutschlands,’ B. iv. 1795, s. 31, on a sub-
variety of the Monck pigeon.
192 This woodcut has been engraved from a beautiful drawing, most kindly
made for me by Mr. Trimen; see also his description of the wonderful amount of
variation in the coloration and shape of the wings of this butterfly, in his,
‘Rhopalocera Africæ Australis,’ p. 186. See also an interesting paper by the Rev.
H. H. Higgins, on the origin of the ocelli in the Lepidoptera in the ‘Quarterly
Journal of Science,’ July, 1868, p. 325.
193 Jerdon, ‘Birds of India,’ vol. iii. p. 517.
194 ‘Variation of Animals and Plants under Domestication,’ vol. i. p. 254.
195 When the Argus pheasant displays his wing-feathers like a great fan, those
nearest to the body stand more upright than the outer ones, so that the shading of

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the ball-and-socket ocelli ought to be slightly different on the different feathers,
in order to bring out their full effect, relatively to the incidence of the light. Mr.
T. W. Wood, who has the experienced eye of an artist, asserts (‘Field,’
Newspaper, May 28, 1870, p. 457) that this is the case; but after carefully
examining two mounted specimens (the proper feathers from one having been
given to me by Mr. Gould for more accurate comparison) I cannot perceive that
this acme of perfection in the shading has been attained; nor can others to whom
I have shewn these feathers recognise the fact.
196 ‘The Reign of Law,’ 1867, p. 247.
197 ‘Introduction to the Trochilidæ,’ 1861, p. 110.
198 Fourth edition, 1866, p. 241.
199 ‘Westminster Review,’ July, 1867. ‘Journal of Travel,’ vol. i. 1868, p. 73.
200 Temminck says that the tail of the female Phasianus Sœmmerringii is only
six inches long, ‘Planches coloriées,’ vol. v. 1838, p. 487 and 488: the
measurements above given were made for me by Mr. Sclater. For the common
pheasant, see Macgillivray, ‘Hist. Brit. Birds,’ vol. i. p. 118-121.
201 Dr. Chapuis, ‘Le Pigeon Voyageur Belge,’ 1865, p. 87.
202 Bechstein, ‘Naturgesch. Deutschlands,’ 1793, B. iii. s. 339.
203 Daines Barrington, however, thought it probable (‘Phil. Transact.’ 1773, p.
164) that few female birds sing, because the talent would have been dangerous to
them during incubation. He adds, that a similar view may possibly account for
the inferiority of the female to the male in plumage.
204 Mr. Ramsay, in ‘Proc. Zoolog. Soc.’ 1868, p. 50.
205 ‘Journal of Travel,’ edited by A. Murray, vol. i. 1868, p. 78.
206 ‘Journal of Travel,’ edited by A. Murray, vol. i. 1868, p. 281.
207 Audubon, ‘Ornithological Biography,’ vol. i. p. 233.
208 Jerdon, ‘Birds of India,’ vol. ii. p. 108. Gould’s ‘Handbook of the Birds of
Australia,’ vol. i. p. 463.
209 For instance, the female Eupetomena macroura has the head and tail dark
blue with reddish loins; the female Lampornis porphyrurus is blackish-green on
the upper surface, with the lores and sides of the throat crimson; the female
Eulampis jugularis has the top of the head and back green, but the loins and the
tail are crimson. Many other instances of highly conspicuous females could be
given. See Mr. Gould’s magnificent work on this family.
210 Mr. Salvin noticed in Guatemala (‘Ibis,’ 1864, p. 375) that humming-birds
were much more unwilling to leave their nests during very hot weather, when the
sun was shining brightly, than during cool, cloudy, or rainy weather.
211 I may specify, as instances of obscurely-coloured birds building concealed
nests, the species belonging to eight Australian genera, described in Gould’s

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‘Handbook of the Birds of Australia,’ vol. i. p. 340, 362, 365, 383, 387, 389, 391,
414.
212 Jerdon, ‘Birds of India,’ vol. i. p. 244.
213 On the nidification and colours of these latter species, see Gould’s
'Handbook,’ &c., vol. i. p. 504, 527.
214 I have consulted, on this subject, Macgillivray’s ‘British Birds,’ and though
doubts may be entertained in some cases in regard to the degree of concealment
of the nest, and of the degree of conspicuousness of the female, yet the following
birds, which all lay their eggs in holes or in domed nests, can hardly be
considered, according to the above standard, as conspicuous: Passer, 2 species;
Sturnus, of which the female is considerably less brilliant than the male; Cinclus;
Motacilla boarula (?); Erithacus (?); Fruticola, 2 sp.; Saxicola; Ruticilla, 2 sp.;
Sylvia, 3 sp.; Parus, 3 sp.; Mecistura; Anorthura; Certhia; Sitta; Yunx;
Muscicapa, 2 sp.; Hirundo, 3 sp.; and Cypselus. The females of the following 12
birds may be considered as conspicuous according to the same standard, viz.,
Pastor, Motacilla alba, Parus major and P. cæruleus, Upupa, Picus, 4 sp.,
Coracias, Alcedo, and Merops.
215 ‘Journal of Travel,’ edited by A. Murray, vol. i. p. 78.
216 See many statements in the ‘Ornithological Biography.’ See, also, some
curious observations on the nests of Italian birds by Eugenio Bettoni, in the ‘Atti
della Società Italiana,’ vol. xi. 1869, p. 487.
217 See his ‘Monograph of the Trogonidæ,’ first edition.
218 Namely Cyanalcyon. Gould’s ‘Handbook of the Birds of Australia,’ vol. i. p.
133; see, also, p. 130, 136.
219 Every gradation of difference between the sexes may be followed in the
parrots of Australia. See Gould’s ‘Handbook,’ &c., vol. ii. p. 14-102.
220 Macgillivray’s ‘British Birds,’ vol. ii. p. 433. Jerdon, ‘Birds of India,’ vol. ii.
p. 282.
221 All the following facts are taken from M. Malherbe’s magnificent
'Monographie des Picidées,’ 1861.
222 Audubon’s ‘Ornithological Biography,’ vol. ii. p. 75; see also the 'Ibis,’ vol.
i. p. 268.
223 Gould’s ‘Handbook of the Birds of Australia,’ vol. ii. p. 109-149.
224 See remarks to this effect in my work on ‘Variation under Domestication,’
vol. ii. chap, xii.
225 The ‘Ibis,’ vol. vi. 1864, p. 122.
226 On Ardetta, Translation of Cuvier’s ‘Règne Animal,’ by Mr. Blyth, footnote,
p. 159. On the Peregrine Falcon, Mr. Blyth, in Charlesworth’s 'Mag. of Nat.
Hist.’ vol. i. 1837, p. 304. On Dicrurus, ‘Ibis,’ 1863, p. 44. On the Platalea,
‘Ibis,’ vol. vi. 1864, p. 366. On the Bombycilla, Audubon’s ‘Ornitholog.

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Biography,’ vol. i. p. 229. On the Palæornis, see, also, Jerdon, ‘Birds of India,’
vol. i. p. 263. On the wild turkey, Audubon, ibid. vol. i. p. 15: I hear from Judge
Caton that in Illinois the female very rarely acquires a tuft.
227 Mr. Blyth has recorded (Translation of Cuvier’s ‘Règne Animal,’ p. 158)
various instances with Lanius, Ruticilla, Linaria, and Anas. Audubon has also
recorded a similar case (‘Ornith. Biog.’ vol. v. p. 519) with Tyranga æstiva.
228 See Gould’s ‘Birds of Great Britain.’
229 In regard to thrushes, shrikes, and woodpeckers, see Mr. Blyth, in
Charlesworth’s ‘Mag. of Nat. Hist.’ vol. i. 1837, p. 304; also footnote to his
translation of Cuvier’s ‘Règne Animal,’ p. 159. I give the case of Loxia from Mr.
Blyth’s information. On thrushes, see also Audubon, 'Ornith. Biography,’ vol. ii.
p. 195. On Chrysococcyx and Chalcophaps, Blyth, as quoted in Jerdon’s ‘Birds
of India,’ vol. iii. p. 485. On Sarkidiornis, Blyth, in ‘Ibis,’ 1867, p. 175.
230 See, for instance, Mr. Gould’s account (‘Handbook of the Birds of
Australia,’ vol. i. p. 133) of Cyanalcyon (one of the Kingfishers) in which,
however, the young male, though resembling the adult female, is less brilliantly
coloured. In some species of Dacelo the males have blue tails, and the females
brown ones; and Mr. R. B. Sharpe informs me that the tail of the young male of
D. Gaudichaudi is at first brown. Mr. Gould has described (ibid. vol. ii. p. 14, 20,
37) the sexes and the young of certain Black Cockatoos and of the King Lory,
with which the same rule prevails. Also Jerdon (‘Birds of India,’ vol. i. p. 260)
on the Palæornis rosa, in which the young are more like the female than the
male. See Audubon (‘Ornith. Biograph.’ vol. ii. p. 475) on the two sexes and the
young of Columba passerina.
231 I owe this information to Mr. Gould who shewed me the specimens; see also
his ‘Introduction to the Trochilidæ,’ 1861, p. 120.
232 Macgillivray, ‘Hist. Brit. Birds,’ vol. v. p. 207-214.
233 See his admirable paper in the ‘Journal of the Asiatic Soc. of Bengal,’ vol.
xix. 1850, p. 223; see also Jerdon, ‘Birds of India,’ vol. i. introduction, p. xxix. In
regard to Tanysiptera, Prof. Schlegel told Mr. Blyth that he could distinguish
several distinct races, solely by comparing the adult males.
234 See also Mr. Swinhoe, in ‘Ibis,’ July, 1863, p. 131; and a previous paper,
with an extract from a note by Mr. Blyth, in ‘Ibis,’ Jan. 1861, p. 52.
235 Wallace, ‘The Malay Archipelago,’ vol. ii. 1869, p. 394.
236 These species are described, with coloured figures, by M. F. Pollen, in ‘Ibis,’
1866, p. 275.
237 ‘Variation of Animals, &c., under Domestication,’ vol. i. p. 251.
238 Macgillivray, ‘Hist. British Birds,’ vol. i. p. 172-174.
239 See, on this subject, chap. xxiii. in the ‘Variation of Animals and Plants
under Domestication.’

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240 Audubon, ‘Ornith. Biography,’ vol. i. p. 193. Macgillivray, ‘Hist. Brit.
Birds,’ vol. iii. p. 85. See also the case before given of Indopicus carlotta.
241 ‘Westminster Review,’ July, 1867, and A. Murray, ‘Journal of Travel,’ 1868,
p. 83.
242 For the Australian species, see Gould’s ‘Handbook,’ &c., vol. ii. p. 178, 180,
186, and 188. In the British Museum specimens of the Australian Plain-wanderer
(Pedionomus torquatus) may be seen, shewing similar sexual differences.
243 Jerdon, ‘Birds of India,’ vol. iii. p. 596. Mr. Swinhoe, in ‘Ibis,’ 1865, p. 542;
1866, p. 131, 405.
244 Jerdon, ‘Birds of India,’ vol. iii. p. 677.
245 Gould’s ‘Handbook of the Birds of Australia,’ vol. ii. p. 275.
246 ‘The Indian Field,’ Sept. 1858, p. 3.
247 ‘Ibis,’ 1866, p. 298.
248 For these several statements, see Mr. Gould’s ‘Birds of Great Britain.’ Prof.
Newton informs me that he has long been convinced, from his own observations
and from those of others, that the males of the above-named species take either
the whole or a large share of the duties of incubation, and that they “shew much
greater devotion towards their young, when in danger, than do the females.” So it
is, as he informs me, with Limosa lapponica and some few other Waders, in
which the females are larger and have more strongly contrasted colours than the
males.
249 The natives of Ceram (Wallace, ‘Malay Archipelago,’ vol. ii. p. 150) assert
that the male and female sit alternately on the eggs; but this assertion, as Mr.
Bartlett thinks, may be accounted for by the female visiting the nest to lay her
eggs.
250 ‘The Student,’ April, 1870, p. 124.
251 See the excellent account of the habits of this bird under confinement, by
Mr. A. W. Bennett, in ‘Land and Water,’ May, 1868, p. 233.
252 Mr. Sclater, on the incubation of the Struthiones, ‘Proc. Zoo. Soc.’ June 9,
1863.
253 For the Milvago, see ‘Zoology of the Voyage of the “Beagle,”’ Birds, 1841,
p. 16. For the Climacteris and nightjar (Eurostopodus), see Gould’s ‘Handbook
of the Birds of Australia,’ vol. i. p. 602 and 97. The New Zealand shieldrake
(Tadorna variegata) offers a quite anomalous case: the head of the female is pure
white, and her back is redder than that of the male; the head of the male is of a
rich dark bronzed colour, and his back is clothed with finely pencilled slate-
coloured feathers, so that he may altogether be considered as the more beautiful
of the two. He is larger and more pugnacious than the female, and does not sit on
the eggs. So that in all these respects this species comes under our first class of
cases; but Mr. Sclater (‘Proc. Zool. Soc.’ 1866, p. 150) was much surprised to
observe that the young of both sexes, when about three months old, resembled in
their dark heads and necks the adult males, instead of the adult females; so that it

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would appear in this case that the females have been modified, whilst the males
and the young have retained a former state of plumage.
254 Jerdon, ‘Birds of India,’ vol. iii. p. 598.
255 Jerdon, ‘Birds of India,’ vol. i. p. 222, 228. Gould’s ‘Handbook of the Birds
of Australia,’ vol. i. 124, 130.
256 Gould, ibid. vol. ii. p. 37, 46, 56.
257 Audubon, ‘Ornith. Biography,’ vol. ii. p. 55.
258 ‘Variation of Animals and Plants under Domestication,’ vol. ii. p. 79.
259 Charlesworth, ‘Mag. of Nat. Hist.’ vol. i. 1837, p. 305, 306.
260 ‘Bulletin de la Soc. Vaudoise des Sc. Nat.’ vol. x. 1869, p. 132. The young of
the Polish swan, Cygnus immutabilis of Yarrell, are always white; but this
species, as Mr. Sclater informs me, is believed to be nothing more than a variety
of the Domestic Swan (Cygnus olor).
261 I am indebted to Mr. Blyth for information in regard to this genus. The
sparrow of Palestine belongs to the sub-genus Petronia.
262 For instance, the males of Tanagra æstiva and Fringilla cyanea require three
years, the male of Fringilla ciris four years, to complete their beautiful plumage.
(See Audubon, ‘Ornith. Biography,’ vol. i. p. 233, 280, 378.) The Harlequin duck
takes three years (ibid. vol. iii. p. 614). The male of the Gold pheasant, as I hear
from Mr. J. Jenner Weir, can be distinguished from the female when about three
months old, but he does not acquire his full splendour until the end of the
September in the following year.
263 Thus the Ibis tantalus and Grus Americanus take four years, the Flamingo
several years, and the Ardea Ludovicana two years, before they acquire their
perfect plumage. See Audubon, ibid. vol. i. p. 221; vol. iii. p. 133, 139, 211.
264 Mr. Blyth, in Charlesworth’s ‘Mag. of Nat. Hist.’ vol. i. 1837, p. 300. Mr.
Bartlett has informed me in regard to gold pheasants.
265 I have noticed the following cases in Audubon’s ‘Ornith. Biography. The
Redstart of America’ (Muscicapa ruticilla, vol. i. p. 203). The Ibis tantalus takes
four years to come to full maturity, but sometimes breeds in the second year (vol.
iii. p. 133). The Grus Americanus takes the same time, but breeds before
acquiring its full plumage (vol. iii. p. 211). The adults of Ardea cærulea are blue
and the young white; and white, mottled, and mature blue birds may all be seen
breeding together (vol. iv. p. 58): but Mr. Blyth informs me that certain herons
apparently are dimorphic, for white and coloured individuals of the same age
may be observed. The Harlequin duck (Anas histrionica, Linn.) takes three years
to acquire its full plumage, though many birds breed in the second year (vol. iii.
p. 614). The White-headed Eagle (Falco leucocephalus, vol. iii. p. 210) is
likewise known to breed in its immature state. Some species of Oriolus
(according to Mr. Blyth and Mr. Swinhoe, in ‘Ibis,’ July, 1863, p. 68) likewise
breed before they attain their full plumage.
266 See the last footnote.

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267 Other animals, belonging to quite distinct classes, are either habitually or
occasionally capable of breeding before they have fully acquired their adult
characters. This is the case with the young males of the salmon. Several
amphibians have been known to breed whilst retaining their larval structure.
Fritz Müller has shewn (‘Facts and Arguments for Darwin,’ Eng. trans. 1869, p.
79) that the males of several amphipod crustaceans become sexually mature
whilst young; and I infer that this is a case of premature breeding, because they
have not as yet acquired their fully-developed claspers. All such facts are highly
interesting, as bearing on one means by which species may undergo great
modifications of character, in accordance with Mr. Cope’s views, expressed
under the terms of the “retardation and acceleration of generic characters;” but I
cannot follow the views of this eminent naturalist to their full extent. See Mr.
Cope, “On the Origin of Genera,” from the ‘Proc. of Acad. Nat. Sc. of
Philadelphia,’ Oct. 1868.
268 Jerdon, ‘Birds of India,’ vol. iii. p. 507, on the peacock. Audubon, ibid. vol.
iii. p. 139, on the Ardea.
269 For illustrative cases see vol. iv. of Macgillivray’s ‘Hist. Brit. Birds;’ on
Tringa, &c., p. 229, 271; on the Machetes, p. 172; on the Charadrius hiaticula,
p. 118; on the Charadrius pluvialis, p. 94.
270 For the goldfinch of N. America, Fringilla tristis, Linn., see Audubon,
‘Ornith. Biography,’ vol. i. p. 172. For the Maluri, Gould’s 'Handbook of the
Birds of Australia,’ vol. i. p. 318.
271 I am indebted to Mr. Blyth for information in regard to the Buphus; see also
Jerdon, ‘Birds of India,’ vol. iii. p. 749. On the Anastomus, see Blyth, in ‘Ibis,’
1867, p. 173.
272 On the Alca, see Macgillivray, ‘Hist. Brit. Birds,’ vol. v. p. 347. On the
Fringilla leucophrys, Audubon, ibid. vol. ii. p. 89. I shall have hereafter to refer
to the young of certain herons and egrets being white.
273 ‘History of British Birds,’ vol. i. 1839, p. 159.
274 Blyth, in Charlesworth’s ‘Mag. of Nat. Hist.’ vol. i. 1837, p. 362; and from
information given to me by him.
275 Audubon, ‘Ornith. Biography,’ vol. i. p. 113.
276 Mr. C. A. Wright, in ‘Ibis,’ vol. vi. 1864, p. 65. Jerdon, ‘Birds of India,’ vol.
i. p. 515.
277 The following additional cases may be mentioned: the young males of
Tanagra rubra can be distinguished from the young females (Audubon, ‘Ornith.
Biography,’ vol. iv. p. 392), and so it is with the nestlings of a blue nuthatch,
Dendrophila frontalis of India (Jerdon, 'Birds of India,’ vol. i. p. 389). Mr. Blyth
also informs me that the sexes of the stonechat, Saxicola rubicola, are
distinguishable at a very early age.
278 ‘Westminster Review,’ July, 1867, p. 5.
279 ‘Ibis,’ 1859, vol. i. p. 429, et seq.

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280 No satisfactory explanation has ever been offered of the immense size, and
still less of the bright colours, of the toucan’s beak. Mr. Bates (‘The Naturalist on
the Amazons,’ vol. ii. 1863, p. 341) states that they use their beak for reaching
fruit at the extreme tips of the branches; and likewise, as stated by other authors,
for extracting eggs and young birds from the nests of other birds. But as Mr.
Bates admits, the beak “can scarcely be considered a very perfectly-formed
instrument for the end to which it is applied.” The great bulk of the beak, as
shewn by its breadth, depth, as well as length, is not intelligible on the view, that
it serves merely as an organ of prehension.
281 Ramphastos carinatus, Gould’s ‘Monograph of Ramphastidæ.’
282 On Larus, Gavia, and Sterna, see Macgillivray, ‘Hist. Brit. Birds,’ vol. v. p.
515, 584, 626. On the Anser hyperboreus, Audubon, ‘Ornith. Biography,’ vol. iv.
p. 562. On the Anastomus, Mr. Blyth, in ‘Ibis,’ 1867, p. 173.
283 It may be noticed that with vultures, which roam far and wide through the
higher regions of the atmosphere, like marine birds over the ocean, three or four
species are almost wholly or largely white, and many other species are black.
This fact supports the conjecture that these conspicuous colours may aid the
sexes in finding each other during the breeding-season.
284 ‘The Journal of Travel,’ edited by A. Murray, vol. i. 1868, p. 286.
285 See Jerdon on the genus Palæornis, ‘Birds of India,’ vol. i. p. 258-260.
286 The young of Ardea rufescens and A. cærulea of the U. States are likewise
white, the adults being coloured in accordance with their specific names.
Audubon (‘Ornith. Biography,’ vol. iii. p. 416; vol. iv. p. 58) seems rather
pleased at the thought that this remarkable change of plumage will greatly
“disconcert the systematists.”
287 I am greatly indebted to the kindness of Mr. Sclater for having looked over
these four chapters on birds, and the two following ones on mammals. By this
means I have been saved from making mistakes about the names of the species,
and from giving any facts which are actually known to this distinguished
naturalist to be erroneous. But of course he is not at all answerable for the
accuracy of the statements quoted by me from various authorities.
288 See Waterton’s account of two hares fighting, ‘Zoologist,’ vol. i. 1843, p.
211. On moles, Bell, ‘Hist. of British Quadrupeds,’ 1st edit. p. 100. On squirrels,
Audubon and Bachman, ‘Viviparous Quadrupeds of N. America,’ 1846, p. 269.
On beavers, Mr. A. H. Green, in ‘Journal of Lin. Soc. Zoolog.’ vol. x. 1869, p.
362.
289 On the battles of seals, see Capt. C. Abbott in ‘Proc. Zool. Soc.’ 1868, p.
191; also Mr. R. Brown, ibid. 1869, p. 436; also L. Lloyd, 'Game Birds of
Sweden,’ 1867, p. 412; also Pennant. On the sperm-whale, see Mr. J. H.
Thompson, in ‘Proc. Zool. Soc.’ 1867, p. 246.
290 See Scrope (‘Art of Deer-stalking,’ p. 17) on the locking of the horns with
the Cervus elaphus. Richardson, in ‘Fauna Bor. Americana,’ 1829, p. 252, says
that the wapiti, moose, and reindeer have been found thus locked together. Sir A.

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Smith found at the Cape of Good Hope the skeletons of two gnus in the same
condition.
291 Mr. Lamont (‘Seasons with the Sea-Horses,’ 1861, p. 143) says that a good
tusk of the male walrus weighs 4 pounds, and is longer than that of the female,
which weighs about 3 pounds. The males are described as fighting ferociously.
On the occasional absence of the tusks in the female, see Mr. R. Brown, ‘Proc.
Zool. Soc.’ 1868, p. 429.
292 Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 283.
293 Mr. R. Brown, in ‘Proc. Zool. Soc.’ 1869, p. 553.
294 Owen on the Cachalot and Ornithorhynchus, ibid. vol. iii. p. 638, 641.
295 On the structure and shedding of the horns of the reindeer, Hoffberg,
'Amœnitates Acad.’ vol. iv. 1788, p. 149. See Richardson, ‘Fauna Bor.
Americana,’ p. 241, in regard to the American variety or species; also Major W.
Ross King, ‘The Sportsman in Canada,’ 1866, p. 80.
296 Isidore Geoffroy St.-Hilaire, ‘Essais de Zoolog. Générale,’ 1841, p. 513.
Other masculine characters, besides the horns, are sometimes similarly
transferred to the female; thus Mr. Boner, in speaking of an old female chamois
(‘Chamois Hunting in the Mountains of Bavaria,’ 1860, 2nd edit. p. 363), says,
“not only was the head very male-looking, but along the back there was a ridge
of long hair, usually to be found only in bucks.”
297 On the Cervulus, Dr. Gray, ‘Catalogue of the Mammalia in British
Museum,’ part iii. p. 220. On the Cervus Canadensis or Wapiti see Hon. J. D.
Caton, ‘Ottawa Acad. of Nat. Sciences,’ May, 1868, p. 9.
298 For instance the horns of the female Ant. Euchore resemble those of a
distinct species, viz. the Ant. Dorcas var. Corine, see Desmarest, 'Mammalogie,’
p. 455.
299 Gray, ‘Catalogue Mamm. Brit. Mus.’ part iii. 1852, p. 160.
300 Richardson, ‘Fauna Bor. Americana,’ p. 278.
301 ‘Land and Water,’ 1867, p. 346.
302 Sir Andrew Smith, ‘Zoology of S. Africa,’ pl. xix. Owen, ‘Anatomy of
Vertebrates,’ vol. iii. p. 624.
303 Sir J. Emerson Tennent, ‘Ceylon,’ 1859, vol. ii. p. 274. For Malacca,
‘Journal of Indian Archipelago,’ vol. iv. p. 357.
304 ‘Calcutta Journal of Nat. Hist.’ vol. ii. 1843, p. 526.
305 Mr. Blyth, in ‘Land and Water,’ March, 1867, p. 134, on the authority of
Capt. Hutton and others. For the wild Pembrokeshire goats see the ‘Field,’ 1869,
p. 150.
306 M. E. M. Bailly, “sur l’usage des Cornes,” &c., ‘Annal. des Sc. Nat.’ tom. ii.
1824, p. 369.

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307 Owen, on the Horns of Red-deer, ‘British Fossil Mammals,’ 1846, p. 478;
‘Forest Creatures,’ by Charles Boner, 1861, p. 76, 62. Richardson on the Horns
of the Reindeer, ‘Fauna Bor. Americana,’ 1829, p. 210.
308 Hon. J. D. Caton (‘Ottawa Acad. of Nat. Science,’ May, 1868, p. 9), says that
the American deer fight with their fore-feet, after “the question of superiority has
been once settled and acknowledged in the herd.” Bailly, “Sur l’usage des
Cornes,” ‘Annales des Sc. Nat.’ tom. ii. 1824, p. 371.
309 See a most interesting account in the Appendix to Hon. J. D. Caton’s paper,
as above quoted.
310 ‘The American Naturalist,’ Dec. 1869, p. 552.
311 Pallas, ‘Spicilegia Zoologica,’ fasc. xiii. 1779, p. 18.
312 Lamont, ‘Seasons with the Sea-Horses,’ 1861, p. 141.
313 See also Corse (‘Philosoph. Transact.’ 1799, p. 212) on the manner in which
the short-tusked Mooknah variety of the elephant attacks other elephants.
314 Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 349.
315 See Rüppell (in ‘Proc. Zoolog. Soc.’ Jan. 12, 1836, p. 3) on the canines in
deer and antelopes, with a note by Mr. Martin on a female American deer. See
also Falconer (‘Palæont. Memoirs and Notes,’ vol. i. 1868, p. 576) on canines in
an adult female deer. In old males of the musk-deer the canines (Pallas, ‘Spic.
Zoolog.’ fasc. xiii. 1779, p. 18) sometimes grow to the length of three inches,
whilst in old females a rudiment projects scarcely half an inch above the gums.
316 Emerson Tennent, ‘Ceylon,’ 1859, vol. ii. p. 275; Owen, ‘British Fossil
Mammals,’ 1846, p. 245.
317 Richardson, ‘Fauna Bor. Americana,’ on the moose, Alces palmata, p. 236,
237; also on the expanse of the horns ‘Land and Water,’ 1869, p. 143. See also
Owen, ‘British Fossil Mammals,’ on the Irish elk, p. 447, 455.
318 ‘Forest Creatures,’ by C. Boner, 1861, p. 60.
319 See the very interesting paper by Mr. J. A. Allen in ‘Bull. Mus. Comp.
Zoolog. of Cambridge; United States,’ vol. ii. No. 1, p. 82. The weights were
ascertained by a careful observer, Capt. Bryant.
320 ‘Animal Economy,’ p. 45.
321 See also Richardson’s ‘Manual on the Dog,’ p. 59. Much valuable
information on the Scottish deerhound is given by Mr. McNeill, who first called
attention to the inequality in size between the sexes, in Scrope’s ‘Art of Deer
Stalking.’ I hope that Mr. Cupples will keep to his intention of publishing a full
account and history of this famous breed.
322 Brehm, ‘Thierleben,’ B. ii. s. 729-732.
323 See Mr. Wallace’s interesting account of this animal, ‘The Malay
Archipelago,’ 1869, vol. i. p. 435.

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324 ‘The Times,’ Nov. 10th, 1857. In regard to the Canada lynx, see Audubon
and Bachman, ‘Quadrupeds of N. America,’ 1846, p. 139.
325 Dr. Murie, on Otaria, ‘Proc. Zoolog. Soc.’ 1869, p. 109. Mr. J. A. Allen, in
the paper above quoted (p. 75), doubts whether the hair, which is longer on the
neck in the male than in the female, deserves to be called a mane.
326 Mr. Boner in his excellent description of the habits of the red-deer in
Germany (‘Forest Creatures,’ 1861, p. 81) says, “while the stag is defending his
rights against one intruder, another invades the sanctuary of his harem, and
carries off trophy after trophy.” Exactly the same thing occurs with seals, see Mr.
J. A. Allen, ibid. p. 100.
327 Mr. J. A. Allen in ‘Bull. Mus. Comp. Zoolog. of Cambridge, United States,’
vol. ii. No. 1, p. 99.
328 ‘Dogs: their Management,’ by E. Mayhew, M.R.C.V.S., 2nd edit. 1864, p.
187-192.
329 Quoted by Alex. Walker ‘On Intermarriage,’ 1838, p. 276; see also p. 244.
330 ‘Traité de l’Héréd. Nat.’ tom. ii. 1850, p. 296.
331 ‘Amœnitates Acad.’ vol. iv. 1788, p. 160.
332 Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 585.
333 Ibid. p. 595.
334 See, for instance, Major W. Ross King (‘The Sportsman in Canada,’ 1866, p.
53, 131) on the habits of the moose and wild reindeer.
335 Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 600.
336 Mr. Green, in ‘Journal of Linn. Soc.’ vol. x. Zoology, 1869, p. 362.
337 C. L. Martin, ‘General Introduction to the Nat. Hist. of Mamm. Animals,’
1841, p. 431.
338 ‘Naturgeschichte der Säugethiere von Paraguay,’ 1830, s. 15, 21.
339 On the sea-elephant, see an article by Lesson, in ‘Dict. Class. Hist. Nat.’
tom. xiii. p. 418. For the Cystophora or Stemmatopus, see Dr. Dekay, ‘Annals of
Lyceum of Nat. Hist. New York,’ vol. i. 1824, p. 94. Pennant has also collected
information from the sealers on this animal. The fullest account is given by Mr.
Brown, who doubts about the rudimentary condition of the bladder in the female,
in ‘Proc. Zoolog. Soc.’ 1868, p. 435.
340 As with the castoreum of the beaver, see Mr. L. H. Morgan’s most
interesting work, ‘The American Beaver,’ 1868, p. 300. Pallas (‘Spic. Zoolog.’
fasc. viii. 1779, p. 23) has well discussed the odoriferous glands of mammals.
Owen (‘Anat. of Vertebrates,’ vol. iii. p. 634) also gives an account of these
glands, including those of the elephant, and (p. 763) those of shrew-mice.
341 Rengger, ‘Naturgeschichte der Säugethiere von Paraguay,’ 1830, s. 355. This
observer also gives some curious particulars in regard to the odour emitted.

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342 Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 632. See, also, Dr. Murie’s
observations on their glands in ‘Proc. Zoolog. Soc.’ 1870, p. 340. Desmarest, On
the Antilope subgutturosa, ‘Mammalogie,’ 1820, p. 455.
343 Pallas, ‘Spicilegia Zoolog.’ fasc. xiii. 1799, p. 24; Desmoulins, 'Dict. Class.
d’Hist. Nat.’ tom. iii. p. 586.
344 Dr. Gray, ‘Gleanings from the Menagerie at Knowsley,’ pl. 28.
345 Judge Caton on the wapiti, ‘Transact. Ottawa Acad. Nat. Sciences,’ 1868, p.
36, 40; Blyth, ‘Land and Water,’ on Capra ægagrus, 1867, p. 37.
346 ‘Hunter’s Essays and Observations,’ edited by Owen, 1861, vol. i. p. 236.
347 See Dr. Gray’s ‘Cat. of Mammalia in British Museum,’ part iii. 1852, p. 144.
348 Rengger, ‘Säugethiere,’ &c., s. 14; Desmarest, ‘Mammalogie,’ p. 66.
349 See the chapters on these several animals in vol. i. of my ‘Variation of
Animals and Plants under Domestication;’ also vol. ii. p. 73; also chap. xx. on
the practice of selection by semi-civilised people. For the Berbura goat, see Dr.
Gray, ‘Catalogue,’ ibid. p. 157.
350 Osphranter rufus, Gould, ‘Mammals of Australia,’ vol. ii. 1863. On the
Didelphis, Desmarest, ‘Mammalogie,’ p. 256.
351 ‘Annals and Mag. of Nat. Hist.’ Nov. 1867, p. 325. On the Mus minutus,
Desmarest, ‘Mammalogie,’ p. 304.
352 J. A. Allen, in ‘Bulletin of Mus. Comp. Zoolog. of Cambridge, United
States,’ 1869, p. 207.
353 Desmarest, ‘Mammalogie,’ 1820, p. 223. On Felis mitis, Rengger, ibid. s.
194.
354 Dr. Murie on the Otaria, ‘Proc. Zool. Soc.’ 1869, p. 108. Mr. R. Brown, on
the P. groenlandica, ibid. 1868, p. 417. See also on the colours of seals,
Desmarest, ibid. p. 243, 249.
355 Judge Caton, in ‘Trans. Ottawa Acad. of Nat. Sciences,’ 1868, p. 4.
356 Dr. Gray, ‘Cat. of Mamm. in Brit. Mus.’ part iii. 1852, p. 134-142; also Dr.
Gray, ‘Gleanings from the Menagerie of Knowsley,’ in which there is a splendid
drawing of the Oreas derbyanus: see the text on Tragelaphus. For the Cape
Eland (Oreas canna), see Andrew Smith, 'Zoology of S. Africa,’ pl. 41 and 42.
There are also many of these antelopes in the Zoological Society’s Gardens.
357 On the Ant. niger, see ‘Proc. Zool. Soc.’ 1850, p. 133. With respect to an
allied species, in which there is an equal sexual difference in colour, see Sir S.
Baker, ‘The Albert Nyanza,’ 1866, vol. ii. p. 327. For the A. sing-sing, Gray,
‘Cat. B. Mus.’ p. 100. Desmarest, Mammalogie,’ p. 468, on the A. caama.
Andrew Smith, ‘Zoology of S. Africa,’ on the Gnu.
358 ‘Ottawa Academy of Sciences,’ May 21, 1868, p. 3, 5.

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359 S. Müller, on the Banteng, ‘Zoog. Indischen Archipel.’ 1839-1844, tab. 35;
see also Raffles, as quoted by Mr. Blyth, in ‘Land and Water,’ 1867, p. 476. On
goats, Dr. Gray, ‘Cat. Brit. Mus.’ p. 146; Desmarest, 'Mammalogie,’ p. 482. On
the Cervus paludosus, Rengger, ibid. s. 345.
360 Sclater, ‘Proc. Zool. Soc.’ 1866, p. 1. The same fact has also been fully
ascertained by MM. Pollen and van Dam.
361 On Mycetes, Rengger, ibid. s. 14; and Brehm, ‘Illustrirtes Thierleben,’ B. i.
s. 96, 107. On Ateles, Desmarest, ‘Mammalogie,’ p. 75. On Hylobates, Blyth,
‘Land and Water,’ 1867, p. 135. On the Semnopithecus, S. Müller, ‘Zoog.
Indischen Archipel.’ tab. x.
362 Gervais, ‘Hist. Nat. des Mammifères,’ 1854, p. 103. Figures are given of the
skull of the male. Desmarest, ‘Mammalogie,’ p. 70. Geoffroy St.-Hilaire and F.
Cuvier, ‘Hist. Nat. des Mamm.’ 1824, tom. i.
363 ‘The Variation of Animals and Plants under Domestication,’ 1868, vol. ii. p.
102, 103.
364 ‘Essays and Observations by J. Hunter,’ edited by Owen, 1861, vol. i. p. 194.
365 Sir S. Baker, ‘The Nile Tributaries of Abyssinia,’ 1867.
366 Fiber zibethicus, Audubon and Bachman, ‘The Quadrupeds of N. America,’
1846, p. 109.
367 ‘Novæ species Quadrupedum e Glirium ordine,’ 1778, p. 7. What I have
called the roe is the Capreolus Sibiricus subecaudatus of Pallas.
368 See the fine plates in A. Smith’s ‘Zoology of S. Africa,’ and Dr. Gray’s
‘Gleanings from the Menagerie of Knowsley.’
369 ‘Westminster Review,’ July 1, 1867, p. 5.
370 ‘Travels in South Africa,’ 1824, vol. ii. p. 315.
371 Dr. Gray, ‘Gleanings from the Menagerie of Knowsley,’ p. 64. Mr. Blyth, in
speaking (‘Land and Water,’ 1869, p. 42) of the hog-deer of Ceylon, says it is
more brightly spotted with white than the common hog-deer, at the season when
it renews its horns.
372 Falconer and Cautley, ‘Proc. Geolog. Soc.’ 1843; and Falconer’s 'Pal.
Memoirs,’ vol. i. p. 196.
373 ‘The Variation of Animals and Plants under Domestication,’ 1868, vol. i. p.
61-64.
374 ‘Proc. Zool. Soc.’ 1862, p. 164. See, also, Dr. Hartmann, ‘Ann. d. Landw.’
Bd. xliii. s. 222.
375 I observed this fact in the Zoological Gardens; and numerous cases may be
seen in the coloured plates in Geoffroy St.-Hilaire and F. Cuvier, ‘Hist. Nat. des
Mammifères,’ tom. i. 1824.
376 Bates, ‘The Naturalist on the Amazons,’ 1863, vol. ii. p. 310.

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377 I have seen most of the above-named monkeys in the Zoological Society’s
Gardens. The description of the Semnopithecus nemæus is taken from Mr. W. C.
Martin’s ‘Nat. Hist. of Mammalia,’ 1841, p. 460; see also p. 475, 523.
378 Schaaffhausen, translation in ‘Anthropological Review,’ Oct. 1868, p. 419,
420, 427.
379 Ecker, translation in ‘Anthropological Review,’ Oct. 1868, p. 351-356. The
comparison of the form of the skull in men and women has been followed out
with much care by Welcker.
380 Ecker and Welcker, ibid. p. 352, 355; Vogt, ‘Lectures on Man,’ Eng. translat.
p. 81.
381 Schaaffhausen, ‘Anthropolog. Review,’ ibid. p. 429.
382 Pruner-Bey, on negro infants, as quoted by Vogt, ‘Lectures on Man,’ Eng.
translat. 1864, p. 189: for further facts on negro infants, as quoted from
Winterbottom and Camper, see Lawrence, ‘Lectures on Physiology,’ &c. 1822, p.
451. For the infants of the Guaranys, see Rengger, ‘Säugethiere,’ &c. s. 3. See
also Godron, ‘De l’Espèce,’ tom. ii. 1859, p. 253. For the Australians, Waitz,
‘Introduct. to Anthropology,’ Eng. translat. 1863, p. 99.
383 Rengger, ‘Säugethiere,’ &c. 1830, s. 49.
384 As in Macacus cynomolgus (Desmarest, ‘Mammalogie,’ p. 65) and in
Hylobates agilis (Geoffroy St.-Hilaire and F. Cuvier, ‘Hist. Nat. des Mamm.’
1824, tom. i. p. 2).
385 ‘Anthropological Review,’ Oct. 1868, p. 353.
386 Mr. Blyth informs me that he has never seen more than one instance of the
beard, whiskers, &c., in a monkey becoming white with old age, as is so
commonly the case with us. This, however, occurred in an aged and confined
Macacus cynomolgus, whose moustaches were “remarkably long and human-
like.” Altogether this old monkey presented a ludicrous resemblance to one of
the reigning monarchs of Europe, after whom he was universally nick-named. In
certain races of man the hair on the head hardly ever becomes grey; thus Mr. D.
Forbes has never seen, as he informs me, an instance with the Aymaras and
Quechuas of S. America.
387 This is the case with the females of several species of Hylobates, see
Geoffroy St.-Hilaire and F. Cuvier, ‘Hist. Nat. des Mamm.’ tom. i. See, also, on
H. lar. ‘Penny Encyclopedia,’ vol. ii. p. 149, 150.
388 The results were deduced by Dr. Weisbach from the measurements made by
Drs. K. Scherzer and Schwarz, see ‘Reise der Novara: Anthropolog. Theil,’
1867, s. 216, 231, 234, 236, 239, 269.
389 ‘Voyage to St. Kilda,’ (3rd edit. 1753) p. 37.
390 Sir J. E. Tennent, ‘Ceylon,’ vol. ii. 1859, p. 107.
391 Quatrefages, ‘Revue des Cours Scientifiques,’ Aug. 29, 1868, p. 630; Vogt,
‘Lectures on Man,’ Eng. translat. p. 127.

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392 On the beards of negroes, Vogt, ‘Lectures,’ &c. ibid. p. 127; Waitz,
'Introduct. to Anthropology,’ Engl. translat. 1863, vol. i. p. 96. It is remarkable
that in the United States (‘Investigations in Military and Anthropological
Statistics of American Soldiers,’ 1869, p. 569) the pure negroes and their crossed
offspring seem to have bodies almost as hairy as those of Europeans.
393 Wallace, ‘The Malay Arch.’ vol. ii. 1869, p. 178.
394 Dr. J. Barnard Davis on Oceanic Races, in ‘Anthropolog. Review,’ April,
1870, p. 185, 191.
395 Catlin, ‘North American Indians,’ 3rd edit. 1842, vol. ii. p. 227. On the
Guaranys, see Azara, ‘Voyages dans l’Amérique Mérid.’ tom. ii. 1809, p. 58; also
Rengger, ‘Säugethiere von Paraguay,’ s. 3.
396 Prof. and Mrs. Agassiz (‘Journey in Brazil,’ p. 530) remark that the sexes of
the American Indians differ less than those of the negroes and of the higher
races. See also Rengger, ibid. p. 3, on the Guaranys.
397 Rütimeyer, ‘Die Grenzen der Thierwelt; eine Betrachtung zu Darwin’s
Lehre,’ 1868, s. 54.
398 ‘A Journey from Prince of Wales Fort,’ 8vo. edit. Dublin, 1796, p. 104. Sir J.
Lubbock (‘Origin of Civilisation,’ 1870, p. 69) gives other and similar cases in
North America. For the Guanas of S. America see Azara, ‘Voyages,’ &c. tom. ii.
p. 94.
399 On the fighting of the male gorillas, see Dr. Savage, in ‘Boston Journal of
Nat. Hist.’ vol. v. 1847, p. 423. On Presbytis entellus, see the ‘Indian Field,’
1859, p. 146.
400 J. Stuart Mill remarks (‘The Subjection of Women,’ 1869, p. 122), “the
things in which man most excels woman are those which require most plodding,
and long hammering at single thoughts.” What is this but energy and
perseverance?
401 An observation by Vogt bears on this subject: he says, it is a “remarkable
circumstance, that the difference between the sexes, as regards the cranial cavity,
increases with the development of the race, so that the male European excels
much more the female, than the negro the negress. Welcker confirms this
statement of Huschke from his measurements of negro and German skulls.” But
Vogt admits (‘Lectures on Man,’ Eng. translat. 1864, p. 81) that more
observations are requisite on this point.
402 Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 603.
403 ‘Journal of the Anthropolog. Soc.’ April, 1869, p. lvii. and lxvi.
404 Dr. Scudder, “Notes on Stridulation,” in ‘Proc. Boston Soc. of Nat. Hist.’
vol. xi. April, 1868.
405 Given in W. C. L. Martin’s ‘General Introduct. to Nat. Hist. of Mamm.
Animals,’ 1841, p. 432; Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 600.
406 Helmholtz, ‘Théorie Phys. de la Musique,’ 1868, p. 187.

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407 Mr. R. Brown, in ‘Proc. Zoo. Soc.’ 1868, p. 410.
408 ‘Journal of Anthropolog. Soc.’ Oct. 1870, p. clv. See also the several later
chapters in Sir John Lubbock’s ‘Prehistoric Times,’ second edition, 1869, which
contain an admirable account of the habits of savages.
409 Since this chapter has been printed I have seen a valuable article by Mr.
Chauncey Wright (‘North Amer. Review,’ Oct. 1870, page 293), who, in
discussing the above subject, remarks, “There are many consequences of the
ultimate laws or uniformities of nature through which the acquisition of one
useful power will bring with it many resulting advantages as well as limiting
disadvantages, actual or possible, which the principle of utility may not have
comprehended in its action.” This principle has an important bearing, as I have
attempted to shew in the second chapter of this work, on the acquisition by man
of some of his mental characteristics.
410 See the very interesting discussion on the Origin and Function of Music, by
Mr. Herbert Spencer, in his collected ‘Essays,’ 1858, p. 359. Mr. Spencer comes
to an exactly opposite conclusion to that at which I have arrived. He concludes
that the cadences used in emotional speech afford the foundation from which
music has been developed; whilst I conclude that musical notes and rhythm were
first acquired by the male or female progenitors of mankind for the sake of
charming the opposite sex. Thus musical tones became firmly associated with
some of the strongest passions an animal is capable of feeling, and are
consequently used instinctively, or through association, when strong emotions
are expressed in speech. Mr. Spencer does not offer any satisfactory explanation,
nor can I, why high or deep notes should be expressive, both with man and the
lower animals, of certain emotions. Mr. Spencer gives also an interesting
discussion on the relations between poetry, recitative, and song.
411 Rengger, ‘Säugethiere von Paraguay,’ s. 49.
412 See an interesting discussion on this subject by Häckel, ‘Generelle Morph.’
B. ii. 1866, s. 246.
413 A full and excellent account of the manner in which savages in all parts of
the world ornament themselves is given by the Italian traveller, Prof.
Mantegazza, ‘Rio de la Plata, Viaggi e Studi,’ 1867, p. 525-545; all the following
statements, when other references are not given, are taken from this work. See,
also, Waitz, ‘Introduct. to Anthropolog.’ Eng. transl. vol. i. 1863, p. 275, et
passim. Lawrence also gives very full details in his ‘Lectures on Physiology,’
1822. Since this chapter was written Sir J. Lubbock has published his 'Origin of
Civilisation,’ 1870, in which there is an interesting chapter on the present
subject, and from which (p. 42, 48) I have taken some facts about savages dyeing
their teeth and hair, and piercing their teeth.
414 Humboldt, ‘Personal Narrative,’ Eng. translat. vol. iv. p. 515; on the
imagination shewn in painting the body, p. 522; on modifying the form of the
calf of the leg, p. 466.
415 ‘The Nile Tributaries,’ 1867; ‘The Albert N’yanza,’ 1866, vol. i. p. 218.
416 Quoted by Prichard, ‘Phys. Hist. of Mankind,’ 4th. edit. vol. i. 1851, p. 321.

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417 On the Papuans, Wallace, ‘The Malay Archipelago,’ vol. ii. p. 445. On the
coiffure of the Africans, Sir S. Baker, ‘The Albert N’yanza,’ vol. i. p. 210.
418 ‘Travels,’ p. 533.
419 ‘The Albert N’yanza,’ 1866, vol. i. p. 217.
420 Livingstone, ‘British Association,’ 1860; report given in the 'Athenæum,’
July 7, 1860, p. 29.
421 Sir S. Baker (ibid. vol. i. p. 210) speaking of the natives of Central Africa
says, “every tribe has a distinct and unchanging fashion for dressing the hair.”
See Agassiz (‘Journey in Brazil,’ 1868, p. 318) on the invariability of the
tattooing of the Amazonian Indians.
422 Rev. R. Taylor, ‘New Zealand and its Inhabitants,’ 1855, p. 152.
423 Mantegazza, ‘Viaggi e Studi,’ p. 542.
424 ‘Travels in S. Africa,’ 1824; vol. i. p. 414.
425 See, for references, ‘Gerland über das Aussterben der Naturvölker,’ 1868, s.
51, 53, 55; also Azara, ‘Voyages,’ &c. tom. ii. p. 116.
426 On the vegetable productions used by the North-Western American Indians,
‘Pharmaceutical Journal,’ vol. x.
427 ‘A Journey from Prince of Wales Fort,’ 8vo. edit. 1796, p. 89.
428 Quoted by Prichard, ‘Phys. Hist. of Mankind,’ 3rd edit. vol. iv. 1844, p. 519;
Vogt, ‘Lectures on Man,’ Eng. translat. p. 129. On the opinion of the Chinese on
the Cingalese, E. Tennent, ‘Ceylon,’ vol. ii. 1859, p. 107.
429 Prichard, as taken from Crawfurd and Finlayson, ‘Phys. Hist. of Mankind,’
vol. iv. p. 534, 535.
430 “Idem illustrissimus viator dixit mihi præcinctorium vel tabula fæminæ,
quod nobis teterrimum est, quondam permagno æstimari ab hominibus in hac
gente. Nunc res mutata est, et censet talem conformationem minime optandam
est.”
431 ‘The Anthropological Review,’ November, 1864, p. 237. For additional
references, see Waitz, ‘Introduct. to Anthropology,’ Eng. translat. 1863, vol. i. p.
105.
432 ‘Mungo Park’s Travels in Africa,’ 4to. 1816, p. 53, 131. Burton’s statement
is quoted by Schaaffhausen, ‘Archiv für Anthropolog.’ 1866, s. 163. On the
Banyai, Livingstone, ‘Travels,’ p. 64. On the Kafirs, the Rev. J. Shooter, ‘The
Kafirs of Natal and the Zulu Country,’ 1857 p. 1.
433 For the Javanese and Cochin-Chinese, see Waitz, ‘Introduct. to
Anthropology,’ Eng. translat. vol. i. p. 305. On the Yura-caras, A. d’Orligny, as
quoted in Prichard, ‘Phys. Hist. of Mankind,’ vol. v. 3rd edit. p. 476.
434 ‘North American Indians,’ by G. Catlin, 3rd edit. 1842, vol. i. p. 49; vol. ii.
p. 227. On the natives of Vancouver Island, see Sproat, 'Scenes and Studies of

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Savage Life,’ 1868, p. 25. On the Indians of Paraguay, Azara, ‘Voyages,’ tom. ii.
p. 105.
435 On the Siamese, Prichard, ibid. vol. iv. p. 533. On the Japanese, Veitch in
‘Gardeners’ Chronicle,’ 1860, p. 1104. On the New Zealanders Mantegazza,
‘Viaggi e Studi,’ 1867, p. 526. For the other nations mentioned, see references in
Lawrence, ‘Lectures on Physiology,’ &c. 1822, p. 272.
436 Lubbock, ‘Origin of Civilisation,’ 1870, p. 321.
437 Dr. Barnard Davis quotes Mr. Pritchard and others for these facts in regard
to the Polynesians, in ‘Anthropological Review,’ April, 1870, p. 185, 191.
438 Ch. Comte has remarks to this effect in his ‘Traité de Législation,’ 3rd edit.
1837, p. 136.
439 The Fuegians, as I have been informed by a missionary who long resided
with them, consider European women as extremely beautiful; but from what we
have seen of the judgment of the other aborigines of America, I cannot but think
that this must be a mistake, unless indeed the statement refers to the few
Fuegians who have lived for some time with Europeans, and who must consider
us as superior beings. I should add that a most experienced observer, Capt.
Burton, believes that a woman whom we consider beautiful is admired
throughout the world, 'Anthropological Review,’ March, 1864, p. 245.
440 ‘Personal Narrative,’ Eng. translat. vol. iv. p. 518, and elsewhere.
Mantegazza, in his ‘Viaggi e Studi,’ 1867, strongly insists on this same principle.
441 On the skulls of the American tribes, see Nott and Gliddon, 'Types of
Mankind,’ 1854, p. 440; Prichard, ‘Phys. Hist. of Mankind,’ vol. i. 3rd edit. p.
321; on the natives of Arakhan, ibid. vol. iv. p. 537. Wilson, ‘Physical
Ethnology,’ Smithsonian Institution, 1863, p. 288; on the Fijians, p. 290. Sir J.
Lubbock (‘Prehistoric Times,’ 2nd edit. 1869, p. 506) gives an excellent résumé
on this subject.
442 On the Huns, Godron, ‘De l’Espèce,’ tom. ii. 1859, p. 300. On the Tahitians,
Waitz, ‘Anthropolog.’ Eng. translat. vol. i. p. 305. Marsden, quoted by Prichard,
‘Phys. Hist. of Mankind,’ 3rd edit. vol. v. p. 67. Lawrence, ‘Lectures on
Physiology,’ p. 337.
443 This fact was ascertained in the ‘Reise der Novara: Anthropolog. Theil,’ Dr.
Weisbach, 1867, s. 265.
444 ‘Smithsonian Institution, 1863, p. 289. On the fashions of Arab women, Sir
S. Baker, ‘The Nile Tributaries,’ 1867, p. 121.
445 ‘The Variation of Animals and Plants under Domestication,’ vol. i. p. 214;
vol. ii. p. 240.
446 Schaaffhausen, ‘Archiv für Anthropologie,’ 1866, s. 164.
447 Mr. Bain has collected (‘Mental and Moral Science,’ 1868, p. 304-314)
about a dozen more or less different theories of the idea of beauty; but none are
quite the same with that here given.

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448 These quotations are taken from Lawrence (‘Lectures on Physiology,’ &c.
1822, p. 393), who attributes the beauty of the upper classes in England to the
men having long selected the more beautiful women.
449 “Anthropologie,” ‘Revue des Cours Scientifiques,’ Oct. 1868, p. 721.
450 ‘The Variation of Animals and Plants under Domestication,’ vol. i. p. 207.
451 Sir J. Lubbock, ‘The Origin of Civilisation,’ 1870, chap. iii. especially p. 60-
67. Mr. M’Lennan, in his extremely valuable work on 'Primitive Marriage,’
1865, p. 163, speaks of the union of the sexes “in the earliest times as loose,
transitory, and in some degree promiscuous.” Mr. M’Lennan and Sir J. Lubbock
have collected much evidence on the extreme licentiousness of savages at the
present time. Mr. L. H. Morgan, in his interesting memoir on the classificatory
system of relationship (‘Proc. American Acad. of Sciences,’ vol. vii. Feb. 1868,
p. 475) concludes that polygamy and all forms of marriage during primeval
times were essentially unknown. It appears, also, from Sir J. Lubbock’s work,
that Bachofen likewise believes that communal intercourse originally prevailed.
452 Address to British Association ‘On the Social and Religious Condition of the
Lower Races of Man,’ 1870, p. 20.
453 ‘Origin of Civilisation,’ 1870, p. 86. In the several works above quoted there
will be found copious evidence on relationship through the females alone, or
with the tribe alone.
454 Brehm (‘Illust. Thierleben,’ B. i. p. 77) says Cynocephalus hamadryas lives
in great troops containing twice as many adult females as adult males. See
Rengger on American polygamous species, and Owen (‘Anat. of Vertebrates,’
vol. iii. p. 746) on American monogamous species. Other references might be
added.
455 Dr. Savage, in ‘Boston Journal of Nat. Hist.’ vol. v. 1845-47, p. 423.
456 ‘Prehistoric Times,’ 1869, p. 424.
457 Mr. M’Lennan, ‘Primitive Marriage,’ 1865. See especially on exogamy and
infanticide, p. 130, 138, 165.
458 Dr. Gerland (‘Ueber das Aussterben der Naturvölker,’ 1868) has collected
much information on infanticide, see especially s. 27, 51, 54. Azara (‘Voyages,’
&c. tom. ii. p. 94, 116) enters in detail on the motives. See also M’Lennan (ibid.
p. 139) for cases in India.
459 ‘Primitive Marriage,’ p. 208; Sir J. Lubbock, ‘Origin of Civilisation,’ p. 100.
See also Mr. Morgan, loc. cit., on former prevalence of polyandry.
460 ‘Voyages,’ &c. tom. ii. p. 92-95.
461 Burchell says (‘Travels in S. Africa, vol. ii. 1824, p. 58), that among the wild
nations of Southern Africa, neither men nor women ever pass their lives in a
state of celibacy. Azara (‘Voyages dans l’Amérique Merid.’ tom. ii. 1809, p. 21)
makes precisely the same remark in regard to the wild Indians of South America.
462 ‘Anthropological Review,’ Jan. 1870, p. xvi.

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463 ‘The Variation of Animals and Plants under Domestication,’ vol. ii. p. 210-
217.
464 An ingenious writer argues, from a comparison of the pictures of Raphael,
Rubens, and modern French artists, that the idea of beauty is not absolutely the
same even throughout Europe: see the ‘Lives of Haydn and Mozart,’ by M.
Bombet, English translat. p. 278.
465 Azara, ‘Voyages,’ &c. tom. ii. p. 23. Dobrizhoffer, ‘An Account of the
Abipones,’ vol. ii. 1822, p. 207. Williams on the Fiji Islanders, as quoted by
Lubbock, ‘Origin of Civilisation,’ 1870, p. 79. On the Fuegians, King and
Fitzroy, ‘Voyages of the Adventure and Beagle,’ vol. ii. 1839, p. 182. On the
Kalmucks, quoted by M’Lennan, 'Primitive Marriage,’ 1865, p. 32. On the
Malays, Lubbock, ibid. p. 76. The Rev. J. Shooter, ‘On the Kafirs of Natal,’
1857, p. 52-60. On the Bushwomen, Burchell, ‘Travels in S. Africa,’ vol. ii.
1824, p. 59.
466 ‘Contributions to the Theory of Natural Selection,’ 1870, p. 346. Mr.
Wallace believes (p. 350) “that some intelligent power has guided or determined
the development of man;” and he considers the hairless condition of the skin as
coming under this head. The Rev. T. R. Stebbing, in commenting on this view
(‘Transactions of Devonshire Assoc. for Science,’ 1870) remarks, that had Mr.
Wallace “employed his usual ingenuity on the question of man’s hairless skin, he
might have seen the possibility of its selection through its superior beauty or the
health attaching to superior cleanliness. At any rate it is surprising that he should
picture to himself a superior intelligence plucking the hair from the backs of
savage men (to whom, according to his own account it would have been useful
and beneficial), in order that the descendants of the poor shorn wretches might
after many deaths from cold and damp in the course of many generations,” have
been forced to raise themselves in the scale of civilisation through the practice of
various arts, in the manner indicated by Mr. Wallace.
467 ‘The Variation of Animals and Plants under Domestication,’ vol. ii. 1868, p.
327.
468 ‘Investigations into Military and Anthropological Statistics of American
Soldiers,’ by B. A. Gould, 1869; p. 568:—Observations were carefully made on
the pilosity of 2129 black and coloured soldiers, whilst they were bathing; and
by looking to the published table, “it is manifest at a glance that there is but little,
if any, difference between the white and the black races in this respect.” It is,
however, certain that negroes in their native and much hotter land of Africa, have
remarkably smooth bodies. It should be particularly observed, that pure blacks
and mulattoes were included in the above enumeration; and this is an unfortunate
circumstance, as in accordance with the principle, the truth of which I have
elsewhere proved, crossed races would be eminently liable to revert to the
primordial hairy character of their early ape-like progenitors.
469 “Ueber die Richtung der Haare am Menschlichen Körper,” in Müller’s
‘Archiv für Anat. und Phys.’ 1837, s. 40.
470 Mr. Sproat (‘Scenes and Studies of Savage Life,’ 1868, p. 25) suggests, with
reference to the beardless natives of Vancouver’s Island, that the custom of
plucking out the hairs on the face, “continued from one generation to another,

Page 303

would perhaps at last produce a race distinguishable by a thin and straggling
growth of beard.” But the custom would not have arisen until the beard had
already become, from some independent cause, greatly reduced. Nor have we
any direct evidence that the continued eradication of the hair would lead to any
inherited effect. Owing to this cause of doubt, I have not hitherto alluded to the
belief held by some distinguished ethnologists, for instance M. Gosse of Geneva,
that artificial modifications of the skull tend to be inherited. I have no wish to
dispute this conclusion; and we now know from Dr. Brown-Séquard’s
remarkable observations, especially those recently communicated (1870) to the
British Association, that with guinea-pigs the effects of operations are inherited.
471 ‘Ueber die Richtung,’ ibid. s. 40.
472 On the “Limits of Natural Selection,” in the ‘North American Review,’ Oct.
1870, p. 295.
473 The Rev. J. A. Picton gives a discussion to this effect in his ‘New Theories
and the Old Faith,’ 1870.

INDEX.

A.
Abbott, C., on the battles of seals, ii. 240.
Abductor of the fifth metatarsal, presence of, in man, i. 128.
Abercrombie, Dr., on disease of the brain affecting speech, i. 58.
Abipones, marriage-customs of the, ii. 373.
Abou-Simbel, caves of, i. 217.
Abortion, prevalence of the practice of, i. 134.
Abstraction, i. 62.
Acalles, stridulation of, i. 384.
Acanthodactylus capensis, sexual differences of colour in, ii. 36.
Accentor modularis, ii. 198.
Acclimatisation, difference of, in different races of men, i. 216.
Achetidæ, stridulation of the, i. 352, 353, 355;
rudimentary stridulating organs in female, i. 359.
Acilius sulcatus, elytra of the female, i. 343.
Acomus, development of spurs in the female of, ii. 162.
Acridiidæ, stridulation of the, i. 352, 356;
rudimentary stridulating organs in female, i. 359.
Acting, i. 232.
Actiniæ, bright colours of, i. 322.
Admiral butterfly, i. 392.
Adoption of the young of other animals by female monkeys, i. 41.
Advancement in the organic scale, Von Baer’s definition of, i. 211.
Aeby, on the difference between the skulls of man and the quadrumana, i. 190.
Æsthetic faculty, not highly developed in savages, i. 64.
Affection, maternal, i. 40;
manifestation of, by animals, i. 40;
parental and filial, partly the result of natural selection, i. 81;

Page 304

shown by birds in confinement, for certain persons, ii. 110;
mutual, of birds, ii. 108.
Africa, probably the birthplace of man, i. 199;
South, crossed population of, i. 225;
South, retention of colour by the Dutch in, i. 242;
South, proportion of the sexes in the butterflies of, i. 310;
tattooing practised in, ii. 339;
Northern, coiffure of natives of, ii. 340.
Agassiz, L., on conscience in dogs, i. 78;
on the coincidence of the races of man with zoological provinces, i. 218;
on the number of species of man, i. 226;
on the courtship of the land-snails, i. 324;
on the brightness of the colours of male fishes during the breeding season, ii. 13;
on the frontal protuberance of the males of Geophagus and Cichla, ii. 13, 20;
on the slight sexual differences of the South Americans, ii. 323;
on the tattooing of the Amazonian Indians, ii. 342.
Age, in relation to the transmission of characters in birds, ii. 183;
variation in accordance with, in birds, ii. 213.
Agelæus phœniceus, ii. 116.
Ageronia feronia, noise produced by, i. 387.
Agrion, dimorphism in, i. 363.
Agrion Ramburii, sexes of, i. 362.
Agrionidæ, difference in the sexes of, i. 362.
Agrotis exclamationis, i. 369.
Ague, tertian, dog suffering from, i. 13.
Aïthurus polytmus, young of, ii. 220.
Ainos, hairiness of the, ii. 321.
Alca torda, young of, ii. 217. Alces palmata, ii. 259.
Alder and Hancock, MM., on the nudibranch mollusca, i. 326.
Algen, Mr., on the stridulation of Scolytus, i. 379.
Allen, J. A., on the relative size of the sexes of Callorhinus ursinus, ii. 260;
on the mane of Otaria jubata, ii. 267;
on the pairing of seals, ii. 279;
on sexual differences in the colour of bats, ii. 286.
Allen, S., on the habits of Hoplopterus, ii. 48;
on the plumes of herons, ii. 82;
on the vernal moult of Herodias bubulcus, ii. 84.
Alligator, courtship of the male, i. 272, ii. 29;
roaring of the male, i. 331.
Amadavat, pugnacity of male, ii. 49.
Amadina Lathami, display of plumage by the male, ii. 95.
Amadina castanotis, display of plumage by the male, ii. 95.
Amazons, butterflies of the, i. 309;
fishes of the, ii. 17.
America, variation in the skulls of aborigines of, i. 108;
wide range of aborigines of, i. 218;
lice of the natives of, i. 220;
general beardlessness of the natives of, ii. 322.
America, North, butterflies of, i. 309;
Indians of, women a cause of strife among the, ii. 324;

Page 305

Indians of, their notions of female beauty, ii. 344, 347.
America, South, character of the natives of, i. 216;
population of parts of, i. 225;
piles of stones in, i. 233;
extinction of the fossil horse of, i. 239;
desert-birds of, ii. 224;
slight sexual difference of the aborigines of, ii. 323;
prevalence of infanticide in, ii. 361.
American languages, often highly artificial, i. 112.
Americans, wide geographical range of, i. 112;
and negroes, difference of, i. 247;
aversion of, to hair on the face, ii. 348;
native, variability of, i. 226.
Ammophila, on the jaws of, i. 342.
Ammotragus tragelaphus, hairy forelegs of, ii. 282, 285.
Amphibia, affinity of, to the ganoid fishes, i. 204;
vocal organs of the, ii. 331.
Amphibians, i. 213, ii. 24;
breeding whilst immature, ii. 215.
Amphioxus, i. 204.
Amphipoda, males sexually mature while young, ii. 215.
Amunoph III., negro character of features of, i. 217.
Anal appendages of insects, i. 342.
Analogous variation in the plumage of birds, ii. 74.
Anas, ii. 180.
Anas acuta, male plumage of, ii. 84.
Anas boschas, male plumage of, ii. 84.
Anas histrionica, ii. 214.
Anastomus oscitans, sexes and young of, ii. 217;
white nuptial plumage of, ii. 228.
Anatidæ, voices of, ii. 60.
Anax junius, difference in the sexes of, i. 362.
Anglo-Saxons, estimation of the beard among the, ii. 349.
Animals, cruelty of savages to, i. 94;
domesticated, more fertile than wild, i. 132;
characters common to man and, i. 185;
domestic, change of breeds of, ii. 369.
Annelida, i. 327.
Annulosa, i. 327.
Anolis cristatellus, male, crest of, ii. 32;
pugnacity of the male, ii. 32;
throat-pouch of, ii. 33.
Anobium tessellatum, sounds produced by, i. 384.
Anser canadensis, ii. 116.
Anser cygnoides, ii. 114;
knob at the base of the beak of, ii. 129.
Anser hyperboreus, whiteness of, ii. 228.
Antelope, prong-horned, horns of, i. 289.
Antelopes, generally polygamous, i. 267;
horns of, i. 289, ii. 245;

Page 306

canine teeth of some male, ii. 241;
use of horns of, ii. 251;
dorsal crests in, ii. 282;
dewlaps of, ii. 284;
winter change of two species of, ii. 299;
peculiar markings of, ii. 299.
Antennæ, furnished with cushions in the male of Penthe, i. 343.
Anthidium manicatum, large male of, i. 347.
Anthocharis cardamines, i. 388, 393, 394;
sexual difference of colour in, i. 409.
Anthocharis genutia, i. 393.
Anthocharis sara, i. 393.
Anthophora acervorum, large male of, i. 347.
Anthophora retusa, difference of the sexes in, i. 366.
Anthus, moulting of, ii. 83.
Anthropidæ, i. 195.
Antigua, Dr. Nicholson’s observations on yellow fever in, i. 245.
Antics of birds, ii. 68.
Antilocapra americana, horns of, i. 289, ii. 245.
Antilope bezoartica, horned females of, ii. 246, 248;
sexual difference in the colour of, ii. 288.
Antilope Dorcas and euchore, ii. 245.
Antilope euchore, horns of, ii. 251.
Antilope montana, rudimentary canines in the young male of, ii. 258.
Antilope niger, sing-sing, caama, and gorgon, sexual differences in the colours of, ii. 289.
Antilope oreas, horns of, i. 289.
Antilope saiga, polygamous habits of, i. 267.
Antilope strepsiceros, horns of, i. 289.
Antilope subgutturosa, absence of suborbital pits in, ii. 280.
Antipathy, shown by birds in confinement, to certain persons, ii. 110.
Ants, i. 186;
playing together, i. 39;
memory in, i. 45;
intercommunication of, by means of the antennæ, i. 58;
large size of the cerebral ganglia in, i. 145;
soldier-, large jaws of, i. 155;
difference of the sexes in, i. 365;
recognition of each other by, after separation, i. 365.
Ants, White, habits of, i. 364.
Anura, ii. 25.
Apatania muliebris, male unknown, i. 314.
Apathus, difference of the sexes in, i. 366.
Apatura Iris, i. 386, 388.
Apes, anthropomorphous, i. 196;
difference of the young, from the adult, i. 13;
building platforms, i. 53;
probable speedy extermination of the, i. 201;
Gratiolet on the evolution of, i. 230;
semi-erect attitude of some, i. 142;
mastoid processes of, i. 143;

Page 307

influence of the jaw-muscles on the physiognomy of, i. 144;
female, destitute of large canines, i. 156;
imitative faculties of, i. 161;
canine teeth of male, ii. 241;
females of some, less hairy beneath than the males, ii. 377.
Apes, long-armed, their mode of progression, i. 143.
Apis mellifica, large male of, i. 347.
Apollo, Greek statues of, ii. 350.
Apoplexy in Cebus Azaræ, i. 12.
Approbation, influence of the love of, i. 86, 92, 164, 165.
Appendages, anal, of insects, i. 342.
Aprosmictus scapulatus, ii. 174.
Aquatic birds, frequency of white plumage in, ii. 229.
Aquila chrysaëtos, ii. 105.
Arab women, elaborate and peculiar coiffure of, ii. 353.
Arabs, gashing of cheeks and temples among the, ii. 339.
Arachnida, i. 337.
Arakhan, artificial widening of the forehead by the natives of, ii. 351.
Arboricola, young of, ii. 190.
Archeopteryx, i. 204.
Arctiidæ,, coloration of the, i. 396.
Ardea asha, rufescens, and cærulea, change of colour in, ii. 231, 232.
Ardea cærulea, breeding in immature plumage, ii. 214.
Ardea gularis, change of plumage in, ii. 232.
Ardea herodias, love-gestures of the male, ii. 68.
Ardea ludoviciana, age of mature plumage in, ii. 213;
continued growth of crest and plumes in the male of, ii. 216.
Ardea nycticorax, cries of, ii. 51.
Ardeola, young of, ii. 190.
Ardetta, changes of plumage in, ii. 179.
Argenteuil, i. 29.
Argus pheasant, ii. 72, 97, 181;
display of plumage by the male, ii. 91;
ocellated spots of the, ii. 134, 141;
gradation of characters in the, ii. 141.
Argyll, Duke of, the fashioning of implements peculiar to man, i. 52;
on the contest in man between right and wrong, i. 104;
on the physical weakness of man, i. 156;
on the primitive civilisation of man, i. 181;
on the plumage of the male Argus pheasant, ii. 91;
on Urosticte Benjamini, ii. 151;
on the nests of birds, ii. 167;
on variety as an object in nature, ii. 230.
Argynnis aglaia, colouring of the lower surface of, i. 396.
Aricoris epitus, sexual differences in the wings of, i. 345.
Aristocracy, increased beauty of the, ii. 356.
Arms, proportions of, in soldiers and sailors, i. 116;
direction of the hair on the, i. 192.
Arms and hands, free use of, indirectly correlated with diminution of canines, i. 144.
Arrest of development, i. 121, 122.

Page 308

Arrow-heads, stone, general resemblance of, i. 233.
Arrows, use of, i. 232.
Arteries, variations in the course of the, i. 108.
Artery, effect of tying, upon the lateral channels, i. 116.
Arthropoda, i. 328.
Arts practised by savages, i. 234.
Ascension, coloured incrustation on the rocks of, i. 326.
Ascidia, affinity of the lancelet to, i. 205;
tadpole-like larvæ of, i. 205.
Ascidians, i. 324;
bright colours of some, i. 322.
Asinus, Asiatic and African species of, ii. 306.
Asinus tæniopus, ii. 306.
Ass, colour-variations of the, ii. 305.
Ateles, effects of brandy on an, i. 12;
absence of the thumb in, i. 140.
Ateles beelzebuth, ears of, i. 23.
Ateles marginatus, colour of the ruff of, ii. 291;
hair on the head of, ii. 109;
on the recognition of a dog by a turkey, ii. 110;
on the selection of a mate by female birds, ii. 307.
Ateuchus, stridulation of, i. 384.
Ateuchus cicatricosus, habits of, i. 376.
Athalia, proportions of the sexes in, i. 314.
Attention, manifestations of, in animals, i. 44.
Audouin, V., on a hymenopterous parasite with a sedentary male, i. 273.
Audubon, J. J., on the pugnacity of male birds, ii. 43, 48;
on Tetrao cupido, ii. 50;
on Ardea nycticorax, ii. 51;
on Sturnella ludoviciana, ii. 51;
on the vocal organs of Tetrao cupido, ii. 56;
on the drumming of the male Tetrao umbellus, ii. 61;
on sounds produced by the nightjar, ii. 63;
on Ardea herodias and Cathartes jota, ii. 68;
on the spring change of colour in some finches, ii. 86;
on Mimus polyglottus, ii. 116;
on the turkey, ii. 119, 121;
on variation in the male scarlet tanager, ii. 126;
on the habits of Pyranga æstiva, ii. 167;
on local differences in the nests of the same species of birds, ii. 171;
on the habits of woodpeckers, ii. 175;
on Bombycilla carolinensis, ii. 180;
on young females of Tanagra æstiva acquiring male characters, ii. 180;
on the immature plumage of thrushes, ii. 185;
on the immature plumage of birds, ii. 186 et seq.;
on birds breeding in immature plumage, ii. 214;
on the growth of the crest and plumes in the male Ardea ludoviciana, ii. 216;
on the change of colour in some species of Ardea, ii. 231;
on the speculum of Mergus cucullatus, ii. 291;
on the musk-rat, ii. 298.

Page 309

Audubon and Bachman, MM., on squirrels fighting, ii. 239;
on the Canadian lynx, ii. 267.
Austen, N. L., on Anolis cristatellus, ii. 32, 33.
Australia, half-castes killed by the natives of, i. 220;
lice of the natives of, i. 220;
not the birthplace of man, i. 199;
prevalence of female infanticide in, ii. 364.
Australia, South, variation in the skulls of aborigines of, i. 108.
Australians, colour of newborn children of, ii. 318;
height of the sexes of, ii. 320;
women a cause of war among the, ii. 323.
Axis deer, sexual difference in the colour of the, ii. 290.
Aymaras, measurements of the, i. 119;
no grey hair among the, ii. 320;
hairlessness of the face in the, ii. 322;
long hair of the, ii. 348.
Azara, on the proportion of men and women among the Guaranys, i. 302;
on Palamedea cornuta, ii. 48;
on the beards of the Guaranys, ii. 322;
on strife for women among the Guanas, ii. 324;
on infanticide, ii. 344, 364;
on the eradication of the eyebrows and eyelashes by the Indians of Paraguay, ii. 348;
on polyandry among the Guanas, ii. 366;
celibacy unknown among the savages of South America, ii. 367;
on the freedom of divorce among the Charruas, ii. 372.

B.
Babbage, C., on the greater proportion of illegitimate female births, i. 302.
Babirusa, tusks of the, ii. 264.
Baboon, employing a mat for shelter against the sun, i. 53;
manifestation of memory by a, i. 45;
protected from punishment by its companions, i. 78;
rage excited in, by reading, i. 42.
Baboon, Cape, mane of the male, ii. 267;
Hamadryas, mane of the male, ii. 267.
Baboons, effects of intoxicating liquors on, i. 12;
ears of, i. 23;
manifestation of maternal affection by, i. 41;
using stones and sticks as weapons, i. 51;
co-operation of, i. 75;
silence of, on plundering expeditions, i. 79;
diversity of the mental faculties in, i. 110;
hands of, i. 139;
habits of, i. 141;
variability of the tail in, i. 150;
apparent polygamy of, i. 266;
polygamous and social habits of, ii. 362.
Bachman, Dr., on the fertility of mulattoes, i. 221.
Baer, K. E. von, on embryonic development, i. 14.
Bagehot, W., on the social virtues among primitive men, i. 93;

Page 310

on the value of obedience, i. 162;
on human progress, i. 166;
on the persistence of savage tribes in classical times, i. 239.
Bailly, E. M., on the fighting of stags, ii. 252;
on the mode of fighting of the Italian buffalo, ii. 250.
Bain, A., on the sense of duty, i. 71;
aid springing from sympathy, i. 77;
on the basis of sympathy, i. 82;
on love of approbation, &c., i. 86;
on the idea of beauty, ii. 354.
Baird, W., on a difference in colour between the males and females of some Entozoa, i. 321.
Baker, Mr., observation on the proportion of the sexes in pheasant-chicks, i. 306.
Baker, Sir S., on the fondness of the Arabs for discordant music, ii. 67;
on sexual difference in the colours of an antelope, ii. 289;
on the elephant and rhinoceros attacking white or grey horses, ii. 295;
on the disfigurements practised by the negroes, ii. 296;
on the gashing of the cheeks and temples practised in Arab countries, ii. 339;
on the coiffure of the North Africans, ii. 340;
on the perforation of the lower lip by the women of Latooka, ii. 341;
on the distinctive characters of the coiffure of central African tribes, ii. 342;
on the coiffure of Arab women, ii. 353.
“Balz” of the Black-cock, ii. 45, 100.
Bantam, Sebright, i. 259, 294.
Banteng, horns of, ii. 247;
sexual differences in the colours of the, ii. 289.
Banyai, colour of the, ii. 346.
Barbarism, primitive, of civilised nations, i. 181.
Barbs, filamentous, of the feathers, in certain birds, ii. 74.
Barr, Mr., on sexual preference in dogs, ii. 272.
Barrington, Daines, on the language of birds, i. 55;
on the clucking of the hen, ii. 51;
on the object of the song of birds, ii. 52;
on the singing of female birds, ii. 54;
on birds acquiring the songs of other birds, ii. 55;
on the muscles of the larynx in song-birds, ii. 55;
on the want of the power of song by female birds, ii. 163.
Barrow, on the widow-bird, ii. 98.
Bartlett, A. D., on the tragopan, i. 270;
on the development of the spurs in Crossoptilon auritum, i. 290;
on the fighting of the males of Plectropterus gambensis, ii. 46;
on the knot, ii. 82;
on display in male birds, ii. 87;
on the display of plumage by the male Polyplectron, ii. 89;
on Crossoptilon auritum and Phasianus Wallichii, ii. 93;
on the habits of Lophophorus, ii. 121;
on the colour of the mouth in Buceros bicornis, ii. 129;
on the incubation of the cassowary, ii. 204;
on the Cape Buffalo, ii. 250;
on the use of the horns of antelopes, ii. 251;
on the fighting of male wart-hogs, ii. 266;

Page 311

on Ammotragus tragelaphus, ii. 282;
on the colours of Cercopithecus cephus, ii. 291;
on the colours of the faces of monkeys, ii. 310;
on the naked surfaces of monkeys, ii. 377.
Bartram, on the courtship of the male alligator, ii. 29.
Basque language, highly artificial, i. 61.
Bate, C. S., on the superior activity of male crustacea, i. 272;
on the proportions of the sexes in crabs, i. 315;
on the chelæ of crustacea, i. 330;
on the relative size of the sexes in crustacea, i. 332;
on the colours of crustacea, i. 335.
Bates, H. W., on variation in the form of the head of Amazonian Indians, i. 111;
on the proportion of the sexes among Amazonian butterflies, i. 309;
on sexual differences in the wings of butterflies, i. 345;
on the field-cricket, i. 353;
on Pyrodes pulcherrimus, i. 367;
on the horns of Lamellicorn beetles, i. 370, 371;
on the colours of Epicaliæ, &c., i. 388;
on the coloration of tropical butterflies, i. 391;
on the variability of Papilio Sesostris and Childrenæ, i. 402;
on male and female butterflies inhabiting different stations, i. 403;
on mimickry, i. 411;
on the caterpillar of a Sphinx, i. 416;
on the vocal organs of the umbrella-bird, ii. 58;
on the toucans, ii. 227;
on Brachyurus calvus, ii. 309.
Batokas, knocking out two upper incisors, ii. 340.
Batrachia, ii. 25;
eagerness of male, i. 272.
Bats, sexual differences in the colour of, ii. 286.
Battle, law of, i. 182;
among beetles, i. 375;
among birds, ii. 40;
among mammals, ii. 239 et seq.;
in man, ii. 323.
Beak, sexual difference in the forms of the, ii. 39;
in the colour of the, ii. 72.
Beaks, of birds, bright colours of, ii. 227.
Beard, development of, in man, ii. 317;
analogy of the, in man and the quadrumana, ii. 319;
variation of the development of the, in different races of men, ii. 321;
estimation of, among bearded nations, ii. 349;
probable origin of the, ii. 379.
Beards, in monkeys, i. 192;
of mammals, ii. 282.
Beautiful, taste for the, in birds, ii. 108;
in the quadrumana, ii. 296.
Beauty, sense of, in animals, i. 63;
appreciation of, by birds, ii. 111;
influence of, ii. 338, 343;

Page 312

variability of the standard of, ii. 370.
Beavan, Lieut., on the development of the horns in Cervus Eldi, i. 288.
Beaver, instinct and intelligence of the, i. 37, 38;
voice of the, ii. 277;
castoreum of the, ii. 279.
Beavers, battles of male, ii. 239.
Bechstein, on female birds choosing the best singers among the males, ii. 52;
on rivalry in song-birds, ii. 53;
on the singing of female birds, ii. 54;
on birds acquiring the songs of other birds, ii. 55;
on pairing the canary and siskin, ii. 115;
on a sub-variety of the monk pigeon, ii. 132;
on spurred hens, ii. 162.
Beddoe, Dr., on causes of difference in stature, i. 115.
Bee-eater, ii. 56.
Bees, i. 73;
destruction of drones and queens by, i. 82;
pollen-baskets and stings of, i. 155;
female, secondary sexual characters of, i. 254;
difference of the sexes in, i. 365.
Beetle, luminous larva of a, i. 345.
Beetles, i. 366;
size of the cerebral ganglia in, i. 145;
dilatation of the fore tarsi in male, i. 343;
blind, i. 367;
stridulation of, i. 378.
Belgium, ancient inhabitants of, i. 237.
Bell, Sir C, on emotional muscles in man, i. 5;
“snarling muscles,” i. 127;
on the hand, i. 141.
Bell, T., on the numerical proportion of the sexes in moles, i. 305;
on the newts, ii. 24;
on the croaking of the frog, ii. 27;
on the difference in the coloration of the sexes in Zootoca vivipara, ii. 36;
on moles fighting, ii. 239.
Bell-bird, sexual difference in the colour of the, ii. 79.
Bell-birds, colours of, ii. 228.
Benevolence, manifested by birds, ii. 109.
Bennett, A. W., on the habits of Dromœus irroratus, ii. 205.
Bennett, Dr., on birds of paradise, ii. 89.
Bernicla antarctica, colours of, ii. 228.
Bernicle gander pairing with a Canada goose, ii. 114.
Bettoni, E., on local differences in the nests of Italian birds, ii. 171.
Bhoteas, colour of the beard in, ii. 319.
Bhringa, disciform tail-feathers of, ii. 83.
Bibio, sexual differences in the genus, i. 349.
Bichat, on beauty, ii. 354.
Bile, coloured, in many animals, i. 323.
Bimana, i. 190.
Birds, imitations of the songs of other birds by, i. 44;

Page 313

dreaming, i. 46;
language of, i. 55;
sense of beauty in, i. 63;
pleasure of, in incubation, i. 79;
male, incubation by, i. 210;
and reptiles, alliance of, i. 213;
sexual differences in the beak of some, i. 255;
migratory, arrival of the male before the female, i. 259;
apparent relation between polygamy and marked sexual differences in, i. 270;
monogamous, becoming polygamous under domestication, i. 270;
eagerness of male in pursuit of the female, i. 272;
wild, numerical proportion of the sexes in, i. 306;
secondary sexual characters of, ii. 38;
difference of size in the sexes of, ii. 43;
fights of male, witnessed by females, ii. 49;
display of male, to captivate the females, ii. 50;
close attention of, to the songs of others, ii. 52;
acquiring the song of their foster-parents, ii. 55;
brilliant, rarely good songsters, ii. 56;
love-antics and dances of, ii. 68;
coloration of, ii. 74 et seqq.;
moulting of, ii. 80 et seqq.;
unpaired, ii. 103;
male, singing out of season, ii. 106;
mutual affection of, ii. 108;
in confinement, distinguish persons, ii. 110;
hybrid, production of, ii. 113;
European, number of species of, ii. 124;
variability of, ii. 124;
gradation of secondary sexual characters in, ii. 135;
obscurely coloured, building concealed nests, ii. 169;
young female, acquiring male characters, ii. 180;
breeding in immature plumage, ii. 214;
moulting of, ii. 214;
aquatic, frequency of white plumage in, ii. 229;
vocal courtship of, ii. 331;
naked skin of the head and neck in, ii. 377.
Birgus latro, habits of, i. 334.
Birkbeck, Mr. on the finding of new mates by Golden Eagles, ii. 105.
Birthplace of man, i. 199.
Births, numerical proportions of the sexes in, in animals and man, i. 263, 265;
male and female, numerical proportion of, in England, i. 300.
Bischoff, Prof., on the agreement between the brains of man and of the Orang, i. 11;
figure of the embryo of the dog, i. 15;
on the convolutions of the brain in the human fœtus, i. 16;
on the difference between the skulls of man and the quadrumana, i. 190.
Bishop, J., on the vocal organs of frogs, ii. 28;
on the vocal organs of corvine birds, ii. 55;
on the trachea of the Merganser, ii. 60.
Bison, American, mane of the male, ii. 267.

Page 314

Bitterns, dwarf, coloration of the sexes of, ii. 179.
Biziura lobata, musky odour of the male, ii. 38;
large size of male, ii. 43.
Blackbird, sexual differences in the, i. 268;
proportion of the sexes in the, i. 307;
acquisition of a song by a, ii. 55;
colour of the beak in the sexes of the, ii. 72, 227;
pairing with a thrush, ii. 113;
colours and nidification of the, ii. 170;
young of the, ii. 219;
sexual difference in coloration of the, ii. 226.
Black-buck, Indian, sexual difference in the colour of the, ii. 288.
Blackcap, arrival of the male, before the female, i. 259;
young of the, ii. 219.
Black-cock, polygamous, i. 269;
proportion of the sexes in the, i. 306;
pugnacity and love-dance of the, ii. 45;
call of the, ii. 60;
moulting of the, ii. 83;
duration of the courtship of the, ii. 100;
sexual difference in coloration of the, ii. 226;
crimson eye-cere of the, ii. 227;
and pheasant, hybrids of, ii. 113.
Black-grouse, characters of young, ii. 185, 194.
Blackwall, J., on the speaking of the magpie, i. 59;
on the desertion of their young by swallows, i. 84;
on the superior activity of male spiders, i. 272;
on the proportion of the sexes in spiders, i. 314;
on sexual variation of colour in spiders, i. 337;
on male spiders, i. 338.
Bladder-nose Seal, hood of the, ii. 278.
Blaine, on the affections of dogs, ii. 270.
Blair, Dr., on the relative liability of Europeans to yellow fever, i. 243.
Blake, C. C., on the jaw from La Naulette, i. 126.
Blakiston, Capt., on the American snipe, ii. 64;
on the dances of Tetrao phasianellus, ii. 69.
Blasius, Dr., on the species of European birds, ii. 124.
Bledius taurus, hornlike processes of male, i. 374.
Bleeding, tendency to profuse, i. 292.
Blenkiron, Mr., on sexual preference in horses, ii. 272.
Blennies, crest developed on the head of male, during the breeding season, ii. 12.
Blethisa multipunctata, stridulation of, i. 379.
Bloch, on the proportions of the sexes in Fishes, i. 308.
Blood, arterial, red colour of, i. 323.
Blood-pheasant, number of spurs in, ii. 46.
Bluebreast, red-throated, sexual differences of the, ii. 195.
Blumenbach, on Man, i. 111;
on the large size of the nasal cavities in American aborigines, i. 119;
on the position of man, i. 190;
on the number of species of man, i. 226.

Page 315

Blyth, E., observations on Indian crows, i. 77;
on the structure of the hand in species of Hylobates, i. 140;
on the ascertainment of the sex of nestling bullfinches by pulling out breast-feathers, ii. 24;
on the pugnacity of the males of Gallinula cristata, ii. 41;
on the presence of spurs in the female Euplocamus erythropthalmus, ii. 46;
on the pugnacity of the amadavat, ii. 49;
on the spoonbill, ii. 60;
on the moulting of Anthus, ii. 83;
on the moulting of bustards, plovers, and Gallus bankiva, ii. 84;
on the Indian honey-buzzard, ii. 126;
on sexual differences in the colour or the eyes of hornbills, ii. 129;
on Oriolus melanocephalus, ii. 178;
on Palæornis javanicus, ii. 179;
on the genus Ardetta, ii. 179;
on the peregrine falcon, ii. 180;
on young female birds acquiring male characters, ii. 180;
on the immature plumage of birds, ii. 185;
on representative species of birds, ii. 190;
on the young of Turnix, ii. 202;
on anomalous young of Lanius rufus and Colymbus glacialis, ii. 211;
on the sexes and young of the sparrows, ii. 212;
on dimorphism in some herons, ii. 214;
on orioles breeding in immature plumage, ii. 214;
on the sexes and young of Buphus and Anastomus, ii. 217;
on the young of the blackcap and blackbird, ii. 219;
on the young of the stonechat, ii. 220;
on the white plumage of Anastomus, ii. 229;
on the horns of Antilope bezoartica, ii. 246;
on the horns of Bovine animals, ii. 247;
on the mode of fighting of Ovis cycloceros, ii. 249;
on the voice of the Gibbons, ii. 276;
on the crest of the male wild goat, ii. 282;
on the colours of Portax picta, ii. 287;
on the colours of Antilope bezoartica, ii. 288;
on the development of the horns in the Koodoo and Eland antelopes, i. 289;
on the colour of the Axis deer, ii. 290;
on sexual difference of colour in Hylobates hoolock, ii. 291;
on the hog-deer, ii. 303;
on the beard and whiskers in a monkey becoming white with age, ii. 319.
Boar, wild, polygamous in India, i. 267;
use of the tusks by the, ii. 256;
fighting of, ii. 263.
Boitard and Corbié, MM., on the transmission of sexual peculiarities in pigeons, i. 283;
on the antipathy shown by some female pigeons to certain males, ii. 118.
Bold, Mr., on the singing of a sterile hybrid canary, ii. 53.
Bombet, on the variability of the standard of beauty in Europe, ii. 370.
Bombus, difference of the sexes in, i. 366.
Bombycidæ, coloration of, i. 394;
pairing of the, i. 401.
Bombycilla carolinensis, red appendages of, ii. 179.

Page 316

Bombyx cynthia, i. 346;
proportion of the sexes in, i. 309, 313;
pairing of, i. 401.
Bombyx mori, difference of size of the male and female cocoons of, i. 346;
pairing of, i. 401.
Bombyx Pernyi, proportion of sexes of, i. 313.
Bombyx Yamamai, i. 346;
M. Personnat on, i. 310;
proportion of sexes of, i. 313.
Bonaparte, C. L., on the call-notes of the wild turkey, ii. 60.
Bond, F., on the finding of new mates by crows, ii. 104.
Bone, implements of, skill displayed in making, i. 138.
Boner, C., on the transfer of male characters to an old female chamois, ii. 245;
on the antlers of the red deer, ii. 252;
on the habits of stags, ii. 259;
on the pairing of red deer, ii. 269.
Bones, increase of, in length and thickness, when carrying a greater weight, i. 116.
Bonnet monkey, i. 192.
Boomerang, i. 183.
Boreus hyemalis, scarcity of the male, i. 314.
Bory St. Vincent, on the number of species of man, i. 226;
on the colours of Labrus pavo, ii. 16.
Bos gaurus, horns of, ii. 247.
Bos primigenius, ii. 240.
Bos sondaicus, horns of, ii. 247;
colours of, ii. 289.
Botocudos, i. 181;
mode of life of, i. 247;
disfigurement of the ears and lower lip of the, ii. 341.
Boucher de Perthes, J. C. de, on the antiquity of man, i. 3.
Bourbon, proportion of the sexes in a species of Papilio from, i. 310.
Bourien, on the marriage-customs of the savages of the Malay Archipelago, ii. 373.
Bovidæ, dewlaps of, ii. 284.
Bower-birds, ii. 102;
habits of the, ii. 69;
ornamented playing-places of, i. 63, ii. 112.
Bows, use of, i. 232.
Brachiopoda, i. 329.
Brachycephalic structure, possible explanation of, i. 148.
Brachyscelus, second pair of antennæ in the male, i. 337.
Brachyura, i. 332.
Brachyurus calvus, scarlet face of, ii. 309.
Brain, of man, agreement of the, with that of lower animals, i. 10;
convolutions of, in the human fœtus, i. 16;
larger in some existing mammals than in their tertiary prototypes, i. 51;
relation of the development of the, to the progress of language, i. 57;
disease of the, affecting speech, i. 58;
influence of development of mental faculties upon the size of the, i. 145;
influence of the development of, on the spinal column and skull, i. 146;
difference in the convolutions of, in different races of men, i. 216.

Page 317

Brakenridge, Dr., on the influence of climate, i. 115.
Braubach, Prof., on the quasi-religious feeling of a dog towards his master, i. 68;
on the self-restraint of dogs, i. 78.
Brauer, F., on dimorphism in Neurothemis, i. 363.
Brazil, skulls found in caves of, i. 218;
population of, i. 225;
compression of the nose by the natives of, ii. 352.
Break between man and the apes, i. 200.
Bream, proportion of the sexes in the, i. 308.
Breeding, age of, in birds, ii. 214.
Breeding season, sexual characters making their appearance in the, in birds, ii. 80.
Brehm, on the effects of intoxicating liquors on monkeys, i. 12;
on the recognition of women by male Cynocephali, i. 13;
on revenge taken by monkeys, i. 40;
on manifestations of maternal affection by monkeys and baboons, i. 41;
on the instinctive dread of monkeys for serpents, i. 42;
on a baboon using a mat for shelter from the sun, i. 53;
on the use of stones as missiles by baboons, i. 51;
on the signal-cries of monkeys, i. 57;
on sentinels posted by monkeys, i. 74;
on co-operation of animals, i. 75;
on an eagle attacking a young Cercopithecus, i. 76;
on baboons in confinement protecting one of their number from punishment, i. 78;
on the habits of baboons when plundering, i. 79;
on the diversity of the mental faculties of monkeys, i. 110;
on the habits of baboons, i. 141;
on polygamy in Cynocephalus and Cebus, i. 266;
on the numerical proportion of the sexes in birds, i. 306;
on the love-dance of the Black-cock, ii. 45;
on Palamedea cornuta, ii. 48;
on the habits of the Black-grouse, ii. 49;
on sound produced by Birds of Paradise, ii. 63;
on assemblages of grouse, ii. 101;
on the finding of new mates by birds, ii. 106;
on the fighting of wild boars, ii. 263;
on the habits of Cynocephalus hamadryas, ii. 362.
Brent, Mr., on the courtship of fowls, ii. 117.
Breslau, numerical proportion of male and female births in, i. 301.
Bridgman, Laura, i. 57.
Brimstone butterfly, i. 393;
sexual difference of colour in the, i. 409.
British, ancient, tattooing practised by, ii. 339.
Broca, Prof., on the occurrence of the supra-condyloid foramen in the human humerus, i. 28;
on the capacity of Parisian skulls at different periods, i. 146;
on the influence of natural selection, i. 152;
on hybridity in man, i. 220;
on human remains from Les Eyzies, i. 237;
on the cause of the difference between Europeans and Hindoos, i. 240.
Brodie, Sir B., on the origin of the moral sense in man, i. 71.
Bronn, H. G., on the copulation of insects of distinct species, i. 342.

Page 318

Bronze period, men of, in Europe, i. 160.
Brown, R., sentinels of seals generally females, i. 74;
on the battles of seals, ii. 240;
on the narwhal, ii. 242;
on the occasional absence of the tusks in the female walrus, ii. 242;
on the bladder-nose seal, ii. 278;
on the colours of the sexes in Phoca grœnlandica, ii. 287;
on the appreciation of music by seals, ii. 333;
on plants used as love-philters, by North American women, ii. 344.
Brown-Séquard, Dr., on the inheritance of the effects of operations by guinea pigs, ii. 380.
Bruce, on the use of the elephant’s tusks, ii. 249.
Brulerie, P. de la, on the habits of Ateuchus cicatricosus, i. 376;
on the stridulation of Ateuchus, i. 384.
Brünnich, on the pied ravens of the Feroe islands, ii. 126.
Bryant, Capt., on the courtship of Callorhinus ursinus, ii. 269.
Bubas bison, thoracic projection of, i. 372.
Bucephalus capensis, difference of the sexes of, in colour, ii. 29.
Buceros, nidification and incubation of, ii. 169.
Buceros bicornis, sexual differences in the colouring of the casque, beak, and mouth in, ii. 129.
Buceros corrugatus, sexual difference in the beak of, ii. 72.
Büchner, L., on the origin of man, i. 4;
on the want of self-consciousness, &c., in low savages, i. 62;
on the use of the human foot as a prehensile organ, i. 142;
on the mode of progression of the apes, i. 142.
Buckland, F., on the numerical proportion of the sexes in rats, i. 305;
on the proportion of the sexes in the trout, i. 308;
on Chimæra monstrosa, ii. 12.
Buckland, W., on the complexity of crinoids, i. 61.
Buckler, W., proportion of sexes of Lepidoptera reared by, i. 313.
Buckinghamshire, numerical proportion of male and female births in, i. 300.
Bucorax abyssinicus, inflation of the neck-wattle of the male, during courtship, ii. 72.
Budytes Raii, i. 260.
Buffalo, Cape, ii. 250.
Buffalo, Indian, horns of the, ii. 247.
Buffalo, Italian, mode of fighting of the, ii. 250.
Buffon, on the number of species of man, i. 226.
Bugs, i. 349.
Buist, R., on the proportion of the sexes in salmon, i. 308;
on the pugnacity of the male salmon, ii. 3.
Bulbul, pugnacity of the male, ii. 41;
display of under tail-coverts by the male, ii. 96.
Bull, mode of fighting of the, ii. 250;
curled frontal hair of the, ii. 282.
Bullfinch, sexual differences in the, i. 269;
piping, ii. 52;
female, singing of the, ii. 54;
courtship of the, ii. 94;
widowed, finding a new mate, ii. 105;
attacking a reed-bunting, ii. 111;
nestling, sex ascertained by pulling out breast-feathers, ii. 214.

Page 319

Bullfinches distinguishing persons, ii. 110;
rivalry of female, ii. 121.
Bulls, two young, attacking an old one, i. 75;
wild, battles of, ii. 240.
Bull-trout, male, colouring of, during the breeding season, ii. 14.
Bunting, reed, head feathers of the male, ii. 95;
attacked by a bullfinch, ii. 111.
Buntings, characters of young, ii. 184.
Buphus coromandus, sexes and young of, ii. 217;
change of colour in, ii. 231, 232.
Burchell, Dr., on the zebra, ii. 302;
on the extravagance of a Bushwoman in adorning herself, ii. 344;
celibacy unknown among the savages of South Africa, ii. 367;
on the marriage-customs of the Bushwomen, ii. 374.
Burke, on the number of species of man, i. 226.
Burmese, colour of the beard in, ii. 319.
Burton, Capt., on negro ideas of female beauty, ii. 346;
on a universal ideal of beauty, ii. 351.
Bushmen, i. 157.
Bushwoman, extravagant ornamentation of a, ii. 344.
Bushwomen, hair of, i. 216;
marriage-customs of, ii. 374.
Busk, Prof. G., on the occurrence of the supra-condyloid foramen in the human humerus, i. 28.
Bustard, throat-pouch of the male, ii. 58;
humming noise produced by a male, ii. 65;
Indian, ear-tufts of a, ii. 73.
Bustards, occurrence of sexual differences and of polygamy among the, i. 269;
love-gestures of the male, ii. 68;
double moult in, ii. 81, 83.
Butler, A. G., on sexual differences in the wings of Aricoris epitus, i. 345;
>on the colouring of the sexes in species of Thecla, i. 389;
on the resemblance of Iphias glaucippe to a leaf, i. 394;
on the rejection of certain moths and caterpillars by lizards and frogs, i. 417.
Butterfly, noise produced by a, i. 387;
Emperor, i. 386, 388;
meadow brown, instability of the ocellated spots of, ii. 132.
Butterflies, proportion of the sexes in, i. 309;
forelegs atrophied in some male, i. 344;
sexual difference in the neuration of the wings of, i. 345;
pugnacity of male, i. 386;
protective resemblances of the lower surface of, i. 392;
display of the wings by, i. 396;
white, alighting upon bits of paper, i. 400;
attracted by a dead specimen of the same species, i. 400;
courtship of, i. 400;
male and female, inhabiting different stations, i. 403.
Buxton, C., observations on macaws, i. 76;
on an instance of benevolence in a parrot, ii. 109.
Buzzard, Indian honey-, variation in the crest of, ii. 126.

Page 320

C.
Cabbage butterflies, i. 393.
Cachalot, large head of the male, ii. 242.
Cadences, musical, perception of, by animals, ii. 333.
Cæcum, i. 27;
large, in the early progenitors of man, i. 206.
Cairina moschata, pugnacity of the male, ii. 43.
Callianassa, chelæ of, figured, i. 330.
Callionymus lyra, characters of the male, ii. 7.
Callorhinus ursinus, relative size of the sexes of, ii. 260;
courtship of, ii. 269.
Calotes nigrilabris, sexual difference in the colour of, ii. 36.
Cambridge, O. Pickard, on the sexes of spiders, i. 315.
Camel, canine teeth of male, ii. 241, 257.
Campbell, J., on the Indian elephant, i. 267, 268;
on the proportion of male and female births in the harems of Siam, i. 303.
Campylopterus hemileucurus, i. 307.
Canaries distinguishing persons, ii. 110.
Canary, polygamy of the, i. 270;
change of plumage in, after moulting, i. 294;
female, selecting the best singing male, ii. 52;
sterile hybrid, singing of a, ii. 53;
female, singing of the, ii. 54;
selecting a greenfinch, ii. 115;
and siskin, pairing of, ii. 115.
Canestrini, G., on rudimentary characters and the origin of man, i. 4;
on rudimentary characters, i. 17;
on the movement of the ear in man, i. 20;
on the variability of the vermiform appendage in man, i. 27;
on the abnormal division of the malar bone in man, i. 124;
on abnormal conditions of the human uterus, i. 124;
on the persistence of the frontal suture in man, i. 125;
on the proportion of the sexes in silk-moths, i. 309, 311.
Canine teeth in man, i. 126;
diminution of, in man, i. 144;
diminution of, in horses, i. 144;
disappearance of, in male ruminants, i. 144;
large, in the early progenitors of man, i. 206. Canines, and horns, inverse development of, ii.
257.
Canoes, use of, i. 137, 234.
Cantharis, difference of colour in the sexes of a species of, i. 367.
Capercailzie, proportion of the sexes in the, i. 306;
pugnacity of the male, ii. 45;
pairing of the, ii. 49;
autumn meetings of the, ii. 54;
call of the, ii. 61;
duration of the courtship of, ii. 100;
behaviour of the female, ii. 121;
inconvenience of black colour to the female, ii. 154;

Page 321

sexual difference in coloration of the, ii. 226;
crimson eye-cere of the male, ii. 227;
polygamous, i. 269.
Capital, i. 169.
Capitonidæ, colours and nidification of the, ii. 171.
Capra ægagrus, ii. 249;
crest of the male, ii. 282;
sexual difference in the colour of, ii. 289.
Capreolus Sibiricus subecaudatus, ii. 298.
Caprice, common to man and animals, i. 65.
Caprimulgus, noise made by the males of some species of, with their wings, ii. 62.
Caprimulgus virginianus, pairing of, ii. 49.
Carabidæ, bright colours of, i. 367.
Carbonnier, on the natural history of the pike, i. 308;
the relative size of the sexes in fishes, ii. 7.
Carcineutes, sexual difference of colour in, ii. 173.
Carcinus mænas, i. 331, 333.
Carduelis elegans, sexual differences of the beak in, ii. 39.
Carnivora, marine, polygamous habits of, i. 268;
sexual differences in the colours of, ii. 286.
Carp, numerical proportion of the sexes in the, i. 308.
Carr, R., on the peewit, ii. 48.
Carrier pigeon, late development of the wattle in the, i. 293.
Carrion beetles, stridulation of, i. 378.
Carus, Prof. V., on the development of the horns in merino sheep, i. 289.
Cassowary, sexes and incubation of the, ii. 204.
Castoreum, ii. 279.
Casuarius galeatus, ii. 204.
Cat, convoluted body in the extremity of the tail of a, i. 30;
sick, sympathy of a dog with a, i. 77.
Cataract in Cebus Azaræ, i. 12.
Catarrh, liability of Cebus Azaræ to, i. 11.
Catarrhine monkeys, i. 196.
Caterpillars, bright colours of, i. 415.
Cathartes aura, ii. 116.
Cathartes jota, love-gestures of the male, ii. 68.
Catlin, G., on the development of the beard among North American Indians, ii. 322;
on the great length of the hair in some North American tribes, ii. 348.
Caton, J. D., on the development of the horns in Cervus virginianus and strongyloceros, i. 288;
on the presence of traces of horns in the female wapiti, ii. 245;
on the fighting of deer, ii. 252;
on the crest of the male wapiti, ii. 282;
on the colours of the Virginian deer, ii. 288;
on sexual differences of colour in the wapiti, ii. 289;
on the spots of the Virginian deer, ii. 303.
Cats, dreaming, i. 46;
tortois-eshell, i. 283, 285, 293;
enticed by valerian, ii. 281;
colours of, ii. 299.
Cattle, domestic, sexual differences of, late developed, i. 293;

Page 322

rapid increase of, in South America, i. 135;
domestic, lighter in winter in Siberia, i. 282;
horns of, i. 289, ii. 247;
numerical proportion of the sexes in, i. 305.
Caudal vertebræ, number of, in macaques and baboons, i. 150;
basal, of monkeys, imbedded in the body, i. 151.
Cebus, maternal affection in a, i. 40;
gradation of species of, i. 227.
Cebus Azaræ, liability of, to the same diseases as man, i. 11;
distinct sounds produced by, i. 53;
early maturity of the female, ii. 318.
Cebus capucinus, polygamous, i. 266;
sexual differences of colour in, ii. 290;
hair on the head of, ii. 307.
Cebus vellerosus, hair on the head of, ii. 307.
Cecidomyidæ, proportions of the sexes in, i. 314.
Celibacy, unknown among the savages of South Africa and South America, ii. 367.
Centipedes, i. 339.
Cephalopoda, absence of secondary sexual characters in, i. 325.
Cephalopterus ornatus, ii. 58, 59.
Cephalopterus penduliger, ii. 59.
Cerambyx heros, stridulant organ of, i. 380.
Ceratophora aspera, nasal appendages of, ii. 34.
Ceratophora Stoddartii, nasal horn of, ii. 34.
Cerceris, habits of, i. 364.
Cercocebus æthiops, whiskers, &c., of, ii. 308.
Cercopithecus, young, seized by an eagle and rescued by the troop, i. 76;
definition of species of, i. 227.
Cercopithecus cephus, sexual difference of colour in, ii. 291, 311.
Cercopithecus cynosurus and griseo-viridis, colour of the scrotum in, ii. 291.
Cercopithecus Diana, sexual differences of colour in, ii. 291, 311, 312.
Cercopithecus griseo-viridis, i. 75.
Cercopithecus petaurista, whiskers, &c., of, ii. 308.
Ceres, of birds, bright colours of, ii. 227.
Ceriornis Temminckii, swelling of the wattles of the male during courtship, ii. 72.
Cervulus, weapons of, ii. 257.
Cervulus moschatus, rudimentary horns of the female, ii. 245.
Cervus alces, i. 288.
Cervus campestris, odour of, ii. 279.
Cervus canadensis, traces of horns in the female, ii. 245;
attacking a man, ii. 253;
sexual difference in the colour of, ii. 289.
Cervus elaphus, battles of male, ii. 240;
horns of, with numerous points, ii. 252.
Cervus Eldi, i. 288.
Cervus mantchuricus, ii. 303.
Cervus paludosus, colours of, ii. 290.
Cervus strongyloceros, i. 288.
Cervus virginianus, i. 288;
horns of, in course of modification, ii. 255.

Page 323

Ceryle, male black-belted in some species of, ii. 173.
Cetacea, nakedness of, i. 148.
Ceylon, frequent absence of beard in the natives of, ii. 321.
Chaffinch, proportion of the sexes in the, i. 306, 307;
courtship of the, ii. 94.
Chaffinches, ii. 53;
new mates found by, ii. 105.
Chalcophaps indicus, characters of young, ii. 185.
Chalcosoma atlas, sexual differences of, i. 368.
Chamæleon, sexual differences in the genus, ii. 34.
Chamæleon bifurcus, ii. 34, 35.
Chamæleon Owenii, ii. 34, 36.
Chameleons, ii. 32.
Chamois, danger-signals of, i. 74;
transfer of male characters to an old female, ii. 245.
Chamæpetes unicolor, modified wing-feather in the male, ii. 64.
Chapuis, Dr., on the transmission of sexual peculiarities in pigeons, i. 283;
on streaked Belgian pigeons, i. 294, ii. 157.
Char, male, colouring of, during the breeding season, ii. 14.
Characters, male, developed in females, i. 280;
natural, artificial exaggeration of, by man, ii. 351;
secondary sexual, transmitted through both sexes, i. 279.
Charadrius hiaticula and pluvialis, sexes and young of, ii. 216.
Chardin on the Persians, ii. 356.
Charms, worn by women, ii. 344.
Charruas, freedom of divorce among the, ii. 372.
Chasmorhynchus, difference of colour in the sexes of, ii. 79;
colours of, ii. 228.
Chastity, early estimation of, i. 96.
Chatterers, sexual differences in, i. 269.
Cheiroptera, absence of secondary sexual characters in, i. 268.
Chelæ of crustacea, i. 330, 336.
Chelonia, sexual differences in, ii. 28.
Chenalopex ægyptiacus, wing-knobs of, ii. 46.
Chera progne, ii. 84, 120.
Chest, proportions of, in soldiers and sailors, i. 117;
large, of the Quechua and Aymara Indians, i. 119.
Chevrotains, canine teeth of, ii. 257.
Chiasognathus, stridulation of, i. 384.
Chiasognathus Grantii, mandibles of, i. 377.
Children, legitimate and illegitimate, proportion of the sexes in, i. 302.
Chiloe, lice of the natives of, i. 220;
population of, i. 225.
Chimæra monstrosa, bony process on the head of the male, ii. 12.
Chimæroid fishes, prehensile organs of male, ii. 1.
Chimpanzee, ii. 323;
ears of the, i. 21;
representatives of the eyebrows in the, i. 25;
platforms built by the, i. 36;
cracking nuts with a stone, i. 51;

Page 324

hands of the, i. 139;
absence of mastoid processes in the, i. 143;
direction of the hair on the arms of the, i. 192;
supposed evolution of the, i. 230;
polygamous and social habits of the, ii. 362.
China, North, idea of female beauty in, ii. 344.
China, Southern, inhabitants of, i. 246.
Chinese, use of flint tools by the, i. 183;
difficulty of distinguishing the races of the, i. 215;
colour of the beard in, ii. 319;
general beardlessness of the, ii. 321;
opinions of the, on the appearance of Europeans and Cingalese, ii. 345, 347;
compression of the feet of, ii. 352.
Chinsurdi, his opinion of beards, ii. 341, 349.
Chlamydera maculata, ii. 70.
Chloëon, pedunculated eyes of the male of, i. 341.
Chloephaga, coloration of the sexes in, ii. 178.
Chlorocœlus Tanana (figured), i. 355.
Chorda Dorsalis, i. 207.
Chough, red beak of the, ii. 227.
Chromidæ, frontal protuberance in male, ii. 13;
sexual differences in colour of, ii. 20.
Chrysemys picta, long claws of the male, ii. 28.
Chrysococcyx, characters of young of, ii. 185.
Chrysomela cerealis, bright colours of, i. 367.
Chrysomelidæ, stridulation of, i. 379.
Cicada pruinosa, i. 351.
Cicada septendecim, i. 351.
Cicadæ, songs of the, i. 350;
rudimentary sound-organs in females of, i. 359.
Cicatrix of a burn, causing modification of the facial bones, i. 147.
Cichla, frontal protuberance of male, ii. 13.
Cimetière du Sud, Paris, i. 28.
Cincloramphus cruralis, large size of male, ii. 43.
Cinclus aquaticus, ii. 170.
Cingalese, Chinese opinion of the appearance of the, ii. 345.
Cirripedes, complemental males of, i. 255.
Civilisation, effects of, upon natural selection, i. 170;
influence of, in the competition of nations, i. 239.
Clanging of Geese, &c., ii. 51.
Claparède, E., on natural selection applied to man, i. 137.
Clarke, on the marriage-customs of the Kalmucks, ii. 373.
Classification, i. 188.
Claus, C., on the sexes of Saphirina, i. 336.
Cleft-palate, inherited, i. 121.
Climacteris erythrops, sexes of, ii. 206.
Climate, i. 115;
cool, favourable to human progress, i. 167;
power of supporting extremes of, by man, i. 237;
want of connexion of, with colour, i. 241.

Page 325

Cloaca, existence of a, in the early progenitors of man, i. 207.
Cloacal passage existing in the human embryo, i. 16.
Club, origin of the, i. 234.
Clucking of fowls, ii. 51.
Clythra 4-punctata, stridulation of, i. 379.
Cobra, ingenuity of a, ii. 31.
Coccus, i. 186.
Coccyx, i. 29, 30;
in the human embryo, i. 16;
convoluted body at the extremity of the, i. 30;
imbedded in the body, i. 151.
Cochin-China, notions of beauty of the inhabitants of, ii. 345, 347.
Cock, game, killing a kite, ii. 44;
blind, fed by its companions, i. 77;
comb and wattles of the, ii. 98;
preference shown by the, for young hens, ii. 121;
game, transparent zone in the hackles of a, ii. 136.
Cock of the rock, ii. 100.
Cockatoos, ii. 226, 228, 230;
nestling, ii. 109;
black, immature plumage of, ii. 188.
Cœlenterata, absence of secondary sexual characters in, i. 321.
Coffee, fondness of monkeys for, i. 12.
Cold, supposed effects of, i. 116;
power of supporting, by man, i. 237.
Coleoptera, i. 366;
stridulant organs of, discussed, i. 381.
Collingwood, C., on the pugnacity of the butterflies of Borneo, i. 386;
on butterflies being attracted by a dead specimen of the same species, i. 400.
Colombia, flattened heads of savages of, ii. 340.
Colonists, success of the English as, i. 179.
Coloration, protective, in birds, ii. 223.
Colour, supposed to be dependent on light and heat, i. 115;
correlation of, with immunity from certain poisons and parasites, i. 242;
purpose of, in lepidoptera, i. 399;
relation of, to sexual functions, in fishes, ii. 14;
difference of, in the sexes of snakes, ii. 29;
sexual differences of, in lizards, ii. 36;
influence of, in the pairing of birds of different species, ii. 115;
relation of, to nidification, ii. 167, 172;
sexual differences of, in mammals, ii. 286, 294;
recognition of, by quadrupeds, ii. 295;
of children, in different races of man, ii. 318;
of the skin in man, ii. 381.
Colours, admired alike by man and animals, i. 64;
bright, due to sexual selection, i. 322;
bright, among the lower animals, i. 322, 323;
bright, protective to butterflies and moths, i. 395;
bright, in male fishes, ii. 7, 13;
transmission of, in birds, ii. 159.

Page 326

Colquhoun, example of reasoning in a retriever, i. 48.
Columba passerina, young of, ii. 188.
Colymbus glacialis, anomalous young of, ii. 211.
Comb, development of, in fowls, i. 295.
Combs and wattles in male birds, ii. 98.
Community, preservation of variations useful to the, by natural selection, i. 155.
Compositæ, gradation of species among the, i. 227.
Comte, C., on the expression of the ideal of beauty by sculpture, ii. 380.
Conditions of life, action of changed, upon man, i. 113;
influence of, on plumage of birds, ii. 196.
Condor, eyes and comb of the, ii. 129.
Conjugations, origin of, i. 61.
Conscience, i. 91, 104;
absence of, in some criminals, i. 92.
Constitution, difference of, in different races of men, i. 216.
Consumption, liability of Cebus Azaræ to, i. 12;
connexion between complexion and, i. 244.
Convergence, i. 230.
Cooing of pigeons and doves, ii. 60.
Cook, Capt., on the nobles of the Sandwich Islands, ii. 356.
Cope, E. D., on the dinosauria, i. 204;
on the origin of genera, ii. 215.
Cophotis ceylanica, sexual differences of, ii. 32, 36.
Copris, i. 370.
Copris Isidis, sexual differences of, i. 369.
Copris lunaris, stridulation of, i. 380.
Corals, bright colours of, i. 322.
Coral-snakes, ii. 31.
Cordylus, sexual difference of colour in a species of, ii. 36.
Corfu, habits of the chaffinch in, i. 307.
Cornelius, on the proportions of the sexes in Lucanus Cervus, i. 313.
Corpora Wolffiana, i. 207;
agreement of, with the kidneys of fishes, i. 16.
Correlated variation, i. 130.
Correlation, influence of, in the production of races, i. 247.
Corse, on the mode of fighting of the elephant, ii. 257.
Corvus corone, ii. 104.
Corvus graculus, red beak of, ii. 227.
Corvus pica, nuptial assembly of, ii. 102.
Corydalis cornutus, large jaws of the male, i. 342.
Cosmetornis, ii. 181.
Cosmetornis vexillarius, elongation of wing-feathers in, ii. 73, 97.
Cotingidæ, sexual differences in, i. 269;
coloration of the sexes of, ii. 177;
resemblance of the females of distinct species of, ii. 192.
Cottus scorpius, sexual differences in, ii. 9.
Counting, origin of, i. 181;
limited power of, in primeval man, i. 234.
Courage, variability of, in the same species, i. 40;
universal high appreciation of, i. 95;

Page 327

importance of, i. 162;
a characteristic of men, ii. 328.
Courtship, greater eagerness of males in, i. 272;
of fishes, ii. 2;
of birds, ii. 50, 100.
Cow, winter change of the, ii. 299.
Crab, devil, i. 332.
Crab, shore, habits of, i. 331.
Crabro cribrarius, dilated tibiæ of the male, i. 343.
Crabs, proportions of the sexes in, i. 315.
Cranz, on the inheritance of dexterity in seal-catching, i. 117.
Crawfurd, on the number of species of man, i. 226.
Crenilabrus massa and C. melops, nests built by, ii. 19.
Crest, origin of, in Polish fowls, i. 284.
Crests, of birds, difference of, in the sexes, ii. 189;
dorsal hairy, of mammals, ii. 282.
Cricket, field-, stridulation of the, i. 353;
pugnacity of male, i. 360.
Cricket, house-, stridulation of the, i. 352, 354.
Crickets, sexual differences in, i. 361.
Crioceridæ, stridulation of the, i. 379.
Crinoids, complexity of, i. 61.
Croaking of frogs, ii. 27.
Crocodiles, musky odour of, during the breeding season, ii. 29.
Crocodilia, ii. 28.
Crossbills, characters of young, ii. 184.
Crosses in man, i. 225.
Crossing of races, effects of the, i. 241.
Crossoptilon auritum, ii. 93, 166, 196;
adornment of both sexes of, i. 290;
sexes alike in, ii. 178.
Crotch, G. R., on the stridulation of beetles, i. 379, 382;
on the stridulation of Heliopathes, i. 383;
on the stridulation of Acalles, i. 384.
Crow Indians, long hair of the, ii. 348.
Crow, young of the, ii. 209.
Crows, ii. 226;
vocal organs of the, ii. 55;
living in triplets, ii. 106.
Crows, carrion, new mates found by, ii. 104.
Crows, Indian, feeding their blind companions, i. 77.
Cruelty of savages to animals, i. 94.
Crustacea, amphipod, males sexually mature while young, ii. 215;
parasitic, loss of limbs by female, i. 255;
prehensile feet and antennæ of, i. 256;
male, more active than female, i. 272;
parthenogenesis in, i. 315;
secondary sexual characters of, i. 328;
auditory hairs of, ii. 333.
Crystal worn in the lower lip by some Central African women, ii. 341.

Page 328

Cuckoo fowls, i. 294.
Culicidæ, i. 254, 349.
Cullen, Dr., on the throat-pouch of the male bustard, ii. 58.
Cultivation of plants, probable origin of, i. 167.
Cupples, Mr., on the numerical proportion of the sexes in dogs, sheep, and cattle, i. 304, 305;
on the Scotch deerhound, ii. 261;
on sexual preference in dogs, ii. 271, 272.
Curculionidæ, sexual difference in length of snout in some, i. 255;
hornlike processes in male, i. 374;
musical, i. 378, 379.
Curiosity, manifestations of, by animals, i. 42.
Curlews, double moult in, ii. 80.
Cursores, comparative absence of sexual differences among the, i. 269.
Curtis, J., on the proportion of the sexes in Athalia, i. 314.
Cuvier, F., on the recognition of women by male quadrumana, i. 13.
Cuvier, G., views of, as to the position of man, i. 190;
on instinct and intelligence, i. 37;
on the number of caudal vertebræ in the mandrill, i. 150;
on the position of the seals, i. 190;
on Hectocotyle, i. 325.
Cyanecula suecica, sexual differences of, ii. 195.
Cyanalcyon, sexual difference in colours of, ii. 173;
immature plumage of, ii. 188.
Cychrus, sounds produced by, i. 382.
Cycnia mendica, sexual difference of colour in, i. 398.
Cygnus ferus, trachea of, ii. 59.
Cygnus olor, white young of, ii. 211.
Cyllo Leda, instability of the ocellated spots of, ii. 133.
Cynanthus, variation in the genus, ii. 125.
Cynipidæ, proportions of the sexes in, i. 314.
Cynocephalus, difference of the young, from the adult, i. 13;
male, recognition of women by, i. 13;
polygamous habits of species of, i. 266.
Cynocephalus chacma, i. 41.
Cynocephalus gelada, i. 51.
Cynocephalus hamadryas, i. 51;
sexual difference of colour in, ii. 291.
Cynocephalus leucophœus, colours of the sexes of, ii. 292.
Cynocephalus mormon, colours of the male, ii. 292, 296, 310.
Cynocephalus porcarius, mane of the male, ii. 267.
Cypridina, proportions of the sexes in, i. 315.
Cyprinidæ, proportion of the sexes in the, i. 308.
Cyprinidæ, Indian, ii. 17.
Cyprinodontidæ, sexual differences in the, ii. 7, 9.
Cyprinus auratus, ii. 16.
Cyprinus phoxinus, spawning of, ii. 15.
Cypris, relations of the sexes in, i. 315.
Cystophora cristata, hood of, ii. 278.

D.

Page 329

Dacelo, sexual difference of colour in, ii. 174.
Dacelo Gaudichaudi, young male of, ii. 188.
Dal-ripa, a kind of ptarmigan, i. 306.
Damalis albifrons, peculiar markings of, ii. 301.
Damalis pygarga, peculiar markings of, ii. 300.
Dampness of climate, supposed influence of, on the colour of the skin, i. 116, 242.
Danaidæ, i. 387.
Dances of birds, ii. 68.
Dancing, i. 232.
Daniell, Dr., his experience of residence in West Africa, i. 245.
Darfur, protuberances artificially produced in, ii. 339.
Darwin, F., on the stridulation of Dermestes murinus, i. 379.
Dasychira pudibunda, sexual difference of colour in, i. 398.
Davis, A. H., on the pugnacity of the male stag-beetle, i. 375.
Davis, J. B., on the capacity of the skull in various races of men, i. 146;
on the beards of the Polynesians, ii. 322.
Death-rate higher in towns than in rural districts, i. 175.
Death-tick, i. 384.
De Candolle, Alph., on a case of inherited power of moving the scalp, i. 20.
Declensions, origin of, i. 61.
Decoration in birds, ii. 71.
Decticus, i. 355.
Deer, spots of young, ii. 184, 303;
horns of, ii. 243, 248;
use of horns of, ii. 252, 263;
size of the horns of, ii. 259;
female, pairing with one male, whilst others are fighting for her, ii. 269;
male, attracted by the voice of the female, ii. 276;
male, odour emitted by, ii. 279;
development of the horns in, i. 288;
horns of a, in course of modification, ii. 255.
Deer, Axis, sexual, difference in the colour of the, ii. 290.
Deer, fallow, different coloured herds of, ii. 295.
Deer, Mantchurian, ii. 303.
Deer, Virginian, ii. 303;
colour of the, not affected by castration, ii. 288;
colours of, ii. 289.
Deerhound, Scotch, greater size of the male, i. 293, ii. 260.
Defensive organs of mammals, ii. 263.
De Geer, C., on a female spider destroying a male, i. 339.
Dekay, Dr., on the bladder-nose seal, ii. 278.
Demerara, yellow fever in, i. 243.
Dendrocygna, ii. 185.
Dendrophila frontalis, young of, ii. 220.
Denny, H., on the lice of domestic animals, i. 219.
Dermestes murinus, stridulation of, i. 379.
Descent traced through the mother alone, ii. 359.
Deserts, protective colouring of animals inhabiting, ii. 224.
Desmarest, on the absence of suborbital pits in Antilope subgutturosa, ii. 280;
on the whiskers of Macacus, ii. 283;

Page 330

on the colour of the opossum, ii. 286;
on the colours of the sexes of Mus minutus, ii. 286;
on the colouring of the ocelot, ii. 287;
on the colours of seals, ii. 287;
on Antilope caama, ii. 289;
on the colours of goats, ii. 290;
on sexual difference of colour in Ateles marginatus, ii. 291;
on the mandrill, ii. 293;
on Macacus cynomolgus, ii. 318.
Desmoulins, on the number of species of man, i. 226;
on the musk-deer, ii. 281.
Desor, on the imitation of man by monkeys, i. 44.
Despine, P., on criminals destitute of conscience, i. 92.
Development, embryonic, of man, i. 14, 16;
correlated, ii. 130.
Devil, not believed in by the Fuegians, i. 67.
Devil-crab, i. 332.
Devonian, fossil insect from the, i. 360.
Dewlaps, of cattle and antelopes, ii. 284.
Diadema, sexual differences of colouring in the species of, i. 388.
Diadema anomala, mimickry by the female of, i. 413.
Diadema bolina, i. 413.
Diamond-beetles, bright colours of, i. 367.
Diastema, occurrence of, in man, i. 126.
Diastylidæ, proportion of the sexes in, i. 315.
Diodorus, on the absence of beard in the natives of Ceylon, ii. 321.
Dicrurus, racket-shaped feathers in, ii. 73;
nidification of, ii. 167.
Dicrurus macrocercus, change of plumage in, ii. 179.
Didelphis opossum, sexual difference in the colour of, ii. 286.
Differences, comparative, between different species of birds of the same sex, ii. 192.
Digits, supernumerary, more frequent in men than in women, i. 276;
supernumerary, inheritance of, i. 285;
supernumerary, early development of, i. 292.
Dimorphism in females of water-beetles, i. 343;
in Neurothemis and Agrion, i. 363.
Dipelicus Cantori, sexual differences of, i. 369.
Diplopoda, prehensile limbs of the male, i. 340.
Dipsas cynodon, sexual difference in the colour of, ii. 29.
Diptera, i. 348.
Disease, generated by the contact of distinct peoples, i. 239.
Diseases common to man and the lower animals, i. 11;
difference of liability to, in different races of men, i. 216;
new, effects of, upon savages, i. 238;
sexually limited, i. 292.
Display, coloration of Lepidoptera for, i. 395;
of plumage by male birds, ii. 86, 96.
Distribution, wide, of man, i. 137;
geographical, as evidence of specific distinctness in man, i. 218.
Disuse, effects of, in producing rudimentary organs, i. 18;

Page 331

and use of parts, effects of, i. 116;
of parts, influence of, on the races of men, i. 247.
Divorce, freedom of, among the Charruas, ii. 372.
Dixon, E. S., on the habits of the guinea-fowl, i. 270;
on the pairing of different species of geese, ii. 114;
on the courtship of peafowl, ii. 121.
Dobrizhoffer, on the marriage-customs of the Abipones, ii. 374.
Dogs, suffering from Tertian ague, i. 13;
memory of, i. 45;
domestic, progress of, in moral qualities, i. 50;
distinct tones uttered by, i. 54;
parallelism between his affection for his master and religious feeling, i. 68;
sociability of the, i. 74;
sympathy of, with a sick cat, i. 77;
sympathy of, with his master, i. 77;
possible use of the hair on the forelegs of the, i. 193;
races of the, i. 229;
diverging when drawing sledges over thin ice, i. 40;
dreaming, i. 46, 158;
exercise of reasoning faculties by, i. 48;
their possession of conscience, i. 78;
numerical proportion of male and female births in, i. 304;
sexual affection between individuals of, ii. 270;
howling at certain notes, ii. 333;
rolling in carrion, ii. 281.
Dolichocephalic structure, possible cause of, i. 148.
Dolphins, nakedness of, i. 148.
Domestic animals, races of, i. 229;
change of breeds of, ii. 369.
Domestication, influence of, in removing the sterility of hybrids, i. 222.
D’Orbigny, A., on the influence of dampness and dryness on the colour of the skin, i. 242;
on the Yura-caras ii. 347.
Dotterel, ii. 203.
Doubleday, E., on sexual differences in the wings of butterflies, i. 345.
Doubleday, H., on the proportion of the sexes in the smaller moths, i. 311;
on the attraction of the males of Lasiocampa quercus and Saturnia carpini by the female, i.
312;
on the proportion of the sexes in the Lepidoptera, i. 312;
on the ticking of Anobium tessellatum, i. 385;
on the structure of Ageronia feronia, i. 387;
on white butterflies alighting upon paper, i. 400.
Douglas, J. W., on the sexual differences of the Hemiptera, i. 349;
on the colours of British Homoptera, i. 352.
Down, of birds, ii. 80.
Draco, gular appendages of, ii. 33.
Dragonet, Gemmeous, ii. 7.
Dragon-flies, caudal appendages of male, i. 344;
relative size of the sexes of, i. 347;
difference in the sexes of, i. 361;
want of pugnacity by the male, i. 364.

Page 332

Drake, breeding plumage of the, ii. 84.
Dreams, i. 46;
a possible source of the belief in spiritual agencies, i. 66.
Drill, sexual difference of colour in the, ii. 291.
Dromœus irroratus, ii. 204.
Dromolæa, Saharan species of, ii. 172.
Drongo shrike, ii. 179.
Drongos, racket-shaped feathers in the tails of, ii. 73, 83.
Dryness, of climate, supposed influence of, on the colour of the skin, i. 242.
Dryopithecus, i. 199.
Duck, harlequin, age of mature plumage in the, ii. 213;
breeding in immature plumage, ii. 214.
Duck, long-tailed, preference of male, for certain females, ii. 122.
Duck, pintail, pairing with a wigeon, ii. 114.
Duck, voice of the, ii. 60;
pairing with a shield-drake, ii. 114;
immature plumage of the, ii. 188.
Duck, wild, sexual differences in the, i. 268;
speculum and male characters of, i. 291;
pairing with a pintail drake, ii. 115.
Ducks, dogs and cats recognised by, ii. 110;
wild, becoming polygamous under partial domestication, i. 270.
Dugong, tusks of, ii. 242;
nakedness of, i. 148.
Dujardin, on the relative size of the cerebral ganglia in insects, i. 145.
Duncan, Dr., on the fertility of early marriages, i. 174.
Dupont, M., on the occurrence of the supra-condyloid foramen in the humerus of man, i. 29.
Durand, J. P., on causes of variation, i. 113.
Dureau de la Malle, on the songs of birds, i. 55;
on the acquisition of an air by blackbirds, ii. 55.
Dutch, retention of their colour by the, in South Africa, i. 242.
Duty, sense of, i. 70.
Duvaucel, female Hylobates washing her young, i. 40.
Dyaks, pride of, in mere homicide, i. 94.
Dynastes, large size of males of, i. 347.
Dynastini, stridulation of, i. 381.
Dytiscus, dimorphism of females of, i. 343;
grooved elytra of the female, i. 343.

E.
Eagle, young Cercopithecus rescued from, by the troop, i. 75.
Eagle, white-headed, breeding in immature plumage, ii. 214.
Eagles, golden, new mates found by, ii. 105.
Ear, motion of the, i. 20;
external shell of the, useless in man, i. 21;
rudimentary point of the, in man, i. 22.
Ears, piercing and ornamentation of the, ii. 341.
Echidna, i. 201.
Echini, bright colours of some, i. 322.
Echinodermata, absence of secondary sexual characters in, i. 321.

Page 333

Ecker, figure of the human embryo, i. 15;
on sexual differences in the pelvis in man, ii. 317;
on the presence of a sagittal crest in Australians, ii. 319.
Edentata, former wide range of, in America, i. 219;
absence of secondary sexual characters in, i. 268.
Edolius, racket-shaped feathers in, ii. 73.
Edwards, Mr., on the proportion of the sexes in North American species of Papilio, i. 309.
Egerton, Sir P., on the use of the antlers of deer, ii. 252;
on the pairing of red deer, ii. 269;
on the bellowing of stags, ii. 275.
Eggs, hatched by male fishes, ii. 20.
Egret, Indian, sexes and young of, ii. 217.
Egrets, breeding plumage of, ii. 82;
white, ii. 228.
Ehrenberg, on the mane of the male Hamadryas baboon, ii. 267.
Ekström, M., on Harelda glacialis, ii. 122.
Elachista rufocinerea, habits of male, i. 311.
Eland, development of the horns of the, i. 289.
Elands, sexual differences of colour in, ii. 288.
Elaphomyia, sexual differences in, i. 349.
Elaphrus uliginosus, stridulation of, i. 379.
Elaps, ii. 31.
Elateridæ, proportions of the sexes in, i. 313.
Elaters, luminous, i. 345.
Elephant, i. 200;
nakedness of the, i. 148;
rate of increase of the, i. 135;
Indian, polygamous habits of the, i. 267;
pugnacity of the male, ii. 240;
tusks of, ii. 242, 243, 248, 249, 258;
Indian, mode of fighting, of the, ii. 257;
male, odour emitted by the, ii. 279;
attacking white or grey horses, ii. 295.
Elevation of abode, modifying influence of, i. 120.
Elimination of inferior individuals, i. 172.
Elk, ii. 249;
winter change of the, ii. 299.
Elk, Irish, horns of the, ii. 259.
Ellice Islands, beards of the natives, ii. 322, 349.
Elliot, R., on the numerical proportion of the sexes in young rats, i. 305;
on the proportion of the sexes in sheep, i. 305.
Elliott, D. G., on Pelecanus erythrorhynchus, ii. 80.
Elliott, Sir W., on the polygamous habits of the Indian wild boar, i. 267.
Ellis, on the prevalence of infanticide in Polynesia, ii. 364.
Elphinstone, Mr., on local differences of stature among the Hindoos, i. 115;
on the difficulty of distinguishing the native races of India, i. 215.
Elytra, of the females of Dytiscus, Acilius, Hydroporus, i. 343.
Emberiza, characters of young, ii. 184.
Emberiza miliaria, ii. 185.
Emberiza schœniculus, ii. 111;

Page 334

head-feathers of the male, ii. 95.
Embryo of man, i. 14, 15;
of the dog, i. 15.
Embryos of mammals, resemblance of the, i. 32.
Emigration, i. 172.
Emotions experienced by the lower animals in common with man, i. 39;
manifested by animals, i. 42.
Emperor moth, i. 398.
Emulation of singing-birds, ii. 53.
Emu, sexes and incubation of, ii. 204.
Endurance, estimation of, i. 95.
Energy, a characteristic of men, ii. 328.
England, numerical proportion of male and female births in, i. 300.
Engleheart, Mr., on the finding of new mates by starlings, ii. 106.
English, success of, as colonists, i. 179.
Engravers, short-sighted, i. 118.
Entomostraca, i. 332.
Entozoa, difference of colour between the males and females of some, i. 321.
Eocene, possible divergence of man during the, i. 200.
Eolidæ, colours of, produced by the biliary glands, i. 323.
Epeira, i. 337.
Epeira nigra, small size of the male of, i. 338.
Ephemeræ, i. 341.
Ephemeridæ, i. 361.
Ephemerina, proportions of the sexes in, i. 314.
Ephippiger vitium, stridulating organs of, i. 354, 358.
Epicalia, sexual differences of colouring in the species of, i. 388.
Equus hemionus, winter change of, ii. 298.
Erateina, coloration of, i. 397.
Erect attitude of man, i. 141, 142.
Eschricht, on the development of hair in man, i. 24;
on a lanuginous moustache in a female fœtus, i. 25;
on the want of definition between the scalp and the forehead in some children, i. 192;
on the arrangement of the hair in the human fœtus, i. 193;
on the hairiness of the face in the human fœtus of both sexes, ii. 379, 380.
Esmeralda, difference of colour in the sexes of, i. 368.
Esox lucius, i. 308.
Esox reticulatus, ii. 14.
Esquimaux, i. 157, 167;
their belief in the inheritance of dexterity in seal-catching, i. 117;
mode of life of, i. 246.
Estrelda amandava, pugnacity of the male, ii. 49.
Eubagis, sexual differences of colouring in the species of, i. 389.
Euchirus longimanus, sound produced by, i. 381.
Eudromias morinellus, ii. 203.
Eulampis jugularis, colours of the female, ii. 168.
Euler, on the rate of increase in the United States, i. 131.
Eumomota superciliaris, racket-shaped feathers in the tail of, ii. 73.
Eupetomena macroura, colours of the female, ii. 168.
Euphema splendida, ii. 174.

Page 335

Euplocamus erythropthalmus, possession of spurs by the female, ii. 46.
Euplœa midamas, mimickry of, by the female of Diadema anomala, i. 413.
Europe, ancient inhabitants of, i. 237.
Europeans, difference of, from Hindoos, i. 240;
hairiness of, probably due to reversion, ii. 378.
Eurostopodus, sexes of, ii. 206.
Eurygnathus, different proportions of the head in the sexes of, i. 344.
Eustephanus, sexual differences of species of, ii. 39;
young of, ii. 220.
Exaggeration of natural characters by man, ii. 351.
Exogamy, ii. 360, 364.
Expression, resemblances in, between man and the apes, i. 191.
Extinction of races, causes of, i. 238.
Eye, destruction of the, i. 116;
change of position in, i. 147;
obliquity of, regarded as a beauty by the Chinese and Japanese, ii. 345.
Eyebrows, elevation of, i. 19;
development of long hairs in, i. 25;
in monkeys, i. 192;
eradicated in parts of South America and Africa, ii. 340;
eradication of, by the Indians of Paraguay, ii. 348.
Eyelids, coloured black, in part of Africa, ii. 339.
Eyelashes, eradication of, by the Indians of Paraguay, ii. 348.
Eyes, difference in the colour of, in the sexes of birds, ii. 128;
pillared, of the male of Chloëon, i. 341.
Eyton, T. C., observations on the development of the horns in the fallow-deer, i. 288.
Eyzies, Les, human remains from, i. 237.

F.
Fabre, M., on the habits of Cerceris, i. 364.
Facial bones, causes of modification of the, i. 147.
Faculties, mental, variation of, in the same species, i. 36;
diversity of, in the same race of men, i. 109;
inheritance of, i. 110;
diversity of, in animals of the same species, i. 110;
of birds, ii. 108.
Fakirs, Indian, tortures undergone by, i. 96.
Falco leucocephalus, ii. 214.
Falco peregrinus, ii. 104, 179.
Falco tinnunculus, ii. 109.
Falcon, peregrine, new mate found by, ii. 104.
Falconer, H., on the mode of fighting of the Indian elephant, ii. 257;
on canines in a female deer, ii. 258;
on Hyomoschus aquaticus, ii. 304.
Falkland islands, horses of, i. 236.
Fallow-deer, different coloured herds of, ii. 295.
Famines, frequency of, among savages, i. 333.
Farr, Dr., on the structure of the uterus, i. 123;
on the effects of profligacy, i. 173;
on the influence of marriage on mortality, i. 175.

Page 336

Farrar, F. W., on the origin of language, i. 56;
on the crossing or blending of languages, i. 60;
on the absence of the idea of God in certain races of men, i. 65;
on early marriages of the poor, i. 173;
on the middle ages, i. 178.
Fashions, long prevalence of, among savages, ii. 343, 352.
Faye, Prof., on the numerical proportion of male and female births in Norway and Russia, i. 301;
on the greater mortality of male children at and before birth, i. 302.
Feathers, modified, producing sounds, ii. 63 et seqq., 163;
elongated, in male birds, ii. 72, 97;
racket-shaped, ii. 73;
barbless and with filamentous barbs in certain birds, ii. 74;
shedding of margins of, ii. 85.
Feeding, high, probable influence of, in the pairing of birds of different species, ii. 115.
Feet, modification of, in man, i. 141;
thickening of the skin on the soles of the, i. 118.
Felis canadensis, throat-ruff of, ii. 267.
Felis pardalis and F. mitis, sexual differences in the colouring of, ii. 287.
Female, behaviour of the, during courtship, i. 273.
Female birds, differences of, ii. 193.
Females, presence of rudimentary male organs in, i. 208;
preference of, for certain males, i. 262;
pursuit of, by males, i. 272;
occurrence of secondary sexual characters in, i. 276;
development of male characters by, i. 280.
Females and males, comparative mortality of, while young, i. 264, 276;
comparative numbers of, i. 261, 263.
Femur and tibia, proportions of, in the Aymara Indians, i. 119.
Ferguson, Mr., on the courtship of fowls, ii. 118.
Fertilization, phenomena of, in plants, i. 273;
in the lower animals, i. 274.
Fevers, immunity of Negroes and Mulattoes from, i. 243.
Fiber zibethicus, protective colouring of, ii. 298.
Fidelity of savages to one another, i. 95;
importance of, i. 162.
Field-slaves, difference of, from house-slaves, i. 246.
Fijians, burying their old and sick parents alive, i. 77;
estimation of the beard among the, ii. 349;
admiration of, for a broad occiput, ii. 352.
Fiji Islands, beards of the natives, ii. 322, 349;
marriage-customs of the, ii. 373.
Filial affection, partly the result of natural selection, i. 81.
Filum terminale, i. 30.
Finch, racket-shaped feathers in the tail of a, ii. 73.
Finches, spring change of colour in, ii. 85;
British, females of the, ii. 193.
Fingers, partially coherent, in species of Hylobates, i. 140.
Finlayson, on the Cochin Chinese, ii. 345.
Fire, use of, i. 137, 183, 234.
Fischer, on the pugnacity of the male of Lethrus cephalotes, i. 376.

Page 337

Fish, proportion of the sexes in, i. 307;
eagerness of male, i. 272.
Fishes, kidneys of, represented by Corpora Wolffiana in the human embryo, i. 16;
male, hatching ova in their mouths, i. 210;
receptacles for ova possessed by, i. 254;
relative size of the sexes in, ii. 7;
freshwater, of the tropics, ii. 17;
protective resemblances in, ii. 18;
nest-building, ii. 19;
spawning of, ii. 19;
sounds produced by, ii. 23, 331;
continued growth of, ii. 216.
Flexor pollicis longus, similar variation of, in man, i. 129.
Flint tools, i. 183.
Flints, difficulty of chipping into form, i. 138.
Florida, Quiscalus major in, i. 307.
Flounder, coloration of the, ii. 18.
Flower, W. H., on the abductor of the fifth metatarsal in apes, i. 128;
on the position of the Seals, i. 190;
on the throat-pouch of the male Bustard, ii. 58.
Fly-catchers, colours and nidification of, ii. 170.
Fœtus, human, woolly covering of the, i. 25;
arrangement of the hair on, i. 193.
Food, influence of, upon stature, i. 115.
Foot, prehensile, in the early progenitors of man, i. 206;
prehensile power of the, retained in some savages, i. 142.
Foramen, supra-condyloid, exceptional occurrence of in the humerus of man, i. 28, 130;
in the early progenitors of man, i. 206.
Forbes, D., on the Aymara Indians, i. 119;
on local variation of colour in the Quechuas, i. 246;
on the hairlessness of the Aymaras and Quechuas, ii. 322;
on the long hair of the Aymaras and Quechuas, ii. 320, 348.
Forel, F., on white young swans, ii. 211.
Formica rufo, size of the cerebral ganglia in, i. 145.
Fossils, absence of, connecting man with the apes, i. 201.
Fowl, occurrence of spurs in the female, i. 280;
game, early pugnacity of, i. 295;
Polish, early development of cranial peculiarities of, i. 295;
variations in plumage of, ii. 74;
examples of correlated development in the, ii. 130;
domestic, breeds and sub-breeds of, ii. 178.
Fowls, spangled Hamburgh, i. 281, 294;
sexual peculiarities in, transmitted only to the same sex, i. 283;
loss of secondary sexual characters by male, i. 284;
inheritance of changes of plumage by, i. 281;
Polish, origin of the crest in, i. 284;
period of inheritance of characters by, i. 294;
cuckoo-, i. 294;
development of the comb in, i. 295;
numerical proportion of the sexes in, i. 306;

Page 338

courtship of, ii. 117;
mongrel, between a black Spanish cock and different hens, ii. 131;
pencilled Hamburgh, difference of the sexes in, ii. 158;
Spanish, sexual differences of the comb in, ii. 158;
spurred, in both sexes, ii. 162.
Fox, W. D., on some half-tamed wild ducks becoming polygamous, and on polygamy in the guinea-
fowl and canary-bird, i. 270;
on the proportion of the sexes in cattle, i. 305;
on the pugnacity of the peacock, ii. 46;
on a nuptial assembly of magpies, ii. 102;
on the finding of new mates by crows, ii. 104;
on partridges living in triplets, ii. 107;
on the pairing of a goose with a Chinese gander, ii. 114.
Foxes, wariness of young, in hunting districts, i. 50;
black, ii. 294.
France, numerical proportion of male and female births in, i. 301.
Francesco, B., on the Simian resemblances of man, i. 4.
Fraser, C., on the different colours of the sexes in a species of Squilla, i. 335.
Fringilla cannabina, ii. 86.
Fringilla ciris, age of mature plumage in, ii. 213.
Fringilla cyanea, age of mature plumage in, ii. 213.
Fringilla leucophrys, young of, ii. 217.
Fringilla spinus, ii. 115.
Fringilla tristis, change of colour in, in spring, ii. 85;
young of, ii. 216.
Fringillidæ, resemblance of the females of distinct species of, ii. 192.
Frogs, ii. 25;
male, temporary receptacles for ova possessed by, i. 254;
ready to breed before the females, i. 260;
vocal organs of, ii. 28.
Frontal bone, persistence of the suture in, i. 124.
Fruits, poisonous, avoided by animals, i. 36.
Fuegians, i. 167, 181;
mental capacity of the, i. 34;
quasi-religious sentiments of the, i. 67;
power of sight in the, i. 118;
skill of, in stone-throwing, i. 138;
resistance of the, to their severe climate, i. 156, 237;
difference of stature among the, i. 115;
mode of life of the, i. 246;
resemblance of, in mental characters, to Europeans, i. 232;
aversion of, to hair on the face, ii. 348;
said to admire European women, ii. 351.
Fulgoridæ, songs of the, i. 351.
Fur, whiteness of, in arctic animals, in winter, i. 282.
Fur-bearing animals, acquired sagacity of, i. 50.

G.
Gallicrex, sexual difference in the colour of the irides in, ii. 128.
Gallicrex cristatus, red caruncle occurring in the male during the breeding-season, ii. 80.

Page 339

Gallinaceæ, frequency of polygamous habits and of sexual differences in the, i. 269;
love-gestures of, ii. 68;
decomposed feathers in, ii. 74;
stripes of young, ii. 184;
comparative sexual differences between the species of, ii. 192, 194;
plumage of, ii. 195.
Gallinaceous birds, weapons of the male, ii. 44;
racket-shaped feathers on the heads of, ii. 73.
Gallinula chloropus, pugnacity of male, ii. 40.
Gallinula cristata, pugnacity of the male, ii. 41.
Galloperdix, spurs of, ii. 46;
development of spurs in the female, ii. 162.
Gallophasis, young of, ii. 190.
Gallus bankiva, ii. 158;
neck-hackles of, ii. 84.
Gallus Stanleyi, pugnacity of the male, ii. 44.
Galls, i. 152.
Galton, Mr., on the struggle between the social and personal impulses, i. 104;
on hereditary genius, i. 111;
on the effects of natural selection on civilised nations, i. 168;
on the sterility of sole daughters, i. 170;
on the degree of fertility of people of genius, i. 171;
on the early marriages of the poor, i. 173;
on the ancient Greeks, i. 177;
on the Middle Ages, i. 178;
on the progress of the United States, i. 179;
on South African notions of beauty, ii. 347.
Gammarus, use of the chelæ of, i. 331.
Gammarus marinus, i. 334.
Gannets, white only when mature, ii. 228.
Ganoidei, i. 204.
Ganoid fishes, i. 212.
Gaour, horns of the, ii. 247.
Gap between man and the apes, i. 200.
Gaper, sexes and young of, ii. 217.
Gardner, on an example of rationality in a Gelasimus, i. 334.
Garrulus glandarius, ii. 104.
Gärtner, on sterility of hybrid plants, i. 223.
Gasteropoda, i. 324;
pulmoniferous, courtship of, i. 324.
Gasterosteus, i. 271;
nidification of, ii. 20.
Gasterosteus leiurus, ii. 2, 14, 20.
Gasterosteus trachurus, ii. 2.
Gastrophora, wings of, brightly coloured beneath, i. 397.
Gauchos, want of humanity among the, i. 101.
Gaudry, M., on a fossil monkey, i. 197.
Gavia, seasonal change of plumage in, ii. 228.
Geese, clanging noise made by, ii. 51;
pairing of different species of, ii. 114;

Page 340

Canada, selection of mates by, ii. 116.
Gegenbaur, C., on the number of digits in the Ichthyopterygia, i. 125;
on the hermaphroditism of the remote progenitors of the vertebrata, i. 207.
Gelasimus, use of the enlarged chela of the male, i. 331;
pugnacity of males of, i. 333;
proportions of the sexes in a species of, i. 315;
rational actions of a, i. 334;
difference of colour in the sexes of a species of, i. 336.
Gemmules, sexual selection of, i. 285.
Genesis, i. 318.
Genius, ii. 328;
hereditary, i. 111.
Genius, fertility of men and women of, i. 171.
Geoffroy-Saint-Hilaire, Isid., on the recognition of women by male quadrumana, i. 13;
on the occurrence of a rudimentary tail in man, i. 29;
on monstrosities, i. 113;
on animal-like anomalies in the human structure, i. 125;
on the correlation of monstrosities, i. 130;
on the distribution of hair in man and monkeys, i. 149;
on the caudal vertebræ of monkeys, i. 150;
on correlated variability, i. 151;
on the classification of man, i. 186;
on the long hair on the heads of species of Semnopithecus, i. 192;
on the hair in monkeys, i. 194;
on the development of horns in female deer, ii. 244;
and F. Cuvier, on the mandrill, ii. 293;
on Hylobates, ii. 318, 320.
Geographical distribution, as evidence of specific distinctions in man, i. 218.
Geometræ, brightly coloured beneath, i. 397.
Geophagus, frontal protuberance of male, ii. 13, 20;
eggs hatched by the male, in the mouth or branchial cavity, ii. 200.
Georgia, change of colour in Germans settled in, i. 246.
Geotrupes, stridulation of, i. 380, 382.
Gerbe, M., on the nest-building of Crenilabrus massa and C. melops, ii. 19.
Gerland, Dr., on the prevalence of infanticide, i. 94; ii. 344, 364;
on the extinction of races, i. 237, 238.
Gervais, P., on the hairiness of the gorilla, i. 149;
on the mandrill, ii. 293.
Gesture-language, i. 232.
Ghost-moth, sexual difference of colour in the, i. 399, 402.
Gibb, Sir D., on differences of the voice in different races of men, ii. 330.
Gibbon, Hoolock, nose of, i. 192.
Gibbons, voice of, ii. 276.
Giraffe, mute, except in the rutting season, ii. 274;
its mode of using the horns, ii. 250.
Giraud-Teulon, on the cause of short sight, i. 118.
Glanders, communicable between man and the lower animals, i. 11.
Glands, odoriferous, in mammals, ii. 279, 281.
Glareola, double moult in, ii. 80.
Glomeris limbata, difference of colour in the sexes of, i. 340.

Page 341

Glowworm, female, apterous, i. 255;
luminosity of the, i. 345.
Gnats, dances of, i. 349.
Gnu, sexual differences in the colour of the, ii. 289.
Goat, male, wild, falling on his horns, ii. 249;
male, odour emitted by, ii. 279;
male, wild, crest of the, ii. 282;
Berbura, mane, dewlap, &c., of the male, ii. 284;
Kemas, sexual difference in the colour of the, ii. 289.
Goats, sexual differences in the horns of, i. 283;
horns of, i. 289, ii. 246;
domestic, sexual differences of, late developed, i. 293;
beards of, ii. 282;
mode of fighting of, ii. 249, 250.
Goat-sucker, Virginian, pairing of the, ii. 49.
Gobies, nidification of, ii. 20.
God, want of the idea of, in some races of men, i. 65.
Godron, M., on variability, i. 112;
on difference of stature, i. 115;
on the want of connexion between climate and the colour of the skin, i. 241;
on the odour of the skin, i. 248;
on the colour of infants, ii. 318.
Goldfinch, ii. 56, 85;
proportion of the sexes in the, i. 307;
sexual differences of the beak in the, ii. 39;
courtship of the, ii. 95.
Goldfinch, North American, young of, ii. 216.
Gold-Fish, ii. 16.
Gomphus, proportions of the sexes in, i. 314;
difference in the sexes of, i. 362.
Gonepteryx Rhamni, i. 393;
sexual difference of colour in, i. 409.
Goodsir, Prof., on the affinity of the lancelet to the ascidians, i. 205.
Goosander, young of, ii. 189.
Goose, Antarctic, colours of the, ii. 228.
Goose, Canada, pairing with a Bernicle gander, ii. 114.
Goose, Chinese, knob on the beak of the, ii. 129.
Goose, Egyptian, ii. 46.
Goose, Sebastopol, plumage of, ii. 74.
Goose, Snow-, whiteness of the, ii. 228.
Goose, Spur-winged, ii. 46.
Gorilla, ii. 323;
semi-erect attitude of the, i. 142;
mastoid processes of the, i. 143;
direction of the hair on the arms of the, i. 192;
supposed evolution of the, i. 230;
polygamy of the, i. 266, ii. 361, 362;
voice of the, ii. 276;
cranium of, ii. 318;
fighting of male, ii. 324.

Page 342

Gosse, P. H., on the pugnacity of the male Humming-birds, ii. 40.
Gosse, M., on the inheritance of artificial modifications of the skull, ii. 380.
Gould, B. A., on variation in the length of the legs in man, i. 108;
measurements of American soldiers, i. 114, 116;
on the proportions of the body and capacity of the lungs in different races of men, i. 216;
on the inferior vitality of mulattoes, i. 221.
Gould, J., on the arrival of male snipes before the females, i. 260;
on the numerical proportion of the sexes in birds, i. 306;
on Neomorpha, ii. 39;
on the species of Eustephanus, ii. 39;
on the Australian Musk-duck, ii. 39;
on the relative size of the sexes in Biziura lobata and Cincloramphus cruralis, ii. 43;
on Lobivanellus lobatus, ii. 48;
on the habits of Menura Alberti, ii. 56;
on the rarity of song in brilliant birds, ii. 58;
on Selasphorus platycercus, ii. 65;
on the Bower-birds, ii. 69, 102;
on the ornamental plumage of the Humming-birds, ii. 78;
on the moulting of the ptarmigan, ii. 83;
on the display of plumage by the male Humming-birds, ii. 86;
on the shyness of adorned male birds, ii. 97;
on the decoration of the bowers of Bower-birds, ii. 112;
on the decoration of their nests by Humming-birds, ii. 112;
on variation in the genus Cynanthus, ii. 125;
on the colour of the thighs in a male parrakeet, ii. 126;
on Urosticte Benjamini, ii. 151, 152;
on the nidification of the Orioles, ii. 168;
on obscurely-coloured birds building concealed nests, ii. 169;
on Trogons and Kingfishers, ii. 173;
on Australian parrots, ii. 174;
on Australian pigeons, ii. 175;
on the moulting of the ptarmigan, ii. 181;
on the immature plumage of birds, ii. 186 et seq.;
on the Australian species of Turnix, ii. 201;
on the young of Aithurus polytmus, ii. 220;
on the colours of the bills of Toucans, ii. 227;
on the relative size of the sexes in the Marsupials of Australia, ii. 260;
on the colours of the Marsupials, ii. 286.
Goureau, on the stridulation of Mutilla europæa, i. 366.
Gout, sexually transmitted, i. 292.
Graba, on the Pied Ravens of the Feroe Islands, ii. 126;
on the Bridled Guillemot, ii. 127.
Gradation of secondary sexual characters in birds, ii. 135.
Grallatores, absence of secondary sexual characters in, i. 270;
double moult in some, ii. 81.
Grallina, nidification of, ii. 169.
Grasshoppers, stridulation of the, i. 356.
Gratiolet, Prof., on the anthropomorphous apes, i. 196;
on the evolution of the anthropomorphous apes, i. 230.
Gray, Asa, on the gradation of species among the Compositæ, i. 227.

Page 343

Gray, J. E., on the caudal vertebræ of monkeys, i. 150;
on the presence of rudiments of horns in the female of Cervulus moschatus, ii. 245;
on the horns of goats and sheep, ii. 246;
on the beard of the Ibex, ii. 283;
on the Berbura goat, ii. 285;
on sexual differences in the coloration of Rodents, ii. 286;
on the colours of the Elands, ii. 288;
on the Sing-sing antelope, ii. 289;
on the colours of goats, ii. 290;
on the Hog-deer, ii. 303.
“Greatest happiness principle,” i. 97, 98.
Greeks, ancient, i. 177.
Green, A. H., on beavers fighting, ii. 239;
on the voice of the beaver, ii. 277.
Greenfinch, selected by a female canary, ii. 115.
Greg, W. R., on the early marriages of the poor, i. 173;
on the Ancient Greeks, i. 178;
on the effects of natural selection on civilised nations, i. 167.
Grenadiers, Prussian, i. 112.
Grey, Sir G., on female infanticide in Australia, ii. 364.
Greyhounds, numerical proportion of the sexes in, i. 263, 265;
numerical proportion of male and female births in, i. 304.
Grouse, red, monogamous, i. 269;
pugnacity of young male, ii. 48;
producing a sound by scraping their wings upon the ground, ii. 61;
duration of courtship of, ii. 100;
colours and nidification of, ii. 170.
Grube, Dr., on the occurrence of the supra-condyloid foramen in the humerus of man, i. 28.
Grus americanus, age of mature plumage in, ii. 213;
breeding in immature plumage, ii. 214.
Grus virgo, trachea of, ii. 60.
Gryllus campestris, i. 353;
pugnacity of male, i. 360.
Gryllus domesticus, i. 354.
Grypus, sexual differences in the beak in, ii. 39.
Guanacoes, battles of, ii. 239;
canine teeth of, ii. 257.
Guanas, strife for women among the, ii. 324;
polyandry among the, ii. 366.
Guanche skeletons, occurrence of the supra-condyloid foramen in the humerus of, i. 29.
Guaranys, proportion of men and women among, i. 302;
colour of newborn children of the, ii. 318;
beards of the, ii. 322.
Guenée, A., on the sexes of Hyperythra, i. 310.
Guilding, L., on the stridulation of the LOCUSTIDÆ, i. 352.
Guillemot, variety of the, ii. 127.
Guinea, sheep of, with males only horned, i. 289.
Guinea-fowl, monogamous, i. 269;
occasional polygamy of the, i. 270;
markings of the, ii. 134.

Page 344

Guinea-pigs, inheritance of the effects of operations by, ii. 380.
Gull, instance of reasoning in a, ii. 108.
Gulls, seasonal change of plumage in, ii. 228;
white, ii. 228.
Günther, Dr., on hermaphroditism in Serranus, i. 208;
on male fishes hatching ova in their mouths, i. 210, ii. 20;
on mistaking infertile female fishes for males, i. 308;
on the prehensile organs of male Plagiostomous fishes, ii. 2;
on the pugnacity of the male salmon and trout, ii. 3;
on the relative size of the sexes in fishes, ii. 7;
on sexual differences in fishes, ii. 8 et seqq.;
on the genus Callionymus, ii. 9;
on a protective resemblance in a Pipe-fish, ii. 18;
on the genus Solenostoma, ii. 22;
on Megalophrys montana, ii. 26;
on the coloration of frogs and toads, ii. 26;
on sexual differences in the Ophidia, ii. 29;
on differences of the sexes of lizards, ii. 32 et seqq.
Gynanisa Isis, ocellated spots of, ii. 132.
Gypsies, uniformity of, in various parts of the world, i. 242.

H.
Habits, bad, facilitated by familiarity, i. 101;
variability of the force of, i. 183.
Häckel, E., on the origin of man, i. 4;
on rudimentary characters, i. 17;
on the canine teeth in man, i. 126;
on death caused by inflammation of the vermiform appendage, i. 28;
on the steps by which man became a biped, i. 142;
on man as a member of the Catarrhine group, i. 199;
on the position of the Lemuridæ, i. 202;
on the genealogy of the Mammalia, i. 203;
on the lancelet, i. 204;
on the transparency of pelagic animals, i. 323;
on the musical powers of women, ii. 337.
Hagen, H., and Walsh, B. D., on American neuroptera, i. 314.
Hair, development of, in man, i. 24;
character of, supposed to be determined by light and heat, i. 116;
distribution of, in man, i. 149, ii. 375;
possibly removed for ornamental purposes, i. 149;
arrangement and direction of, i. 192;
of the early progenitors of man, i. 206;
different texture of, in distinct races, i. 216;
and skin, correlation of colour of, i. 248;
development of, in mammals, ii. 281;
management of, among different peoples, ii. 340;
great length of, in some North American tribes, ii. 348;
elongation of the, on the human head, ii. 380.
Hairiness, difference of, in the sexes, in man, ii. 320;
variation of, in races of men, ii. 321.

Page 345

Hairs and excretory pores, numerical relation of, in sheep, i. 248.
Hairy family, Siamese, ii. 378.
Hamadryas baboon, turning over stones, i. 75;
mane of the male, ii. 267.
Hamilton, C., on the cruelty of the Kafirs to animals, i. 94;
on the engrossment of the women by the Kafir chiefs, ii. 369.
Hammering, difficulty of, i. 138.
Hancock, A., on the colours of the nudibranch mollusca, i. 326.
Hands, larger at birth, in the children of labourers, i. 117;
structure of, in the quadrumana, i. 139;
and arms, freedom of, indirectly correlated with diminution of canines, i. 144.
Handwriting, inherited, i. 58.
Harcourt, E. Vernon, on Fringilla cannabina, ii. 86.
Harelda glacialis, ii. 122.
Hare, protective colouring of the, ii. 298. Hares, battles of male, ii. 239.
Harlan, Dr., on the difference between field- and house-slaves, i. 246.
Harris, J. M., on the relation of complexion to climate, i. 245.
Harris, T. W., on the Katy-did locust, i. 353;
on the stridulation of the grasshoppers, i. 357;
on Œcanthus nivalis, i. 361;
on the colouring of Lepidoptera, i. 396;
on the colouring of Saturnia Io, i. 398.
Harry-long-legs, pugnacity of male, i. 349.
Hartman, Dr., on the singing of Cicada septendecim, i. 351.
Haughton, S., on a variation of the flexor pollicis longus in man, i. 129.
Hawks, feeding orphan nestling, ii. 107.
Hayes, Dr., on the diverging of sledge-dogs on thin ice, i. 46.
Head, altered position of, to suit the erect attitude of man, i. 143;
hairiness of, in man, i. 149;
processes of, in male beetles, i. 370;
artificial alterations of the form of the, ii. 351.
Hearne, on strife for women among the North American Indians, ii. 324;
on the North American Indians’ notion of female beauty, ii. 344;
repeated elopements of a North American woman, ii. 372.
Heart, in the human embryo, i. 16.
Heat, supposed effects of, i. 116.
Hectocotyle, i. 325.
Hedge-warbler, ii. 198;
young of the, ii. 209.
Heel, small projection of, in the Aymara Indians, i. 120.
Hegt, M., on the development of the spurs in peacocks, i. 290.
Heliconidæ, i. 387;
mimickry of, by other butterflies, i. 411.
Heliopathes, stridulation peculiar to the male, i. 383.
Heliothrix auriculata, young of, ii. 188, 189.
Helix pomatia, example of individual attachment in, i. 325.
Hellins, J., proportions of sexes of Lepidoptera reared by, i. 313.
Helmholtz, on the vibration of the auditory hairs of crustacea, ii. 333.
Hemiptera, i. 349.
Hemitragus, beardless in both sexes, ii. 283.

Page 346

Hepburn, Mr., on the autumn song of the water-ouzel, ii. 54.
Hepialus humuli, sexual difference of colour in the, i. 399, 402.
Herbs, poisonous, avoided by animals, i. 36.
Hermaphroditism of embryos, i. 207.
Herodias bubulcus, vernal moult of, ii. 84.
Heron, love-gestures of a, ii. 68.
Heron, Sir R., on the habits of peafowl, ii. 119, 120, 152.
Herons, decomposed feathers in, ii. 74;
breeding plumage of, ii. 82, 83;
young of the, ii. 208;
sometimes dimorphic, ii. 214;
continued growth of crest and plumes in the males of some, ii. 216;
change of colour in some, ii. 231.
Hetærina, difference in the sexes of, i. 362;
proportion of the sexes in, i. 314.
Heterocerus, stridulation of, i. 379.
Hewitt, Mr. on a game-cock killing a kite, ii. 44;
on the recognition of dogs and cats by ducks, ii. 110;
on the pairing of a wild duck with a pintail drake, ii. 115;
on the courtship of fowls, ii. 117;
on the coupling of pheasants with common hens, ii. 122.
Hindoo, his horror of breaking his caste, i. 99, 103.
Hindoos, local difference of stature among, i. 115;
difference of, from Europeans, i. 240;
colour of the beard in, ii. 319.
Hipparchia Janira, instability of the ocellated spots of, ii. 132.
Hipparchiæ, i. 387.
Hippocampus, development of, i. 210;
marsupial receptacles of the male, ii. 21.
Hippopotamus, nakedness of, i. 148.
Hips, proportions of, in soldiers and sailors, i. 117.
Hodgson, S., on the sense of duty, i. 71.
Hoffberg, on the horns of the reindeer, ii. 244;
on sexual preferences shown by reindeer, ii. 273.
Hog, wart-, ii. 265;
river-, ii. 266.
Hog-deer, ii. 303.
Holland, Sir H., on the effects of new diseases, i. 238.
Homologous structures, correlated variation of, i. 130.
Homoptera, i. 350;
stridulation of the, and orthoptera, discussed, i. 360.
Honduras, Quiscalus major in, i. 307.
Honey-buzzard of India, variation in the crest of, ii. 126.
Honey-suckers, moulting of the, ii. 83;
Australian, nidification of, ii. 169.
Honour, law of, i. 99.
Hooker, Jos., on the colour of the beard in man, ii. 319.
Hoolock Gibbon, nose of, i. 192.
Hoopoe, ii. 56;
sounds produced by the male, ii. 62.

Page 347

Hoplopterus armatus, wing-spurs of, ii. 48.
Hornbill, African, inflation of the neck-wattle of the male during courtship, ii. 72.
Hornbills, sexual difference in the colour of the eyes in, ii. 129;
nidification and incubation of, ii. 169.
Horne, C., on the rejection of a brightly-coloured locust by lizards and birds, i. 361.
Horns, of deer, ii. 243, 248, 259;
and canine teeth, inverse development of, ii. 257;
sexual differences of, in sheep and goats, i. 283;
loss of, in female merino sheep, i. 284;
development of, in deer, i. 288;
development of, in antelopes, i. 289;
from the head and thorax, in male beetles, i. 370.
Horse, polygamous, i. 267;
canine teeth of male, ii. 241;
winter change of the, ii. 298;
fossil, extinction of the, in South America, i. 239.
Horses, dreaming, i. 46;
rapid increase of, in South America, i. 135;
diminution of canine teeth in, i. 144;
of the Falkland Islands and Pampas, i. 236;
numerical proportion of the sexes in, i. 263, 265;
lighter in winter in Siberia, i. 282;
sexual preferences in, ii. 272;
pairing preferentially the same colour, ii. 295;
numerical proportion of male and female births in, i. 303;
formerly striped, ii. 305.
Hottentot women, peculiarities of, i. 225.
Hottentots, lice of, i. 220;
readily become musicians, ii. 334;
notions of female beauty of the, ii. 345;
compression of nose by, ii. 352.
House-slaves, difference of, from field-slaves, i. 246.
Huber, P., on ants playing together, i. 39;
on memory in ants, i. 45;
on the intercommunication of ants, i. 58;
on the recognition of each other by ants after separation, i. 365.
Huc, on Chinese opinions of the appearance of Europeans, ii. 345.
Human kingdom, i. 186.
Human sacrifices, i. 68.
Humanity, unknown among some savages, i. 94;
deficiency of, among savages, i. 101.
Humboldt, A. von, on the rationality of mules, i. 48;
on a parrot preserving the language of a lost tribe, i. 236;
on the cosmetic arts of savages, ii. 339, 340;
on the exaggeration of natural characters by man, ii. 351;
on the red painting of American Indians, ii. 352.
Hume, D., on sympathetic feelings, i. 85.
Humming-bird, racket-shaped feathers in the tail of a, ii. 73;
display of plumage by the male, ii. 86.
Humming-birds, ornament their nests, i. 63, ii. 112;

Page 348

polygamous, i. 269;
proportion of the sexes in, i. 307, ii. 221;
sexual differences in, ii. 39, 40, 151;
pugnacity of male, ii. 40;
modified primaries of male, ii. 65;
coloration of the sexes of, ii. 78;
young of, ii. 220;
nidification of the, ii. 168;
colours of female, ii. 168.
Humphreys, H. N., on the habits of the Stickle-back, i. 271, ii. 2.
Hunger, instinct of, i. 89.
Huns, ancient, flattening of the nose by the, ii. 352.
Hunter, J., on the number of species of man, i. 226;
on secondary sexual characters, i. 253;
on the general behaviour of female animals during courtship, i. 273;
on the muscles of the larynx in song-birds, ii. 55;
on the curled frontal hair of the Bull, ii. 282;
on the rejection of an ass by a female zebra, ii. 295.
Hunter, W. W., on the recent rapid increase of the Santali, i. 133;
on the Santali, i. 241.
Hussey, Mr., on a partridge distinguishing persons, ii. 110.
Hutchinson, Col., example of reasoning in a retriever, i. 48.
Hutton, Capt., on the male wild goat falling on his horns, ii. 249.
Huxley, T. H., on the structural agreement of man with the apes, i. 3;
on the agreement of the brain in man with that of lower animals, i. 10;
on the adult age of the Orang, i. 13;
on the embryonic development of man, i. 14;
on the origin of man, i. 4, 17;
on variation in the skulls of the natives of Australia, i. 108;
on the abductor of the fifth metatarsal in apes, i. 128;
on the position of man, i. 191;
on the sub-orders of primates, i. 195;
on the Lemuridæ, i. 202;
on the Dinosauria, i. 204;
on the amphibian affinities of the Ichthyosaurians, i. 204;
on variability of the skull in certain races of man, i. 226;
on the races of man, i. 229.
Hybrid birds, production of, ii. 113.
Hydrophobia communicable between man and the lower animals, i. 11.
Hydroporus, dimorphism of females of, i. 343.
Hyelaphus porcinus, ii. 303.
Hygrogonus, ii. 21.
Hyla, singing species of, ii. 27.
Hylobates, maternal affection in a, i. 40;
absence of the thumb in, i. 140;
upright progression of some species of, i. 143;
direction of the hair on the arms of species of, i. 192;
females of, less hairy below than males, ii. 320.
Hylobates agilis, i. 140;
hair on the arms of, i. 193;

Page 349

musical voice of the, ii. 277;
superciliary ridge of, ii. 318;
voice of, ii. 332.
Hylobates hoolock, sexual difference of colour in, ii. 291.
Hylobates lar, i. 140;
hair on the arms of, i. 193.
Hylobates leuciscus, i. 140.
Hylobates syndactylus, i. 140;
laryngeal sac of, ii. 276.
Hymenoptera, i. 364;
large size of the cerebral ganglia in, i. 145;
classification of, i. 188;
sexual differences in the wings of, i. 345;
aculeate, relative size of the sexes of, i. 347.
Hymenopteron, parasitic, with a sedentary male, i. 272.
Hyomoschus aquaticus, ii. 304.
Hyperythra, proportion of the sexes in, i. 310.
Hypogymna dispar, sexual difference of colour in, i. 398.
Hypopyra, coloration of, i. 397.

I.
Ibex, male, falling on his horns, ii. 249;
beard of the, ii. 283.
Ibis, scarlet, young of the, ii. 208;
white, change of colour of naked skin in, during the breeding season, ii. 80.
Ibis tantalus, age of mature plumage in, ii. 213;
breeding in immature plumage, ii. 214, 215.
Ibises, decomposed feathers in, ii. 74;
white, ii. 228, and black, ii. 230.
Ichneumonidæ, difference of the sexes in, i. 365.
Ichthyopterygia, i. 125.
Ichthyosaurians, i. 204.
Ideas, general, i. 62.
Idiots, microcephalous, imitative faculties of, i. 57;
microcephalous, their characters and habits, i. 121.
Iguana tuberculata, ii. 32.
Iguanas, ii. 32.
Illegitimate and legitimate children, proportion of the sexes in, i. 302.
Imagination, existence of, in animals, i. 45.
Imitation, i. 39;
of man by monkeys, i. 44;
tendency to, in monkeys, microcephalous idiots and savages, i. 56;
influence of, i. 161.
Immature plumage of birds, ii. 183, 187.
Implacentata, i. 202.
Implements, employed by monkeys, i. 51;
fashioning of, peculiar to man, i. 52.
Impregnation, period of, influence of, upon sex, i. 303.
Improvement, progressive, man alone supposed to be capable of, i. 49.
Incisor teeth, knocked out or filed by some savages, ii. 340.

Page 350

Increase, rate of, i. 131;
necessity of checks in, i. 135.
Indecency, hatred of, a modern virtue, i. 96.
India, difficulty of distinguishing the native races of, i. 215;
Cyprinidæ of, ii. 17;
colour of the beard in races of men of, ii. 319.
Indian, North American, honoured for scalping a man of another tribe, i. 93.
Individuality, i. 62.
Individuation, i. 318.
Indopicus carlotta, colours of the sexes of, ii. 175.
Infanticide, prevalence of, i. 94, 134;
supposed cause of, ii. 344;
prevalence and causes of, ii. 363 et seq.
Inferiority, supposed physical, of man, i. 156.
Inflammation of the bowels, occurrence of, in Cebus Azaræ, i. 12.
Inheritance, i. 110;
of effects of use of vocal and mental organs, i. 58;
of moral tendencies, i. 102, 104;
of long and short sight, i. 118;
laws of, i. 279;
sexual, i. 285;
sexually limited, ii. 154.
Inquisition, influence of the, i. 179.
Insanity, hereditary, i. 111.
Insect, fossil, from the Devonian, i. 360.
Insectivora, ii. 286;
absence of secondary sexual characters in, i. 268.
Insects, relative size of the cerebral ganglia in, i. 145;
male, appearance of, before the females, i. 260;
pursuit of female, by the males, i. 272;
period of development of sexual characters in, i. 291;
secondary sexual characters of, i. 341;
stridulation of, ii. 331.
Insessores, vocal organs of, ii. 55.
Instep, depth of, in soldiers and sailors, i. 117.
Instinct and intelligence, i. 37.
Instinct, migratory, vanquishing the maternal, i. 83, 90.
Instinctive actions, the result of inheritance, i. 80.
Instinctive impulses, difference of the force of, i. 87, 89;
and moral impulses, alliance of, i. 88.
Instincts, i. 36;
complex origin of, through natural selection, i. 38;
possible origin of some, i. 38;
acquired, of domestic animals, i. 79;
variability of the force of, i. 83;
difference of force between the social and other, i. 89, 104;
utilised for new purposes, ii. 335.
Instrumental music of birds, ii. 61, 66.
Intellect, influence of, in natural selection in civilised society, i. 171.
Intellectual faculties, their influence on natural selection in man, i. 158;

Page 351

probably perfected through natural selection, i. 160.
Intelligence, Mr. H. Spencer on the dawn of, i. 37.
Intemperance, no reproach among savages, i. 96;
its destructiveness, i. 172.
Intoxication in monkeys, i. 12.
Iphias glaucippe, i. 394.
Iris, sexual difference in the colour of the, in birds, ii. 72, 128.
Ischio-pubic muscle, i. 127.
Ithaginis cruentus, number of spurs in, ii. 46.
Iulus, tarsal suckers of the males of, i. 340.

J.
Jackals learning to bark from dogs, i. 44.
Jack-snipe, coloration of the, ii. 226.
Jacquinot, on the number of species of man, i. 226.
Jaeger, Dr., on the difficulty of approaching herds of wild animals, i. 74;
on the increase of length in bones, i. 116;
on the deposition of a male Silver pheasant on account of spoiled plumage, ii. 120.
Jaguars, black, ii. 294.
Janson, E. W., on the proportions of the sexes in Tomicus villosus, i. 314;
on stridulant beetles, i. 379.
Japan, encouragement of licentiousness in, i. 134.
Japanese, general beardlessness of the, ii. 321;
aversion of the, to whiskers, ii. 349.
Jardine, Sir W., on the Argus pheasant, ii. 72, 97.
Jarrold, Dr., on modifications of the skull induced by unnatural position, i. 147.
Javanese, relative height of the sexes of, ii. 320;
notions of female beauty, ii. 347.
Jaw, influence of the muscles of the, upon the physiognomy of the apes, i. 144.
Jaws, smaller in the same ratio with the extremities, i. 117;
influence of food upon the size of, i. 118;
diminution of, in man, i. 144;
in man, reduced by correlation, ii. 325.
Jay, young of the, ii. 209;
Canada, young of the, ii. 209.
Jays, new mates found by, ii. 104;
distinguishing persons, ii. 110.
Jeffreys, J. Gwyn, on the form of the shell in the sexes of the Gasteropoda, i. 324;
on the influence of light upon the colours of shells, i. 326.
Jelly-fish, bright colours of some, i. 322.
Jenner, Dr., on the voice of the rook, ii. 61;
on the finding of new mates by magpies, ii. 103;
on retardation of the generative organs in birds, ii. 107.
Jenyns, L., on the desertion of their young by swallows, i. 84;
on male birds singing after the proper season, ii. 107.
Jerdon, Dr., on birds dreaming, ii. 46;
on the pugnacity of the male bulbul, ii. 41;
on the pugnacity of the male Ortygornis gularis, ii. 44;
on the spurs of Galloperdix, ii. 46;
on the habits of Lobivanellus, ii. 48;

Page 352

on the spoonbill, ii. 60;
on the drumming of the Kalij pheasant, ii. 63;
on Indian bustards, ii. 65;
on Otis bengalensis, ii. 69;
on the ear-tufts of Sypheotides auritus, ii. 73;
on the double moults of certain birds, ii. 82;
on the moulting of the honey-suckers, ii. 83;
on the moulting of bustards, plovers, and drongos, ii. 84;
on display in male birds, ii. 86;
on the spring change of colour in some finches, ii. 86;
on the display of the under tail-coverts by the male bulbul, ii. 96;
on the Indian honey-buzzard, ii. 126;
on sexual differences in the colour of the eyes of hornbills, ii. 129;
on the markings of the Tragopan pheasant, ii. 134;
on the nidification of the Orioles, ii. 168;
on the nidification of the hornbills, ii. 169;
on the Sultan yellow-tit, ii. 174;
on Palæornis javanicus, ii. 180;
on the immature plumage of birds, ii. 186 et seq.;
on representative species of birds, ii. 190;
on the habits of Turnix, ii. 202;
on the continued increase of beauty of the peacock, ii. 216;
on coloration in the genus Palæornis, ii. 231.
Jevons, W. S., on the migrations of man, i. 135.
Jews, ancient, use of flint tools by the, i. 183;
uniformity of, in various parts of the world, i. 242;
numerical proportion of male and female births among the, i. 301;
ancient, tattooing practised by, ii. 339.
Johnstone, Lieut., on the Indian elephant, i. 268.
Jollofs, fine appearance of the, ii. 357.
Jones, Albert, proportion of sexes of Lepidoptera, reared by, i. 313.
Juan Fernandez, humming-birds of, ii. 221.
Junonia, sexual differences of colouring in species of, i. 389.
Jupiter, Greek statues of, ii. 350.

K.
Kafir skull, occurrence of the diastema in a, i. 126.
Kafirs, their cruelty to animals, i. 94;
lice of the, i. 220;
colour of the, ii. 347;
engrossment of the handsomest women by the chiefs of the, ii. 369;
marriage-customs of the, ii. 373.
Kalij pheasant, drumming of the male, ii. 62;
young of, ii. 190.
Kallima, resemblance of, to a withered leaf, i. 392.
Kalmucks, aversion of, to hairs on the face, ii. 349;
marriage-customs of the, ii. 373.
Kangaroo, great red, sexual difference in the colour of, ii. 286.
Kant, Imm., on duty, i. 70;
on self-restraint, i. 86;

Page 353

on the number of species of man, i. 226.
Katy-did, stridulation of the, i. 352.
Keller, Dr., on the difficulty of fashioning stone implements, i. 138.
Kestrels, new mates found by, ii. 104.
Kidney, i. 116.
King, W. R., on the vocal organs of Tetrao cupido, ii. 56;
on the drumming of grouse, ii. 63;
on the reindeer, ii. 244;
on the attraction of male deer by the voice of the female, ii. 276.
King and Fitzroy, on the marriage-customs of the Fuegians, ii. 374.
King-crows, nidification of, ii. 167.
Kingfisher, ii. 56;
racket-shaped feathers in the tail of a, ii. 73.
Kingfishers, colours and nidification of the, ii. 171, 173, 176;
immature plumage of the, ii. 188, 190;
young of the, ii. 209.
King Lory, ii. 174;
immature plumage of the, ii. 188.
Kingsley, C., on the sounds produced by Umbrina, ii. 23.
Kirby and Spence, on the courtship of insects, i. 272;
on sexual differences in the length of the snout in curculionidæ, i. 255;
on the elytra of Dytiscus, i. 343;
on peculiarities in the legs of male insects, i. 344;
on the relative size of the sexes in insects, i. 345;
on the luminosity of insects, i. 345;
on the Fulgoridæ, i. 351;
on the habits of Termites, i. 364;
on difference of colour in the sexes of beetles, i. 367;
on the horns of the male lamellicorn beetles, i. 371;
on hornlike processes in male curculionidæ, i. 374;
on the pugnacity of the male stag-beetle, i. 375.
Kite, killed by a game-cock, ii. 44.
Knot, retention of winter plumage by the, ii. 82.
Knox, R., on the semilunar fold, i. 23;
on the occurrence of the supra-condyloid foramen in the humerus of man, i. 28;
on the features of the young Memnon, i. 217.
Koala, length of the cæcum in, i. 27.
Kölreuter, on the sterility of hybrid plants, i. 223.
Kobus ellipsiprymnus, proportion of the sexes in, i. 305.
Koodoo, development of the horns of the, i. 289;
markings of the, ii. 300.
Köppen, F. T., on the migratory locust, i. 352.
Kordofan, protuberances artificially produced in, ii. 339.
Kowalevsky, A., on the affinity of the Ascidia to the Vertebrata, i. 205.
Kowalevsky, W., on the pugnacity of the male Capercailzie, ii. 45;
on the pairing of the Capercailzie, ii. 49.
Krause, on a convoluted body at the extremity of the tail in a Macacus and a cat, i. 30.
Kuppfer, Prof., on the affinity of the Ascidia to the Vertebrata, i. 205.

L.

Page 354

Labidocera Darwinii, prehensile organs of the male, i. 329.
Labrus, splendid colours of the species of, ii. 16.
Labrus mixtus, sexual differences in, ii. 9.
Labrus pavo, ii. 16.
Lacertilia, sexual differences of, ii. 32.
Lafresnaye, M. de, on Birds of Paradise, ii. 78.
Lamarck, on the origin of man, i. 4.
Lamellibranchiata, i. 324.
Lamellicorn beetles, hornlike processes from the head and thorax of, i. 370, 373;
analogy of, to Ruminants, i. 373;
influence of sexual selection on, i. 377.
Lamellicornia, stridulation of, i. 380.
Lamont, Mr., on the tusks of the Walrus, ii. 242;
on the use of its tusks by the Walrus, ii. 257.
Lampornis porphyrurus, colours of the female, ii. 168.
Lancelet, i. 204, 212.
Landois, H., on the production of sound by the Cicadæ, i. 351;
on the stridulating organ of the Crickets, i. 354;
on Decticus, i. 355;
on the stridulating organs of the Acridiidæ, i. 356;
on the presence of rudimentary stridulating organs in some female Orthoptera, i. 359;
on the stridulation of Necrophorus, i. 378;
on the stridulant organ of Cerambyx heros, i. 380;
on the stridulating organs in the Coleoptera, i. 382;
on the ticking of Anobium, i. 385;
on the stridulant organ of Geotrupes, i. 380.
Language an art, i. 55;
articulate, origin of, i. 56;
relation of the progress of, to the development of the brain, i. 57;
effects of inheritance in production of, i. 58;
complex structure of, among barbarous nations, i. 61;
natural selection in, i. 61;
gesture, i. 232;
primeval, i. 235;
of a lost tribe preserved by a parrot, i. 236.
Languages, presence of rudiments in, i. 60;
classification of, i. 60;
variability of, i. 60;
crossing or blending of, i. 60;
complexity of, no test of perfection or proof of special creation, i. 62;
resemblance of, evidence of community of origin, i. 189.
Languages and species, identity of evidence of their gradual development, i. 59.
Lanius, ii. 180;
characters of young, ii. 185.
Lanius rufus, anomalous young of, ii. 211.
Lankester, E. R., on comparative longevity, i. 168, 171;
on the destructive effects of intemperance, i. 173.
Lanugo, of the human fœtus, i. 25; ii. 375.
Lapponian language, highly artificial, i. 61.
Lark, proportion of the sexes in the, i. 307;

Page 355

female, singing of the, ii. 54.
Larks, attracted by a mirror, ii. 112.
Lartet, E., on the size of the brain in mammals, i. 51;
comparison of cranial capacities of skulls of recent and tertiary mammals, i. 146;
on Dryopithecus, i. 199.
Larus, seasonal change of plumage in, ii. 228.
Larva, luminous, of a Brazilian beetle, i. 345.
Larynx, muscles of the, in song-birds, ii. 55.
Lasiocampa quercus, attraction of males by the female, i. 311;
sexual difference of colour in, i. 398.
Latham, R. G., on the migrations of man, i. 136.
Latooka, perforation of the lower lip by the women of, ii. 341.
Laurillard, on the abnormal division of the malar bone in man, i. 124.
Lawrence, W., on the superiority of savages to Europeans in power of sight, i. 118;
on the colour of negro infants, ii. 318;
on the fondness of savages for ornaments, ii. 338;
on beardless races, ii. 349;
on the beauty of the English aristocracy, ii. 357.
Layard, E. L., on an instance of rationality in a Cobra, ii. 30;
on the pugnacity of Gallus Stanleyi, ii. 44.
Laycock, Dr., on vital periodicity, i. 12.
Leaves, decaying, tints of, i. 323.
Lecky, Mr., on the sense of duty, i. 71;
on suicide, i. 94;
on the practice of celibacy, i. 96;
his view of the crimes of savages, i. 97;
on the gradual rise of morality, i. 103.
Leconte, J. L., on the stridulant organ in the Coprini and Dynastini, i. 381.
Lee, H., on the numerical proportion of the sexes in the trout, i. 308.
Leg, calf of the, artificially modified, ii. 340.
Legitimate and illegitimate children, proportion of the sexes in, i. 302.
Legs, variation of the length of the, in man, i. 108;
proportions of, in soldiers and sailors, i. 116;
fore, atrophied in some male butterflies, i. 344;
peculiarities of, in male insects, i. 344.
“Lek” of the black-cock and capercailzie, ii. 100.
Lemoine, Albert, on the origin of language, i. 56.
Lemur macaco, sexual difference of colour in, ii. 290.
Lemuridæ, i. 195;
their origin, i. 213;
position and derivation of the, i. 202;
ears of the, i. 23;
variability of the muscles in the, i. 128.
Lemurs, uterus in the, i. 123;
tailless species of, i. 194.
Leopards, black, ii. 294.
Lepidoptera, i. 386;
numerical proportions of the sexes in the, i. 309;
colouring of, i. 387;
ocellated spots of, ii. 132.

Page 356

Lepidosiren, i. 204, 212.
Lenguas, disfigurement of the ears of the, ii. 341.
Leptorhynchus angustatus, pugnacity of male, i. 375.
Leptura testacea, difference of colour in the sexes of, i. 367.
Lequay, on the occurrence of the supra-condyloid foramen in the humerus of man, i. 29.
Leroy, on the wariness of young foxes in hunting-districts, i. 50;
on the desertion of their young by swallows, i. 84.
Lesse, valley of the, i. 29.
Lesson, on the Birds of Paradise, i. 269, ii. 98;
on the sea-elephant, ii. 278.
Lestis bombylans, difference of the sexes in, i. 366.
Lethrus cephalotes, pugnacity of the males of, i. 371, 376.
Leuckart, R., on the vesicula prostatica, i. 31;
on the influence of the age of parents on the sex of offspring, i. 302.
Levator claviculæ muscle, i. 128.
Libellula depressa, colour of the male, i. 363.
Libellulidæ, relative size of the sexes of, i. 347;
difference in the sexes of, i. 361.
Lice of domestic animals and man, i. 219.
Licentiousness, prevalence of, among savages, i. 96;
a check upon population, i. 134.
Lichtenstein, on Chera progne, ii. 120.
Life, inheritance at corresponding periods of, i. 280, 285.
Light, supposed effects of, i. 116;
influence of, upon the colours of shells, i. 326.
Lilford, Lord, the ruff attracted by bright objects, ii. 111.
Limosa lapponica, ii. 204.
Linaria, ii. 180.
Linaria montana, i. 307.
Linnæus, views of, as to the position of man, i. 190.
Linnet, numerical proportion of the sexes in the, i. 307;
crimson forehead and breast of the, ii. 86;
courtship of the, ii. 94.
Linyphia, i. 337.
Lion, polygamous, i. 268;
mane of the, defensive, ii. 266;
roaring of the, ii. 275.
Lions, stripes of young, ii. 183.
Lips, piercing of the, by savages, ii. 341.
Lithobius, prehensile appendages of the female, i. 340.
Lithosia, coloration in, i. 396.
Littorina littorea, i. 324.
Livingstone, Dr., on the influence of dampness and dryness on the colour of the skin, i. 242;
on the liability of negroes to tropical fevers after residence in a cold climate, i. 243;
on the spur-winged goose, ii. 47;
on weaver-birds, ii. 63;
on an African nightjar, ii. 73, 97;
on the battle-scars of South African male mammals, ii. 239;
on the removal of the upper incisors by the Batokas, ii. 340;
on the perforation of the upper lip by the Makalolo, ii. 342;

Page 357

on the Banyai, ii. 347.
Livonia, numerical proportion of male and female births in, i. 301.
Lizards, relative size of the sexes of, ii. 32;
gular pouches of, ii. 33.
Lloyd, L., on the polygamy of the capercailzie and bustard, i. 269;
on the numerical proportion of the sexes in the capercailzie and blackcock, i. 306;
on the salmon, ii. 5;
on the colours of the sea-scorpion, ii. 9;
on the pugnacity of male grouse, ii. 45;
on the capercailzie and black-cock, ii. 49, 54;
on the call of the capercailzie, ii. 61;
on assemblages of grouse and snipes, ii. 101;
on the pairing of a shield-drake with a common duck, ii. 114;
on the battles of seals, ii. 240;
on the elk, ii. 249.
Lobivanellus, wing-spurs in, ii. 48.
Local influences, effect of, upon stature, i. 114.
Lockwood, Mr., on the development of Hippocampus, i. 210.
Locust, bright-coloured, rejected by lizards and birds, i. 361.
Locust, migratory, i. 352.
Locustidæ, stridulation of the, i. 352, 354;
descent of the, i. 356.
Longicorn beetles, difference of the sexes of, in colour, i. 367;
stridulation of, i. 380.
Lonsdale, Mr., on an example of personal attachment in Helix pomatia, i. 325.
Lophobranchii, marsupial receptacles of the male, ii. 21.
Lophophorus, habits of, ii. 121.
Lophorina atra, sexual difference in coloration of, ii. 226.
Lophornis ornatus, ii. 76.
Lord, J. K., on Salmo lycaodon, ii. 5.
Lory, King, ii. 174;
immature plumage of the, ii. 188.
Love-antics and dances of birds, ii. 68.
Lowne, B. T., on Musca vomitoria, i. 145, 349.
Loxia, characters of young of, ii. 184.
Lubbock, Sir J., on the antiquity of man, i. 3;
on the origin of man, i. 4;
on the mental capacity of savages, i. 34;
on the origin of implements, i. 52;
on the simplification of languages, i. 62;
on the absence of the idea of God among certain races of men, i. 65;
on the origin of the belief in spiritual agencies, i. 66;
on superstitions, i. 69;
on the sense of duty, i. 71;
on the practice of burying the old and sick among the Fijians, i. 77;
non-prevalence of suicide among the lowest barbarians, i. 94;
on the immorality of savages, i. 97;
on Mr. Wallace’s claim to the origination of the idea of natural selection, i. 137;
on the absence of remorse among savages, i. 164;
on the former barbarism of civilised nations, i. 181;

Page 358

on improvements in the arts among savages, i. 182;
on resemblances of the mental characters in different races of men, i. 232;
on the power of counting in primeval man, i. 234;
on the arts practised by savages, i. 234;
on the prehensile organs of the male Labidocera Darwinii, i. 329;
on Chloëon, i. 341;
on Smynthurus luteus, i. 348;
on strife for women among the North American Indians, ii. 324;
on music, ii. 334;
on the ornamental practices of savages, ii. 338;
on the estimation of the beard among the Anglo-Saxons, ii. 349;
on artificial deformation of the skull, ii. 352;
on “communal marriages,” ii. 358, 360;
on exogamy, ii. 360, 364;
on the Veddahs, ii. 363;
on polyandry, ii. 365.
Lucanidæ, variability of the mandibles in the male, i. 376.
Lucanus, large size of males of, i. 347.
Lucanus cervus, numerical proportion of sexes of, i. 313;
weapons of the male, i. 375.
Lucanus elaphus, use of mandibles of, i. 377;
large jaws of male, i. 342.
Lucas, Prosper, on sexual preference in horses and bulls, ii. 272.
Lunar periods, i. 212.
Lund, Dr., on skulls found in Brazilian caves, i. 218.
Lungs, enlargement of, in the Quechua and Aymara Indians, i. 119;
a modified swim-bladder, i. 207;
different capacity of in races of man, i. 216.
Luminosity in insects, i. 345.
Luschka, Prof., on the termination of the coccyx, i. 30.
Lust, instinct of, i. 89.
Luxury, comparatively innocuous, i. 171.
Lycæna, sexual differences of colouring in species of, i. 390.
Lyell, Sir C., on the antiquity of man, i. 3;
on the origin of man, i. 4;
on the parallelism of the development of species and languages, i. 59;
on the extinction of languages, i. 60;
on the Inquisition, i. 178;
on the fossil remains of vertebrata, i. 201;
on the fertility of mulattoes, i. 221.
Lynx, Canadian, throat-ruff of the, ii. 267.
Lyre-bird, assemblies of, ii. 101.

M.
Macacus, ears of, i. 23;
convoluted body in the extremity of the tail of, i. 30;
variability of the tail in species of, i. 150;
whiskers of species of, ii. 283.
Macacus cynomolgus, superciliary ridge of, ii. 318;
beard and whiskers of, becoming white with age, ii. 319.

Page 359

Macacus inornatus, i. 151.
Macacus lasiotus, facial spots of, ii. 308.
Macacus radiatus, i. 192.
Macacus rhesus, sexual difference in the colour of, ii. 293, 310.
Macalister, Prof., on variations of the palmaris accessorius muscle, i. 109;
on muscular abnormalities in man, i. 128, 129;
on the greater variability of the muscles in men than in women, i. 275.
Macaws, Mr. Buxton’s observations on, i. 76;
screams of, ii. 61.
McCann, J., on mental individuality, i. 63.
McClelland, J., on the Indian cyprinidæ, ii. 17.
Macculloch, Col., on an Indian village without any female children, ii. 364.
Macculloch, Dr., on tertian ague in a dog, i. 13.
Macgillivray, W., on the vocal organs of birds, i. 59;
on the Egyptian goose, ii. 48;
on the habits of woodpeckers, ii. 63;
on the habits of the snipe, ii. 64;
on the whitethroat, ii. 69;
on the moulting of the snipes, ii. 82;
on the moulting of the anatidæ, ii. 85;
on the finding of new mates by magpies, ii. 103;
on the pairing of a blackbird and thrush, ii. 113;
on pied ravens, ii. 126;
on the guillemots, ii. 127;
on the colours of the tits, ii. 174;
on the immature plumage of birds, ii. 186 et seqq.
Machetes, sexes and young of, ii. 216.
Machetes pugnax, numerical proportion of the sexes in, i. 306;
supposed to be polygamous, i. 270;
pugnacity of the male, ii. 41;
double moult in, ii. 81.
Mackintosh, on the moral sense, i. 70.
MacLachlan, R., on Apatania muliebris and Boreus hyemalis, i. 314;
on the anal appendages of male insects, i. 342;
on the pairing of dragon-flies, i. 347;
on dragon-flies, i. 362, 363;
on dimorphism in Agrion, i. 363;
on the want of pugnacity in male dragon-flies, i. 364;
on the ghost-moth in the Shetland Islands, i. 402.
McLennan, Mr., on the origin of the belief in spiritual agencies, i. 66;
on the prevalence of licentiousness among savages, i. 96, ii. 358;
on infanticide, i. 134, ii. 363;
on the primitive barbarism of civilised nations, i. 181;
on traces of the custom of the forcible capture of wives, i. 182, ii. 365;
on polyandry, ii. 365.
McNeill, Mr., on the use of the antlers of deer, ii. 252;
on the Scotch deerhound, ii. 261;
on the long hairs of the throat of the stag, ii. 268;
on the bellowing of stags, ii. 274.
Macrorhinus proboscideus, structure of the nose of, ii. 278.

Page 360

Magpie, power of speech of, i. 59;
stealing bright objects, ii. 112;
nuptial assemblies of, ii. 102;
new mates found by, ii. 103;
young of the, ii. 209;
coloration of the, ii. 230.
Magpies, vocal organs of the, ii. 55.
Maillard, M., on the proportion of the sexes in a species of Papilio from Bourbon, i. 310.
Maine, Mr., on the absorption of one tribe by another, i. 159;
on the want of a desire for improvement, i. 166.
Makalolo, perforation of the upper lip by the, ii. 341.
Malar bone, abnormal division of, in man, i. 124.
Malay, Archipelago, marriage-customs of the savages of the, ii. 373.
Malays, line of separation between the Papuans and the, i. 218;
general beardlessness of the, ii. 321;
staining of the teeth among, ii. 339;
aversion of some, to hairs on the face, ii. 349.
Malays and Papuans, contrasted characters of, i. 216.
Male animals, struggles of, for the possession of the females, i. 259, 260;
eagerness of, in courtship, i. 272, 273;
generally more modified than female, i. 272, 275;
differ in the same way from females and young, i. 285.
Male characters, developed in females, i. 280;
transfer of, to female birds, ii. 193.
Male, sedentary, of a hymenopterous parasite, i. 272.
Malefactors, i. 172.
Males, presence of rudimentary female organs in, i. 208.
Males and females, comparative mortality of, while young, i. 264, 276;
comparative numbers of, i. 261, 263.
Malherbe, on the woodpeckers, ii. 174.
Malthus, T., on the rate of increase of population, i. 131, 132, 134.
Maluridæ, nidification of the, ii. 169.
Malurus, young of, ii. 216.
Mammæ, i. 254;
rudimentary, in male mammals, i. 17, 30, 208, 209, 210;
supernumerary, in women, i. 125;
of male human subject, i. 130.
Mammalia, Prof. Owen’s classification of, i. 187;
genealogy of the, i. 203.
Mammals, secondary sexual characters of, ii. 239;
weapons of, ii. 241;
recent and tertiary, comparison of cranial capacity of, i. 146;
relative size of the sexes of, ii. 260;
pursuit of female, by the males, i. 272;
parallelism of, with birds in secondary sexual characters, ii. 297;
voices of, used especially during the breeding season, ii. 331.
Man, variability of, i. 108;
erroneously regarded as more domesticated than other animals, i. 111;
definitive origin of, i. 235;
migrations of, i. 135;

Page 361

wide distribution of, i. 137;
causes of the nakedness of, i. 149;
supposed physical inferiority of, i. 156;
numerical proportions of the sexes in, i. 264;
a member of the Catarrhine group, i. 198;
early progenitors of, i. 206;
secondary sexual characters of, ii. 316;
primeval condition of, ii. 367.
Mandans, correlation of colour and texture of hair in the, i. 248.
Mandible, left, enlarged in the male of Taphroderes distortus, i. 344.
Mandibles, use of the, in Ammophila, i. 342;
large, of Corydalis cornutus, i. 342;
large, of male Lucanus elaphus, i. 342.
Mandrill, number of caudal vertebræ in the, i. 150;
colours of the male, ii. 292, 296, 310.
Mantegazza, Prof., on the ornaments of savages, ii. 338 et seqq.;
on the beardlessness of the New Zealanders, ii. 349;
on the exaggeration of natural characters by man, ii. 351.
Mantell, W., on the engrossment of pretty girls by the New Zealand chiefs, ii. 369.
Mantis, pugnacity of species of, i. 360.
Marcus Aurelius, on the origin of the moral sense, i. 71;
on the influence of habitual thoughts, i. 101.
Mareca penelope, ii. 114.
Marks, retained throughout groups of birds, ii. 131.
Marriage, influence of, upon morals, i. 96;
restraints upon, among savages, i. 133;
influence of, on mortality, i. 175;
development of, ii. 361.
Marriages, communal, ii. 358, 360;
early, i. 174, 175.
Marshall, Mr., on the brain of a Bushwoman, i. 216.
Marsupials, i. 202;
possession of nipples by, i. 209;
their origin from Monotremata, i. 213;
uterus of, i. 122;
development of the nictitating membrane in, i. 23;
abdominal sacks of, i. 254;
relative size of the sexes of, ii. 260;
colours of, ii. 286.
Marsupium, rudimentary, in male marsupials, i. 208.
Martin, W. C. L., on alarm manifested by an orang at the sight of a turtle, i. 43;
on the hair in Hylobates, i. 194;
on a female American deer, ii. 258;
on the voice of Hylobates agilis, ii. 277;
on Semnopithecus nemæus, ii. 312.
Martin, on the beards of the inhabitants of St. Kilda, ii. 321.
Martins deserting their young, i. 84.
Martins, C., on death caused by inflammation of the vermiform appendage, i. 28.
Mastoid processes in man and apes, i. 143.
Maudsley, Dr., on the influence of the sense of smell in man, i. 24;

Page 362

on Laura Bridgman, i. 58;
on the development of the vocal organs, i. 59.
Mayers, W. F., on the domestication of the goldfish in China, ii. 17.
Mayhew, E., on the affection between individuals of different sexes in the dog, ii. 270.
Maynard, C. J., on the sexes of Chrysemys picta, ii. 28.
Meckel, on correlated variation of the muscles of the arm and leg, i. 130.
Medicines, effect produced by, the same in man and in monkeys, i. 12.
Medusæ, bright colours of some, i. 322.
Megalithic structures, prevalence of, i. 233.
Megalophrys montana, sexual differences in, ii. 26, 27.
Megapicus validus, sexual difference of colour in, ii. 174.
Megasoma, large size of males of, i. 347.
Meigs, Dr. A., on variation in the skulls of the natives of America, i. 108.
Meinecke, on the numerical proportion of the sexes in butterflies, i. 309.
Meliphagidæ, Australian, nidification of, ii. 169.
Melita, secondary sexual characters of, i. 331.
Meloë, difference of colour in the sexes of a species of, i. 367.
Memory, manifestations of, in animals, i. 45.
Memnon, young, i. 217.
Mental characters, difference of, in different races of men, i. 216.
Mental faculties, variation of, in the same species, i. 36, 110;
diversity of, in the same race of men, i. 109;
inheritance of, i. 110;
similarity of the, in different races of man, i. 232;
of birds, ii. 108.
Mental powers, difference of, in the two sexes in man, ii. 326.
Menura Alberti, ii. 102;
song of, ii. 55.
Menura superba, ii. 101, 102;
long tails of both sexes of, ii. 164.
Merganser, trachea of the male, ii. 60.
Mergus cucullatus, speculum of, i. 291.
Mergus merganser, young of, ii. 189.
Merganser serrator, male plumage of, ii. 85.
Metallura, splendid tail-feathers of, ii. 152.
Methoca ichneumonides, large male of, i. 347.
Meves, M., on the drumming of the snipe, ii. 63.
Mexicans, civilisation of the, not foreign, i. 183.
Meyer, on a convoluted body at the extremity of the tail in a Macacus and a cat, i. 30.
Meyer, Dr. A., on the copulation of phryganidæ of distinct species, i. 342.
Migrations of man, effects of, i. 135.
Migratory instinct of birds, i. 79;
vanquishing the maternal, i. 83, 90.
Mill, J. S., on the origin of the moral sense, i. 71;
on the “greatest happiness principle,” i. 97;
on the difference of the mental powers in the sexes of man, ii. 328.
Millipedes, i. 339.
Milne-Edwards, H., on the use of the enlarged chela of the male Gelasimus, i. 331.
Milvago leucurus, sexes and young of, ii. 205.
Mimickry, i. 411.

Page 363

Mimus polyglottus, ii. 109.
Mind, difference of, in man and the highest animals, i. 104;
similarity of the, in different races, i. 232.
Minnow, proportion of the sexes in the, i. 308, 309.
Minnows, spawning habits of, ii. 15.
Mirror, larks attracted by, ii. 112.
Mivart, St. George, on the reduction of organs, i. 18;
on the ears of the lemuroidea, i. 23;
on variability of the muscles in lemuroidea, i. 128, 136;
on the caudal vertebræ of monkeys, i. 150;
on the classification of the primates, i. 196;
on the orang and on man, i. 197;
on differences in the lemuroidea, i. 198;
on the crest of the male newt, ii. 24.
Mocking-thrush, partial migration of, ii. 109;
young of the, ii. 219.
Modifications, unserviceable, i. 153.
Moles, numerical proportion of the sexes in, i. 305;
battles of male, ii. 239.
Mollienesia petenensis, sexual difference in, ii. 9.
Mollusca, beautiful colours and shapes of, i. 326;
absence of secondary sexual characters in the, i. 324.
Molluscoida, i. 205, 324.
Monacanthus scopas and M. Peronii, sexual differences in, ii. 12.
Mongolians, perfection of the senses in, i. 119.
Monkey, protecting his keeper from a baboon, i. 78, 87;
bonnet-, i. 192;
rhesus, sexual difference in colour of the, ii. 293, 310;
moustache-, colours of the, ii. 291.
Monkeys, liability of, to the same diseases as man, i. 11;
male, recognition of women by, i. 13;
revenge taken by, i. 40;
maternal affection in, i. 40;
variability of the faculty of attention in, i. 44;
using stones and sticks, i. 51;
imitative faculties of, i. 56;
signal-cries of, i. 57;
sentinels posted by, i. 74;
diversity of the mental faculties in, i. 110;
mutual kindnesses of, i. 75;
hands of the, i. 139, 140;
breaking hard fruits with stones, i. 140;
basal caudal vertebræ of, imbedded in the body, i. 151;
human characters of, i. 191;
gradation of species of, i. 227;
beards of, ii. 283;
ornamental characters of, ii. 306;
analogy of sexual differences of, with those of man, ii. 318;
different degrees of difference in the sexes of, ii. 323;
expression of emotions by, ii. 336;

Page 364

generally monogamous habits of, ii. 361;
polygamous habits of some, ii. 362;
naked surfaces of, ii. 376;
American, manifestation of reason in, i. 47;
American, direction of the hair on the arms of some, i. 192.
Monogamy, not primitive, i. 182.
Monogenists, i. 228.
Mononychus pseudacori, stridulation of, i. 382.
Monotremata, i. 202;
development of the nictitating membrane in, i. 23;
lactiferous glands of, i. 209;
connecting mammals with reptiles, i. 213.
Monstrosities, analogous, in man and lower animals, i. 113;
caused by arrest of development, i. 121;
correlation of, i. 130;
transmission of, i. 224.
Montagu, G., on the habits of the black and red grouse, i. 269;
on the pugnacity of the ruff, ii. 41;
on the singing of birds, ii. 52;
on the double moult of the male pintail, ii. 84.
Monteiro, Mr., on Bucorax abyssinicus, ii. 72.
Montes de Oca, M., on the pugnacity of male Humming-birds, ii. 40.
Monticola cyanea, ii. 172.
Monuments, as traces of extinct tribes, i. 237.
Moose, battles of, ii. 240;
horns of the, an incumbrance, ii. 259.
Moral and instinctive impulses, alliance of, i. 88.
Moral faculties, their influence on natural selection in man, i. 158.
Moral rules, distinction between the higher and lower, i. 100.
Moral sense, origin of the, i. 102;
so-called, derived from the social instincts, i. 97, 98.
Moral tendencies, inheritance of, i. 102.
Morality, supposed to be founded in selfishness, i. 97;
test of, the general welfare of the community, i. 98;
gradual rise of, i. 103;
influence of a high standard of, i. 166.
Morgan, L. H., on the Beaver, i. 37;
on the reasoning powers of the Beaver, i. 46;
on the forcible capture of wives, i. 182;
on the castoreum of the beaver, ii. 279;
marriage unknown in primeval times, ii. 359;
on Polyandry, ii. 365.
Morris, F. O., on hawks feeding an orphan nestling, ii. 107.
Mortality, comparative, of females and males, i. 264, 276, 302.
Morton, on the number of species of man, i. 226.
Moschus moschiferus, odoriferous organs of, ii. 280.
Motacillæ, Indian, young of, ii. 190.
Moths, i. 394;
absence of mouth in some male, i. 254;
apterous female, i. 255;

Page 365

male, prehensile use of the tarsi by, i. 256;
male, attracted by females, i. 311;
coloration of, i. 397;
sexual differences of colour in, i. 398.
Motmot, racket-shaped feathers in the tail of a, ii. 73.
Moult, double, ii. 181;
double annual, in birds, ii. 80.
Moulting of birds, ii. 214.
Moults, partial, ii. 83.
Moustache-monkey, colours of the, ii. 291, 311.
Moustaches, in monkeys, i. 192.
Mud-turtle, long claws of the male, ii. 28.
Mulattoes, persistent fertility of, i. 221;
immunity of, from yellow fever, i. 243.
Mule, sterility and strong vitality of the, i. 221.
Mules, rational, i. 48.
Müller, Ferd., on the Mexicans and Peruvians, i. 183.
Müller, Fritz, on astomatous males of Tanais, i. 255;
on the disappearance of spots and stripes in adult mammals, ii. 305;
on the proportions of the sexes in some crustacea, i. 315;
on secondary sexual characters in various crustaceans, i. 328 et seqq.;
on the luminous larva of a beetle, i. 345;
musical contest between male Cicadæ, i. 351;
on the sexual maturity of young amphipod crustacea, ii. 215.
Müller, J., on the nictitating membrane and semilunar fold, i. 23.
Müller, Max, on the origin of language, i. 56;
struggle for life among the words, &c., of languages, i. 60.
Müller, S., on the Banteng, ii. 290;
on the colours of Semnopithecus chrysomelas, ii. 291.
Muntjac-deer, weapons of the, ii. 257.
Murie, J., on the reduction of organs, i. 18;
on the ears of the Lemuroidea, i. 23;
on variability of the muscles in the Lemuroidea, i. 128, 136;
basal caudal vertebræ of Macacus inornatus imbedded in the body, i. 151;
on differences in the Lemuroidea, i. 198;
on the throat-pouch of the male Bustard, ii. 58;
on the mane of Otaria jubata, ii. 267;
on the suborbital pits of Ruminants, ii. 280;
on the colours of the sexes in Otaria nigrescens, ii. 287.
Murray, A., on the Pediculi of different races of men, i. 219.
Murray, T. A., on the fertility of Australian women with white men, i. 220.
Mus coninga, i. 50.
Mus minutus, sexual difference in the colour of, ii. 286.
Musca vomitoria, i. 145.
Muscicapa grisola, ii. 170.
Muscicapa luctuosa, ii. 170.
Muscicapa ruticilla, breeding in immature plumage, ii. 214.
Muscle, ischio-pubic, i. 127.
Muscles, rudimentary, occurrence of, in man, i. 19;
variability of the, i. 109;

Page 366

effects of use and disuse upon, i. 116;
animal-like abnormalities of, in man, i. 127;
correlated variation of, in the arm and leg, i. 130;
variability of, in the hands and feet, i. 136;
of the jaws, influence of, on the physiognomy of the Apes, i. 144;
habitual spasms of, causing modifications of the facial bones, i. 147;
of the early progenitors of man, i. 206;
greater variability of the, in men than in women, i. 275.
Musculus sternalis, Prof. Turner on the, i. 19.
Music, i. 232;
of birds, ii. 51;
discordant, love of savages for, ii. 67;
different appreciation of, by different peoples, ii. 333;
origin of, ii. 333, 337;
effects of, ii. 335.
Musical cadences, perception of, by animals, ii. 333;
powers of man, ii. 330 et seqq.
Musk-deer, canine teeth of male, ii. 241, 256, 257;
male, odoriferous organs of the, ii. 280;
winter change of the, ii. 299.
Musk-duck, Australian, ii. 38;
large size of male, ii. 43;
of Guiana, pugnacity of the male, ii. 43.
Musk-ox, horns of, ii. 247.
Musk-rat, protective resemblance of the, to a clod of earth, ii. 298.
Musophagæ, colours and nidification of the, ii. 171;
both sexes of, equally brilliant, ii. 177.
Mussels opened by monkeys, i. 140.
Mustela, winter change of two species of, ii. 298.
Mutilations, healing of, i. 13.
Mutilla europæa, stridulation of, i. 366.
Mutillidæ, absence of ocelli in female, i. 341.
Mycetes caraya, polygamous, i. 266;
vocal organs of, ii. 277;
beard of, ii. 283;
sexual differences of colour in, ii. 290;
voice of, ii. 332.
Mycetes seniculus, sexual differences of colour in, ii. 290.
Myriapoda, i. 339.

N.
Nägeli, on the influence of natural selection on plants, i. 152;
on the gradation of species of plants, i. 227.
Nails, coloured yellow or purple in part of Africa, ii. 339.
Naples, greater proportion of female illegitimate children in, i. 301.
Narwhal, tusks of the, ii. 242, 248.
Nasal cavities, large size of, in American aborigines, i. 119.
Nascent organs, i. 18.
Nathusius, H. von, on the improved breeds of pigs, i. 230;
on the breeding of domestic animals, ii. 370.

Page 367

Natural selection, its effects on the early progenitors of man, i. 136;
influence of, on man, i. 151, 154;
limitation of the principle, i. 152;
influence of, on social animals, i. 155;
Mr. Wallace on the limitation of, by the influence of the mental faculties in man, i. 158;
influence of, in the progress of the United States, i. 179.
Natural and sexual selection contrasted, i. 278.
Naulette, jaw from, large size of the canines in, i. 126.
Neanderthal skull, capacity of the, i. 146.
Neck, proportion of, in soldiers and sailors, i. 117.
Necrophorus, stridulation of, i. 378, 382.
Nectarinia, young of, ii. 190.
Nectariniæ, nidification of, ii. 169;
moulting of the, ii. 83.
Negro, resemblance of a, to Europeans, in mental characters, i. 232.
Negro-women, their kindness to Mungo Park, i. 95.
Negroes, character of, i. 216;
lice of, i. 220;
blackness of, i. 224, ii. 381;
variability of, i. 225, 226;
immunity of, from yellow fever, i. 243;
difference of, from Americans, i. 247;
disfigurements of the, ii. 296;
colour of newborn children of, ii. 318;
comparative beardlessness of, ii. 321;
readily become musicians, ii. 334;
appreciation of beauty of their women by, ii. 344, 346;
idea of beauty among, ii. 350;
compression of the nose by some, ii. 352.
Neolithic period, i. 183.
Neomorpha, sexual difference of the beak in, ii. 39.
Nephila, i. 337.
Nests, made by fishes, ii. 19;
decoration of, by Humming-birds, ii. 112.
Neumeister, on a change of colour in pigeons after several moultings, i. 294.
Neuration, difference of, in the two sexes of some butterflies and hymenoptera, i. 345.
Neuroptera, i. 314, 361.
Neurothemis, dimorphism in, i. 363.
New Zealand, expectation by the natives of, of their extinction, i. 240;
practice of tattooing in, ii. 342;
aversion of natives of, to hairs on the face, ii. 349;
pretty girls engrossed by the chiefs in, ii. 369.
Newton, A., on the throat-pouch of the male bustard, ii. 58;
on the difference between the females of two species of Oxynotus, ii. 193;
on the habits of the phalarope, dotterel, and godwit, ii. 204.
Newts, ii. 24.
Nicholson, Dr., on the non-immunity of dark Europeans from yellow fever, i. 245.
Nictitating membrane, i. 23, 207.
Nidification, of fishes, ii. 19;
relation of, to colour, ii. 167, 172;

Page 368

of British birds, ii. 169.
Night-heron, cries of the, ii. 51.
Nightingale, arrival of the male before the female, i. 259;
object of the song of the, ii. 52.
Nightingales, new mates found by, ii. 105.
Nightjar, selection of a mate by the female, ii. 116;
Australian, sexes of, ii. 206;
coloration of the, ii. 226.
Nightjars, noise made by some male, with their wings, ii. 62;
elongated feathers in, ii. 73, 97.
Nilghau, sexual differences of colour in the, ii. 287.
Nilsson, Prof., on the resemblance of stone arrow-heads from various places, i. 233;
on the development of the horns in the reindeer, i. 288.
Nipples, absence of, in Monotremata, i. 209.
Nitzsch, C. L., on the down of birds, ii. 80.
Noctuæ, brightly-coloured beneath, i. 397.
Noctuidæ, coloration of, i. 394.
Nordmann, A., on Tetrao urogalloides, ii. 100.
Nomadic habits, unfavourable to human progress, i. 167.
Norway, numerical proportion of male and female births in, i. 301.
Nose, resemblance of, in man and the apes, i. 192;
piercing and ornamentation of the, ii. 341;
flattening of the, ii. 352;
very flat, not admired in negroes, ii. 350.
Nott and Gliddon, on the features of Rameses II., i. 217;
on the features of Amunoph III., i. 218;
on skulls from Brazilian caves, i. 218;
on the immunity of negroes and mulattoes from yellow fever, i. 243;
on the deformation of the skull among American tribes, ii. 352.
Nudibranch mollusca, bright colours of, i. 326.
Numerals, Roman, i. 182.
Nunemaya, natives of, bearded, ii. 322, 349.

O.
Obedience, value of, i. 162.
Observation, powers of, possessed by birds, ii. 109.
Occupations, sometimes a cause of diminished stature, i. 115;
effect of, upon the proportions of the body, i. 116.
Ocelli, absence of, in female Mutillidæ, i. 341.
Ocelli of birds, formation and variability of the, ii. 132.
Ocelot, sexual differences in the colouring of the, ii. 287.
Ocyphaps lophotes, ii. 96.
Odonata, i. 314.
Odonestis potatoria, sexual difference of colour in, i. 398.
Odour, correlation of, with colour of skin, i. 248;
emitted by snakes in the breeding-season, ii. 30;
of mammals, ii. 278.
Œcanthus nivalis, difference of colour in the sexes of, i. 361.
Oidemia, ii. 226, 227.
Olivier, on sounds produced by Pimelia striata, i. 385.

Page 369

Omaloplia brunnea, stridulation of, i. 381.
Onitis furcifer, processes of anterior femora of the male, and on the head and thorax of the female,
i. 372.
Onthophagus, i. 370.
Onthophagus rangifer, sexual differences of, i. 369;
variation in the horns of the male, i. 370.
Ophidia, sexual differences of, ii. 29.
Opossum, wide range of, in America, i. 219.
Optic nerve, atrophy of the, caused by destruction of the eye, i. 116.
Orang-Outan, ii. 323;
Bischoff on the agreement of the brain of the, with that of man, i. 11;
adult age of the, i. 13;
ears of the, i. 21;
vermiform appendage of, i. 27;
platforms built by the, i. 36;
alarmed at the sight of a turtle, i. 43;
using a stick as a lever, i. 51;
using missiles, i. 52;
using the leaves of the Pandanus as a night covering, i. 53;
hands of the, i. 139;
absence of mastoid processes in the, i. 143;
direction of the hair on the arms of the, i. 192;
its aberrant characters, i. 197;
supposed evolution of the, i. 230;
voice of the, ii. 276;
monogamous habits of the, ii. 361;
male, beard of the, ii. 284.
Oranges, treatment of, by monkeys, i. 139.
Orange-tip butterfly, i. 388, 393, 394.
Orchestia Darwinii, dimorphism of males of, i. 332.
Orchestia Tucuratinga, limbs of, i. 330, 331, 337.
Ordeal, i. 68.
Oreas canna, colours of, ii. 288.
Oreas Derbyanus, colours of, ii. 288, 299.
Organs, prehensile, i. 256;
utilised for new purposes, ii. 335.
Organic scale, von Baer’s definition of progress in, i. 211.
Orioles, nidification of, ii. 167.
Oriolus, species of, breeding in immature plumage, ii. 214, 215.
Oriolus melanocephalus, coloration of the sexes in, ii. 178.
Ornaments, prevalence of similar, i. 233;
fondness of savages for, ii. 338;
of male birds, ii. 50.
Ornamental characters, equal transmission of, to both sexes, in mammals, ii. 297;
of monkeys, ii. 306.
Ornithoptera crœsus, i. 310.
Ornithorhynchus, i. 200;
spur of the male, ii. 242;
reptilian tendency of, i. 204.
Orocetes erythrogastra, young of, ii. 219.

Page 370

Orrony, Grotto of, i. 28.
Orsodacna atra, difference of colour in the sexes of, i. 368.
Orthoptera, i. 352;
metamorphosis of, i. 292;
stridulating, auditory apparatus of, i. 353;
colours of, i. 360;
rudimentary stridulating organs in female, i. 359;
stridulation of the, and Homoptera, discussed, i. 360.
Ortygornis gularis, pugnacity of the male, ii. 44.
Oryctes, stridulation of, i. 381;
sexual differences in the stridulant organs of, i. 383.
Oryx leucoryx, use of the horns of, ii. 251, 263.
Osphranter rufus, sexual difference in the colour of, ii. 286.
Ostrich, African, sexes and incubation of the, ii. 205.
Ostriches, stripes of young, ii. 184.
Otaria jubata, mane of the male, ii. 267.
Otaria nigrescens, difference in the coloration of the sexes of, ii. 287.
Otis bengalensis, love-antics of the male, ii. 68.
Otis tarda, polygamous, i. 269;
throat-pouch of the male, ii. 58.
Ouzel, ring-, colours and nidification of the, ii. 179.
Ouzel, water-, colours and nidification of the, ii. 170.
Ovibos moschatus, horns of, ii. 247.
Ovipositor of insects, i. 254.
Ovis cycloceros, mode of fighting of, ii. 249.
Ovule of man, i. 14.
Owen, Prof., on the Corpora Wolffiana, i. 16;
on the great toe in man, i. 16;
on the nictitating membrane and semilunar fold, i. 23;
on the development of the posterior molars in different races of man, i. 26;
on the length of the cæcum in the Koala, i. 27;
on the coccygeal vertebræ, i. 29;
on rudimentary structures belonging to the reproductive system, i. 31;
on abnormal conditions of the human uterus, i. 123;
on the number of digits in the Ichthyopterygia, i. 125;
on the canine teeth in man, i. 126;
on the walking of the chimpanzee and orang, i. 139;
on the mastoid processes in the higher apes, i. 143;
on the hairiness of elephants in elevated districts, i. 149;
on the caudal vertebræ of monkeys, i. 150;
classification of mammalia, i. 187;
on the hair in monkeys, i. 194;
on the piscine affinities of the Ichthyosaurians, i. 204;
on polygamy and monogamy among the antelopes, i. 267;
on the horns of Antilocapra americana, i. 289;
on the musky odour of crocodiles during the breeding season, ii. 29;
on the scent-glands of snakes, ii. 30;
on the Dugong, Cachalot and Ornithorhynchus, ii. 242;
on the antlers of the red deer, ii. 252;
on the dentition of the camelidæ, ii. 257;

Page 371

on the tusks of the Mammoth, ii. 258;
on the horns of the Irish elk, ii. 259;
on the voice in the giraffe, porcupine, and stag, ii. 274;
on the laryngeal sac of the gorilla and orang, ii. 276;
on the odoriferous glands of mammals, ii. 279, 280;
on the effects of emasculation on the vocal organs of men, ii. 330;
on the voice of Hylobates agilis, ii. 332;
on American monogamous monkeys, ii. 362.
Owls, white, new mates found by, ii. 105.
Oxynotus, difference of the females of two species of, ii. 193.

P.
Pachydermata, i. 268.
Paget, on the abnormal development of hairs in man, i. 25;
on the thickness of the skin on the soles of the feet of infants, i. 118.
Painting, i. 232.
Palæmon, chelæ of a species of, i. 331.
Palæornis, sexual differences of colour in, ii. 231.
Palæornis Javanicus, colour of beak of, ii. 179.
Palæornis rosa, young of, ii. 188.
Palamedea cornuta, spurs on the wings, ii. 47.
Paleolithic period, i. 183.
Palestine, habits of the chaffinch in, i. 307.
Pallas, on the perfection of the senses in the Mongolians, i. 119;
on the want of connexion between climate and the colour of the skin, i. 241;
on the polygamous habits of Antilope Saiga, i. 267;
on the lighter colour of horses and cattle in winter in Siberia, i. 282;
on the tusks of the musk-deer, ii. 256, 258;
on the odoriferous glands of mammals, ii. 279;
on the odoriferous glands of the musk-deer, ii. 280;
on winter changes of colour in mammals, ii. 298;
on the ideal of female beauty in North China, ii. 344.
Palmaris accessorius muscle, variations of the, i. 109.
Pampas, horses of the, i. 236.
Pangenesis, hypothesis of, i. 280, 284.
Panniculus carnosus, i. 19.
Papilio, sexual differences of colouring in species of, i. 389;
proportion of the sexes in North American species of, i. 309;
coloration of the wings in species of, i. 396.
Papilio ascanius, i. 389.
Papilio Sesostris and Childrenæ, variability of, i. 402.
Papilio Turnus, i. 310.
Papilionidæ, variability in the, i. 402.
Papuans, line of separation between the, and the Malays, i. 218;
beards of the, ii. 322;
hair of, ii. 340.
Papuans and Malays, contrast in characters of, i. 216.
Paradise, Birds of, ii. 100, 181;
supposed by Lesson to be polygamous, i. 260;
rattling of their quills by, ii. 61;

Page 372

racket-shaped feathers in, ii. 73;
sexual differences in colour of, ii. 76;
decomposed feathers in, ii. 74, 97;
display of plumage by the male, ii. 88.
Paradisea apoda, barbless feathers in the tail of, ii. 74;
plumage of, ii. 78;
and P. papuana, divergence of the females of, ii. 192.
Paradisea rubra, ii. 75, 78.
Paraguay, Indians of, eradication of eyebrows and eyelashes by, ii. 348.
Parrakeet, Australian, variation in the colour of the thighs of a male, ii. 126.
Parallelism of development of species and languages, i. 59.
Parasites on man and animals, i. 12;
as evidence of specific identity or distinctness, i. 219;
immunity from, correlated with colour, i. 242.
Parental affection, partly a result of natural selection, i. 81.
Parents, age of, influence upon sex of offspring, i. 302.
Parinæ, sexual difference of colour in, ii. 174.
Park, Mungo, negro-women teaching their children to love the truth, i. 95;
his treatment by the negro-women, i. 95, 326;
on negro opinions of the appearance of white men, ii. 346.
Parrot, racket-shaped feathers in the tail of a, ii. 73;
instance of benevolence in a, ii. 109.
Parrots, imitative faculties of, i. 44;
change of colour in, i. 152;
living in triplets, ii. 106;
affection of, ii. 108;
colours of, ii. 223;
sexual differences of colour in, ii. 231;
colours and nidification of the, ii. 171, 174, 176;
immature plumage of the, ii. 188;
musical powers of, ii. 335.
Parthenogenesis in the Tenthredinæ, i. 314;
in Cynipidæ, i. 314;
in crustacea, i. 315.
Partridge, monogamous, i. 269;
proportion of the sexes in the, i. 306;
female, ii. 194.
“Partridge-dances,” ii. 68.
Partridges, living in triplets, ii. 106;
spring coveys of male, ii. 107;
distinguishing persons, ii. 110.
Parus cæruleus, ii. 174.
Passer, sexes and young of, ii. 212.
Passer brachydactylus, ii. 212.
Passer domesticus, ii. 170, 212.
Passer montanus, ii. 170, 212.
Patagonians, self-sacrifice by, i. 88.
Patterson, Mr., on the AGRIONIDÆ, i. 362.
Paulistas of Brazil, i. 225.
Pavo cristatus, i. 290; ii. 136.

Page 373

Pavo muticus, i. 290, ii. 136;
possession of spurs by the female, ii. 46, 162.
Pavo nigripennis, ii. 120.
Payaguas Indians, thin legs and thick arms of the, i. 117.
Payan, Mr., on the proportion of the sexes in sheep, i. 305.
Peacock, polygamous, i. 269;
sexual characters of, i. 290;
pugnacity of the, ii. 46;
rattling of the quills by, ii. 61;
elongated tail-coverts of the, ii. 72, 97;
love of display of the, ii. 68, 87, 135;
ocellated spots of the, ii. 135;
inconvenience of long tail of the, to the female, ii. 154, 164, 165;
continued increase of beauty of the, ii. 216.
Peacock-butterfly, i. 392.
Peafowl, preference of females for a particular male, ii. 120;
first advances made by the female, ii. 120.
Pediculi of domestic animals and man, i. 219.
Pedigree of man, i. 213.
Pedionomus torquatus, sexes of, ii. 201.
Peewit, wing-tubercles of the male, ii. 48.
Pelagic animals, transparency of, i. 323.
Pelecanus erythrorhynchus, horny crest on the beak of the male, during the breeding season, ii. 80.
Pelecanus onocrotalus, spring plumage of, ii. 85.
Pelelé, ii. 341.
Pelican, blind, fed by his companions, i. 77;
young, guided by old birds, i. 77;
pugnacity of the male, ii. 43.
Pelicans, fishing in concert, i. 75.
Pelobius Hermanni, stridulation of, i. 380, 382.
Pelvis, alteration of, to suit the erect attitude of man, i. 143;
differences of the, in the sexes in man, ii. 317.
Penelope nigra, sound produced by the male, ii. 64.
Pennant, on the battles of seals, ii. 240;
on the bladder-nose seal, ii. 278.
Penthe, antennal cushions of the male, i. 343.
Perch, brightness of male, during breeding season, ii. 13.
Peregrine Falcon, new mate found by, ii. 104.
Period of variability, relation of, to sexual selection, i. 296.
Periodicity, vital, Dr. Laycock on, i. 12.
Periods, lunar, followed by functions in man and animals, i. 12, 212.
Periods of life, inheritance at corresponding, i. 280, 285.
Perisoreus canadensis, young of, ii. 209.
Peritrichia, difference of colour in the sexes of a species of, i. 367.
Periwinkle, i. 324.
Pernis cristata, ii. 126.
Perseverance, a characteristic of man, ii. 328.
Persians, said to be improved by intermixture with Georgians and Circassians, ii. 357.
Personnat, M., on Bombyx Yamamai, i. 310.
Peruvians, civilisation of the, not foreign, i. 183.

Page 374

Petrels, colours of, ii. 230.
Petrocincla cyanea, young of, ii. 219.
Petronia, ii. 212.
Pfeiffer, Ida, on Javanese ideas of beauty, ii. 347.
Phacochœrus æthiopicus, tusks and pads of, ii. 265.
Phalanger, Vulpine, black varieties of the, ii. 294.
Phalaropus fulicarius, ii. 203.
Phalaropus hyperboreus, ii. 203.
Phanæus, i. 373.
Phanæus carnifex, variation of the horns of the male, i. 370.
Phanæus faunus, sexual differences of, i. 369.
Phanæus lancifer, i. 370.
Phasgonura viridissima, stridulation of, i. 354, 356.
Phasianus Sœmmerringii, ii. 157.
Phasianus versicolor, ii. 89.
Phasianus Wallichii, ii. 93, 196.
Phasmidæ, mimickry of leaves by the, i. 414.
Pheasant, polygamous, i. 269;
production of hybrids with the common fowl, ii. 122;
and black grouse, hybrids of, ii. 113;
immature plumage of the, ii. 188.
Pheasant, Argus, ii. 72, 181;
display of plumage by the male, ii. 91;
ocellated spots of the, ii. 134, 141;
gradation of characters in the, ii. 141.
Pheasant, Blood-, ii. 46.
Pheasant, Cheer, ii. 93, 195.
Pheasant, Eared, i. 290, ii. 93, 195;
sexes alike in the, ii. 178;
length of the tail in the, ii. 166.
Pheasant, Golden, display of plumage by the male, ii. 89;
sex of young, ascertained by pulling out head-feathers, ii. 214;
age of mature plumage in the, ii. 213.
Pheasant, Kalij, drumming of the male, ii. 62.
Pheasant, Reeve’s, length of the tail in, ii. 166.
Pheasant, Silver, sexual coloration of the, ii. 228;
triumphant male, deposed on account of spoiled plumage, ii. 120.
Pheasant, Sœmmerring’s, ii. 156, 166.
Pheasant, Tragopan, ii. 72;
display of plumage by the male, ii. 91;
markings of the sexes of the, ii. 134.
Pheasants, period of acquisition of male characters in the family of the, i. 290;
proportion of sexes in chicks of, i. 306;
length of the tail in, ii. 156, 164, 166.
Philodromus, i. 337.
Philters, worn by women, ii. 344.
Phoca grœnlandica, sexual difference in the coloration of, ii. 287.
Phœnicura ruticilla, ii. 105.
Phosphorescence of insects, i. 345.
Phryganidæ, copulation of distinct species of, i. 342.

Page 375

Phryniscus nigricans, ii. 25.
Physical inferiority, supposed, of man, i. 156.
Pickering, on the number of species of man, i. 226.
Picton, J. A., on the soul of man, ii. 395.
Picus auratus, ii. 43.
Pieridæ, mimickry by female, i. 413.
Pieris, i. 393.
Pigeon, carrier, late development of the wattle in, i. 293;
domestic, breeds and sub-breeds of, ii. 178;
pouter, late development of the crop in, i. 293;
female, deserting a weakened mate, i. 262.
Pigeons, nestling, fed by the secretion of the crop of both parents, i. 210;
changes of plumage in, i. 281;
transmission of sexual peculiarities in, i. 283;
changing colour after several moultings, i. 294;
numerical proportion of the sexes in, i. 306;
cooing of, ii. 60;
variations in plumage of, ii. 74;
display of plumage by male, ii. 96;
local memory of, ii. 109;
antipathy of female, to certain males, ii. 118;
pairing of, ii. 118, 119;
profligate male and female, ii. 119;
wing-bars and tail-feathers of, ii. 131;
supposititious breed of, ii. 155;
pouter and carrier, peculiarities of predominant in males, ii. 158;
nidification of, ii. 168;
immature plumage of the, ii. 188;
Australian, ii. 175;
Belgian, with black-streaked males, i. 285, 293; ii. 157.
Pigs, origin of the improved breeds of, i. 230;
numerical proportion of the sexes in, i. 305;
stripes of young, ii. 184, 303;
sexual preference shown by, ii. 273.
Pike, American, brilliant colours of the male, during the breeding season, ii. 14.
Pike, male, devoured by females, i. 308.
Pike, L. O., on the psychical elements of religion, i. 68.
Pimelia striata, sounds produced by the female, i. 385.
Pintail Drake, plumage of, ii. 84;
pairing with a wild duck, ii. 115.
Pintail Duck, pairing with a Wigeon, ii. 114.
Pipe-fish, filamentous, ii. 18;
marsupial receptacles of the male, ii. 21.
Pipits, moulting of the, ii. 83.
Pipra, modified secondary wing-feathers of male, ii. 65.
Pipra deliciosa, ii. 65, 66.
Pirates stridulus, stridulation of, i. 350.
Pithecia leucocephala, sexual differences of colour in, ii. 290.
Pithecia Satanas, beard of, ii. 283, 284, 285;
resemblance of, to a negro, ii. 381.

Page 376

Pits, suborbital, of Ruminants, ii. 280.
Pittidæ, nidification of, ii. 167.
Placentata, i. 202.
Plagiostomous fishes, ii. 1.
Plain-wanderer, Australian, ii. 201.
Planariæ, bright colours of some, i. 322.
Plantain-eaters, colours and nidification of the, ii. 171;
both sexes of, equally brilliant, ii. 177.
Plants, cultivated, more fertile than wild, i. 132;
Nägeli, on natural selection in, i. 152;
male flowers of, mature before the female, i. 260;
phenomena of fertilisation in, i. 273;
relation between number and size of seeds in, i. 317.
Platalea, ii. 60;
change of plumage in, ii. 179.
Platyblemnus, i. 361.
Platycercus, young of, ii. 209.
Platyphyllum concavum, i. 352, 356.
Platyrrhine monkeys, i. 196.
Platysma myoides, i. 19.
Plecostomus, head-tentacles of the male of a species of, ii. 10.
Plecostomus barbatus, peculiar beard of the male, ii. 10.
Plectropterus gambensis, spurred wings of, ii. 46.
Ploceus, ii. 54.
Plovers, wing-spurs of, ii. 48;
double moult in, ii. 83.
Plumage, changes of, inheritance of, by fowls, i. 281;
tendency to analogous variation in, ii. 74;
display of, by male birds, ii. 86, 96;
changes of, in relation to season, ii. 180;
immature, of birds, ii. 183, 187;
colour of, in relation to protection, ii. 223.
Plumes on the head in birds, difference of, in the sexes, ii. 164.
Pneumora, structure of, i. 357.
Podica, sexual difference in the colour of the irides of, ii. 128.
Poeppig, on the contact of civilised and savage races, i. 239.
Poison, avoidance of, by animals, i. 49.
Poisonous fruits and herbs avoided by animals, i. 36.
Poisons, immunity from, correlated with colour, i. 242.
Polish fowls, origin of the crest in, i. 284.
Pollen and van Dam, on the colours of Lemur macaco, ii. 290.
Polyandry, ii. 365;
in certain cyprinidæ, i. 309;
among the elateridæ, i. 313.
Polydactylism in man, i. 125.
Polygamy, influence of, upon sexual selection, i. 265;
superinduced by domestication, i. 270;
supposed increase of female births by, i. 303;
in the stickleback, ii. 2.
Polygenists, i. 228.

Page 377

Polynesia, prevalence of infanticide in, ii. 364.
Polynesians, aversion of, to hairs on the face, ii. 349;
wide geographical range of, i. 112;
difference of stature among the, i. 115;
crosses of, i. 225;
variability of, i. 225;
heterogeneity of the, i. 241.
Polyplectron, display of plumage by the male, i. 89;
number of spurs in, ii. 46;
gradation of characters in, ii. 137;
female of, ii. 194.
Polyplectron chinquis, ii. 90, 138, 139.
Polyplectron Hardwickii, ii. 138, 139.
Polyplectron malaccense, ii. 139, 140.
Polyplectron Napoleonis, ii. 138, 140.
Polyzoa, i. 324.
Pontoporeia affinis, i. 329.
Porcupine, mute, except in the rutting season, ii. 274.
Pores, excretory, numerical relation of, to the hairs in sheep, i. 248.
Porpitæ, bright colours of some, i. 322.
Portax picta, dorsal crest and throat-tuft of, ii. 282;
sexual differences of colour in, ii. 287, 299.
Portunus puber, pugnacity of, i. 332.
Potamochoerus penicillatus, tusks and facial knobs of the, ii. 266.
Pouchet, G., on the ratio of instinct and intelligence, i. 37;
on the instincts of ants, i. 187;
on the caves of Abou-Simbel, i. 217;
on the immunity of negroes from yellow fever, i. 243.
Pouter pigeon, late development of the large crop in, i. 293.
Power, Dr., on the different colours of the sexes in a species of Squilla, i. 335.
Powys, Mr., on the habits of the chaffinch in Corfu, i. 307.
Pre-eminence of man, i. 137.
Preference for males by female birds, ii. 113, 122;
shown by mammals, in pairing, ii. 268.
Prehensile organs, i. 256.
Presbytis entellus, fighting of the male, ii. 324.
Preyer, Dr., on supernumerary mammæ in women, i. 125.
Prichard, on the difference of stature among the Polynesians, i. 115;
on the connection between the breadth of the skull in the Mongolians and the perfection of
their senses, i. 119;
on the capacity of British skulls of different ages, i. 146;
on the flattened heads of the Colombian savages, ii. 340;
on Siamese notions of beauty, ii. 345;
on the beardlessness of the Siamese, ii. 349;
on the deformation of the head among American tribes and the natives of Arakhan, ii. 352.
Primary sexual organs, i. 254.
Primates, i. 190;
sexual differences of colour in, ii. 290.
Primogeniture, evils of, i. 170.
Primula, relation between number and size of seeds in, i. 317.

Page 378

Prionidæ, difference of the sexes in colour, i. 367.
Proctotretus multimaculatus, ii. 26, 37.
Proctotretus tenuis, sexual difference in the colour of, ii. 37.
Profligacy, i. 173.
Progenitors, early, of man, i. 206.
Progress, not the normal rule in human society, i. 166;
elements of, i. 177.
Prong-horn, horns of, i. 289.
Proportions, difference of, in distinct races, i. 216.
Protective colouring in butterflies, i. 392;
in lizards, ii. 37;
in birds, ii. 197, 223;
in mammals, ii. 298, 299.
Protective nature of the dull colouring of female Lepidoptera, i. 403, 405, 414.
Protective resemblances in fishes, ii. 18.
Protozoa, absence of secondary sexual characters in, i. 321.
Pruner-Bey, on the occurrence of the supra-condyloid foramen in the humerus of man, i. 29;
on the colour of negro infants, ii. 318.
Prussia, numerical proportion of male and female births in, i. 301.
Psocus, proportions of the sexes in, i. 314.
Ptarmigan, monogamous, i. 269;
summer and winter plumage of the, ii. 81, 83;
nuptial assemblages of, ii. 101;
triple moult of the, ii. 181;
protective coloration of, ii. 198.
Puff-birds, colours and nidification of the, ii. 171.
Pugnacity of fine-plumaged male birds, ii. 93.
Pumas, stripes of young, ii. 183.
Puppies learning from cats to clean their faces, i. 44.
Pycnonotus hæmorrhous, pugnacity of the male, ii. 41;
display of under tail coverts by the male, ii. 96.
Pyranga æstiva, male aiding in incubation, ii. 167.
Pyrodes, difference of the sexes in colour, i. 367.

Q.
Quadrumana, hands of, i. 139;
differences between man and the, i. 190;
dependence of, on climate, i. 218;
sexual differences of colour in, ii. 290;
ornamental characters of, ii. 306;
analogy of sexual differences of, with those of man, ii. 318;
fighting of males for the females, ii. 324;
monogamous habits of, ii. 361;
beards of the, ii. 378.
Quain, R., on the variation of the muscles in man, i. 109.
Quatrefages, A. de, on the occurrence of a rudimentary tail in man, i. 29;
on the moral sense as a distinction between man and animals, i. 70;
on variability, i. 112;
on the fertility of Australian women with white men, i. 221;
on the Paulistas of Brazil, i. 225;

Page 379

on the evolution of the breeds of cattle, i. 230;
on the Jews, i. 242;
on the liability of negroes to tropical fevers after residence in a cold climate, i. 243;
on the difference between field- and house-slaves, i. 246;
on the influence of climate on colour, i. 246;
on the Ainos, ii. 321;
on the women of San-Giuliano, ii. 357.
Quechua Indians, i. 119;
local variation of colour in the, i. 246;
no grey hair among the, ii. 320;
hairlessness of the, ii. 322;
long hair of the, ii. 348.
Querquedula acuta, ii. 114.
Quiscalus major, proportions of the sexes of, in Florida and Honduras, i. 307.

R.
Rabbit, white tail of the, ii. 298.
Rabbits, danger-signals of, i. 74;
domestic, elongation of the skull in, i. 147;
modification of the skull in, by the lopping of the ear, i. 147;
numerical proportion of the sexes in, i. 305.
Races, distinctive characters of, i. 215;
or species of man, i. 217;
crossed, fertility or sterility of, i. 220;
of man, variability of the, i. 225;
of man, resemblance of, in mental characters, i. 232;
formation of, i. 235;
of man, extinction of, i. 236;
effects of the crossing of, i. 240;
of man, formation of the, i. 240;
of man, children of the, ii. 318;
beardless, aversion of, to hairs on the face, ii. 349.
Raffles, Sir S., on the Banteng, ii. 290.
Rafts, use of, i. 137, 234.
Rage, manifested by animals, i. 40.
Raia batis, teeth of, ii. 6.
Raia clavata, female spined on the back, ii. 2;
sexual difference in the teeth of, ii. 6.
Raia maculata, teeth of, ii. 6.
Rails, spur-winged, ii. 48.
Ram, mode of fighting of the, ii. 249;
African, mane of an, ii. 284;
fat-tailed, ii. 284.
Rameses II., i. 217.
Ramsay, Mr., on the Australian Musk-duck, ii. 38;
on the Regent-bird, ii. 113;
on the incubation of Menura superba, ii. 165.
Rana esculenta, vocal sacs of, ii. 28.
Rat, common, general dispersion of, a consequence of superior cunning, i. 50;
supplantation of the native, in New Zealand, by the European rat, i. 240;

Page 380

common, said to be polygamous, i. 268;
numerical proportion of the sexes in, i. 305.
Rats, enticed by essential oils, ii. 281.
Rationality of birds, ii. 108.
Rattle-snakes, difference of the sexes in the, ii. 29;
said to use their rattles as a sexual call, ii. 30.
Raven, vocal organs of the, ii. 55;
stealing bright objects, ii. 112;
pied, of the Feroe Islands, ii. 120.
Rays, prehensile organs of male, ii. 1.
Razor-bill, young of the, ii. 217.
Reade, Winwood, on the Guinea sheep, i. 289;
non-development of horns in castrated male Guinea sheep, ii. 247;
on the occurrence of a mane in an African ram, ii. 285;
on the negroes’ appreciation of the beauty of their women, ii. 344;
on the admiration of negroes for a black skin, ii. 346;
on the idea of beauty among negroes, ii. 350;
on the Jollofs, ii. 357;
on the marriage-customs of the negroes, ii. 374.
Reason, in animals, i. 46.
Redstart, American, breeding in immature plumage, ii. 214.
Redstarts, new mates found by, ii. 105.
Reduvidæ, stridulation of, i. 350.
Reed-bunting, head-feathers of the male, ii. 95;
attacked by a bullfinch, ii. 111.
Reefs, fishes frequenting, ii. 17.
Regeneration, partial, of lost parts in man, i. 13.
Regent-bird, ii. 112.
Reindeer, antlers of, with numerous points, ii. 252;
sexual preferences shown by, ii. 273;
horns of the, i. 288;
winter change of the, ii. 299;
battles of, ii. 240;
horns of the female, ii. 243.
Relationship, terms of, ii. 360.
Religion, deficiency of, among certain races, i. 65;
psychical elements of, i. 68.
Remorse, i. 91;
deficiency of, among savages, i. 164.
Rengger, on the diseases of Cebus Azaræ, i. 11;
on maternal affection in a Cebus, i. 40;
revenge taken by monkeys, i. 40;
on the reasoning powers of American monkeys, i. 47;
on the use of stones by monkeys for cracking hard nuts, i. 51;
on the sounds uttered by Cebus Azaræ, i. 54;
on the signal-cries of monkeys, i. 57;
on the diversity of the mental faculties of monkeys, i. 110;
on the Payaguas Indians, i. 117;
on the inferiority of Europeans to savages in their senses, i. 118;
on the polygamous habits of Mycetes caraya, i. 266;

Page 381

on the voice of the howling monkeys, ii. 277;
on the odour of Cervus campestris, ii. 279;
on the beards of Mycetes caraya and Pithecia Satanas, ii. 283;
on the colours of Felis mitis, ii. 287;
on the colours of Cervus paludosus, ii. 290;
on sexual differences of colour in Mycetes, ii. 291;
on the colour of the infant Guaranys, ii. 318;
on the early maturity of the female of Cebus azaræ, ii. 318;
on the beards of the Guaranys, ii. 322, 323;
on the emotional notes employed by monkeys, ii. 336;
on American polygamous monkeys, ii. 362.
Representative species, of birds, ii. 190, 191.
Reproduction, unity of phenomena of, throughout the mammalia, i. 13;
period of, in birds, ii. 214.
Reproductive system, rudimentary structures in the, i. 30;
accessory parts of, i. 207.
Reptiles, ii. 28.
Reptiles and birds, alliance of, i. 213.
Resemblances, small, between man and the apes, i. 191.
Retrievers, exercise of reasoning faculties by, i. 48.
Revenge, manifested by animals, i. 40.
Reversion, i. 122;
perhaps the cause of some bad dispositions, i. 173.
Rhagium, difference of colour in the sexes of a species of, i. 367.
Ramphastos carinatus, ii. 227.
Rhinoceros, nakedness of, i. 148;
horns of, ii. 247;
horns of, used defensively, ii. 263;
attacking white or grey horses, ii. 295.
Rhynchæa, sexes and young of, ii. 202.
Rhynchæa australis, ii. 203.
Rhynchæa bengalensis, ii. 203.
Rhynchæa capensis, ii. 202.
Rhythm, perception of, by animals, ii. 333.
Richard, M., on rudimentary muscles in man, i. 19.
Richardson, Sir J., on the pairing of Tetrao umbellus, ii. 49;
on Tetrao urophasianus, ii. 58;
on the drumming of grouse, ii. 63;
on the dances of Tetrao phasianellus, ii. 69;
on assemblages of grouse, ii. 101;
on the battles of male deer, ii. 240;
on the reindeer, ii. 244;
on the horns of the musk-ox, ii. 247;
on antlers of the reindeer with numerous points, ii. 252;
on the moose, ii. 259.
Richardson, on the Scotch deerhound, ii. 261.
Richter, Jean Paul, on imagination, i. 45.
Riedel, on profligate female pigeons, ii. 119.
Ring-ouzel, colours and nidification of the, ii. 170.
Ripa, Father, on the difficulty of distinguishing the races of the Chinese, i. 215.

Page 382

Rivalry, in singing, between male birds, ii. 53.
River-hog, African, tusks and knobs of the, ii. 266.
Rivers, analogy of, to islands, i. 204.
Roach, brightness of male during breeding-season, ii. 13.
Robbery, of strangers, considered honourable, i. 94.
Robertson, Mr., remarks on the development of the horns in the roebuck and red-deer, i. 288.
Robin, pugnacity of the male, ii. 40;
autumn song of the, ii. 54;
female, singing of the, ii. 54;
attacking other birds with red in their plumage, ii. 111;
young of the, ii. 208.
Robinet, on the difference of size of the male and female cocoons of the silk-moth, i. 346.
Rodents, uterus in the, i. 123;
absence of secondary sexual characters in, i. 268;
sexual differences in the colours of, ii. 286.
Roe, winter change of the, ii. 299.
Rolle, F., on the origin of man, i. 4;
on a change in German families settled in Georgia, i. 246.
Roller, ii. 56.
Romans, ancient, gladiatorial exhibitions of the, i. 101.
Rook, voice of the, ii. 61.
Rössler, Dr., on the resemblance of the lower surface of butterflies to the bark of trees, i. 392.
Rostrum, sexual difference in the length of, in some weevils, i. 255.
Rudimentary organs, i. 17;
origin of, i. 32.
Rudiments, presence of, in languages, i. 60.
Rudolph, on the want of connexion between climate and the colour of the skin, i. 241.
Ruff, supposed to be polygamous, i. 270;
proportion of the sexes in the, i. 306;
pugnacity of the, ii. 41, 48;
double moult in, ii. 81, 84;
duration of dances of, ii. 100;
attraction of the, to bright objects, ii. 111.
Ruminants, male, disappearance of canine teeth in, i. 144, ii. 325;
generally polygamous, i. 266;
analogy of Lamellicorn beetles to, i. 373;
suborbital pits of, ii. 280;
sexual differences of colour in, ii. 287.
Rupicola crocea, display of plumage by the male, ii. 87.
Rüppell, on canine teeth in deer and antelopes, ii. 258.
Russia, numerical proportion of male and female births in, i. 301.
Ruticilla, ii. 180.
Rütimeyer, Prof., on the physiognomy of the apes, i. 149;
on the sexual differences of monkeys, ii. 323.
Rutlandshire, numerical proportion of male and female births in, i. 301.

S.
Sachs, Prof., on the behaviour of the male and female elements in fertilisation, i. 274.
Sacrifices, Human, i. 182.
Sagittal crest in male apes and Australians, ii. 319.

Page 383

Sahara, birds of the, ii. 172;
animal inhabitants of the, ii. 224.
Sailors, growth of, delayed by conditions of life, i. 114;
long-sighted, i. 118.
Sailors and soldiers, difference in the proportions of, i. 116.
St. John, Mr., on the attachment of mated birds, ii. 108.
St. Kilda, beards of the inhabitants of, ii. 321.
Salmo eriox, and S. umbla, colouring of the male, during the breeding season, ii. 14.
Salmo lycaodon, ii. 4.
Salmo salar, ii. 4.
Salmon, leaping out of fresh water, i. 83;
male, ready to breed before the female, i. 260;
proportion of the sexes in, i. 308;
male, pugnacity of the, ii. 3;
male, characters of, during the breeding season, ii. 3, 14;
spawning of the, ii. 19;
breeding of immature male, ii. 215.
Salvin, O., on the Humming-birds, i. 269, ii. 168;
on the numerical proportion of the sexes in Humming-birds, i. 307, ii. 221;
on Chamæpetes and Penelope, ii. 64;
on Selasphorus platycercus, ii. 65;
on Pipra deliciosa, ii. 66;
on Chasmorhynchus, ii. 79.
Samoa Islands, beardlessness of the natives of, ii. 322, 349.
Sand-skipper, i. 334.
Sandwich Islands, variation in the skulls of the natives of the, i. 108;
superiority of the nobles in the, ii. 356.
Sandwich Islanders, lice of, i. 219.
San-Giuliano, women of, ii. 357.
Santali, recent rapid increase of the, i. 133;
Mr. Hunter on the, i. 241.
Saphirina, characters of the males of, i. 335.
Sarkidiornis melanonotus, characters of the young, ii. 185.
Sars, O., on Pontoporeia offinis, i. 329.
Saturnia carpini, attraction of males by the female, i. 311.
Saturnia Io, difference of coloration in the sexes of, i. 398.
Saturniidæ, coloration of the, i. 396, 398.
Savage, Dr., on the fighting of the male gorillas, ii. 324;
on the habits of the gorilla, ii. 363.
Savage and Wyman, on the polygamous habits of the gorilla, i. 266.
Savages, imitative faculties of, i. 57, 161;
causes of low morality of, i. 97;
uniformity of, exaggerated, i. 111;
long-sighted, i. 118;
rate of increase among, usually small, i. 132;
retention of the prehensile power of the feet by, i. 142;
tribes of, supplanting one another, i. 160;
improvements in the arts among, i. 182;
arts of, i. 234;
fondness of, for rough music, ii. 67;

Page 384

attention paid by, to personal appearance, ii. 338;
relation of the sexes among, ii. 363.
Saw-fly, pugnacity of a male, i. 364.
Saw-flies, proportions of the sexes in, i. 314.
Saxicola rubicola, young of, ii. 220.
Scalp, motion of the, i. 20.
Scent-glands in snakes, ii. 30.
Schaaffhausen, Prof., on the development of the posterior molars in different races of man, i. 26;
on the jaw from La Naulette, i. 126;
on the correlation between muscularity and prominent supra-orbital ridges, i. 130;
on the mastoid processes of man, i. 143;
on modifications of the cranial bones, i. 147;
on human sacrifices, i. 182;
on the probable speedy extermination of the anthropomorphous apes, i. 201;
on the ancient inhabitants of Europe, i. 237;
on the effects of use and disuse of parts, i. 247;
on the superciliary ridge in man, ii. 316;
on the absence of race-differences in the infant skull in man, ii. 318;
on ugliness, ii. 354.
Schaum, H., on the elytra of Dytiscus and Hydroporus, i. 343.
Schelver, on dragon-flies, i. 363.
Schiodte, on the stridulation of Heterocerus, i. 379.
Schlegel, F. von, on the complexity of the languages of uncivilised peoples, i. 61.
Schlegel, Prof., on Tanysiptera, ii. 190.
Schleicher, Prof., on the origin of language, i. 56.
Schleiden, Prof., on the rattle-snake, ii. 30.
Schomburgk, Sir R., on the pugnacity of the male musk-duck of Guiana, ii. 43;
on the courtship of Rupicola crocea, ii. 87.
Schoolcraft, Mr., on the difficulty of fashioning stone implements, i. 138.
Sclater, P. L., on modified secondary wing-feathers in the males of Pipra, ii. 65;
on elongated feathers in nightjars, ii. 73;
on the species of Chasmorhynchus, ii. 79;
on the plumage of Pelecanus onocrotalus, ii. 85;
on the plantain-eaters, ii. 177;
on the sexes and young of Tadorna variegata, ii. 206;
on the colours of Lemur macaco, ii. 290;
on the stripes in asses, ii. 305.
Scolecida, absence of secondary sexual characters in, i. 321.
Scolopax frenata, tail-feathers of, ii. 64.
Scolopax gallinago, drumming of, ii. 63.
Scolopax javensis, tail-feathers of, ii. 64.
Scolopax major, assemblies of, ii. 101.
Scolopax Wilsonii, sound produced by, ii. 64.
Scolytus, stridulation of, i. 379.
Scoter-duck, black, sexual difference in coloration of the, ii. 226;
bright beak of male, ii. 227.
Scott, J., on the colour of the beard in man, ii. 319.
Scrope, on the pugnacity of the male salmon, ii. 3;
on the battles of stags, ii. 240.
Scudder, S. H., imitation of the stridulation of the Orthoptera, i. 353;

Page 385

on the stridulation of the ACRIDIIDÆ, i. 356;
on a Devonian insect, i. 360;
on stridulation, ii. 331.
Sculpture, expression of the ideal of beauty by, ii. 350.
Sea-anemonies, bright colours of, i. 322.
Sea-bear, polygamous, i. 268.
Sea-elephant, male, structure of the nose of the, ii. 278;
polygamous, i. 268.
Sea-lion, polygamous, i. 268.
Seal, bladder-nose, ii. 278.
Seals, their sentinels generally females, i. 74;
evidence furnished by, on classification, i. 190;
sexual differences in the coloration of, ii. 287;
appreciation of music by, ii. 333;
battles of male, ii. 240;
canine teeth of male, ii. 241;
polygamous habits of, i. 268;
pairing of, ii. 269;
sexual peculiarities of, ii. 277.
Sea-scorpion, sexual differences in, ii. 9.
Season, changes of colour in birds, in accordance with the, ii. 80;
changes of plumage of birds in relation to, ii. 180.
Seasons, inheritance at corresponding, i. 282.
Sebituani, ii. 340.
Sebright Bantam, i. 294.
Secondary sexual characters, i. 253;
relations of polygamy to, i. 266;
gradation of, in birds, ii. 135;
transmitted through both sexes, i. 279.
Sedgwick, W., on hereditary tendency to produce twins, i. 133.
Seemann, Dr., on the different appreciation of music by different peoples, ii. 334;
on the effects of music, ii. 335.
Selasphorus platycercus, acuminate first primary of the male, ii. 65.
Selby, P. J., on the habits of the black and red grouse, i. 269.
Selection, double, i. 276.
Selection of male by female birds, ii. 99, 122.
Selection, methodical, of Prussian grenadiers, i. 112.
Selection, sexual, influence of, on the colouring of Lepidoptera, i. 403;
explanation of, i. 256, 260, 271.
Selection, sexual and natural, contrasted, i. 278.
Self-command, habit of, inherited, i. 92;
estimation of, i. 95.
Self-consciousness, i. 62.
Self-preservation, instinct of, i. 89.
Self-sacrifice, by savages, i. 88;
estimation of, i. 95.
Semilunar fold, i. 23.
Semnopithecus, i. 197;
long hair on the heads of species of, i. 192; ii. 380.
Semnopithecus chrysomelas, sexual differences of colour in ii. 291.

Page 386

Semnopithecus comatus, ornamental hair on the head of, ii. 307.
Semnopithecus frontatus, beard, &c., of, ii. 308.
Semnopithecus nasica, nose of, i. 192.
Semnopithecus nemæus, colouring of, ii. 310.
Semnopithecus rubicundus, ornamental hair on the head of, ii. 306.
Senses, inferiority of Europeans to savages in the, i. 118.
Sentinels, i. 74, 82.
Serpents, instinctively dreaded by apes and monkeys, i. 37, 42.
Serranus, hermaphroditism in, i. 208.
Sex, inheritance limited by, i. 282.
Sexes, relative proportions of, in man, i. 300, ii. 320;
probable relation of the, in primeval man, ii. 362.
Sexual characters, secondary, i. 253;
relations of polygamy to, i. 266;
transmitted through both sexes, i. 279; gradation of, in birds, ii. 135.
Sexual and natural selection, contrasted, i. 278.
Sexual characters, effects of the loss of, i. 284;
limitation of, i. 284.
Sexual differences in man, i. 14.
Sexual selection, explanation of, i. 256, 260, 271;
influence of, on the colouring of Lepidoptera, i. 403;
action of, in mankind, ii. 368.
Sexual similarity, i. 277.
Sharks, prehensile organs of male, ii. 1.
Sharpe, R. B., on Tanysiptera sylvia, ii. 165;
on Ceryle, ii. 173;
on the young male of Dacelo Gaudichaudi, ii. 188.
Shaw, Mr., on the pugnacity of the male salmon, ii. 3.
Shaw, J., on the decorations of birds, ii. 71.
Sheep, danger-signals of, i. 74;
sexual differences in the horns of, i. 283;
horns of, i. 289, ii. 246, 259;
domestic, sexual differences of, late developed, i. 293;
numerical proportion of the sexes in, i. 304;
mode of fighting of, ii. 249;
arched foreheads of some, ii. 284.
Sheep, Merino, loss of horns in females of, i. 284;
horns of, i. 289.
Shells, difference in form of, in male and female Gasteropoda, i. 324;
beautiful colours and shapes of, i. 326.
Shield-drake, pairing with a common duck, ii. 114;
New Zealand, sexes and young of, ii. 206.
Shooter, J., on the Kafirs, ii. 347;
on the marriage-customs of the Kafirs, ii. 373.
Shrew-mice, odour of, ii. 279.
Shrike, Drongo, ii. 179.
Shrikes, characters of young, ii. 185.
Shuckard, W. E., on sexual differences in the wings of Hymenoptera, i. 345.
Shyness of adorned male birds, ii. 97.
Siagonium, proportions of the sexes in, i. 314;

Page 387

dimorphism in males of, i. 374.
Siam, proportion of male and female births in, i. 303.
Siamese, general beardlessness of the, ii. 321;
notions of beauty of the, ii. 345;
hairy family of, ii. 378.
Siebold, C. T. von, on the auditory apparatus of the stridulant orthoptera, i. 353.
Sight, inheritance of long and short, i. 118.
Signal-cries of monkeys, i. 57.
Silk-moth, difference of size of the male and female cocoons of the, i. 346;
pairing of the, i. 401;
male, fertilising two or three females, i. 406;
proportion of the sexes in, i. 309, 311;
Ailanthus, Prof. Canestrini, on the destruction of its larvæ by wasps, i. 311.
Simiadæ, i. 195;
their origin and divisions, i. 213.
Similarity, sexual, i. 277.
Singing of the Cicadæ and Fulgoridæ, i. 351;
of tree-frogs, ii. 27;
of birds, object of the, ii. 52.
Sirenia, nakedness of, i. 148.
Sirex juvencus, i. 365.
Siricidæ, difference of the sexes in, i. 365.
Siskin, ii. 85;
pairing with a canary, ii. 115.
Sitana, throat-pouch of the males of, ii. 33, 36.
Size, relative, of the sexes of insects, i. 345.
Skin, movement of the, i. 19;
nakedness of, in man, i. 148;
colour of the, i. 241.
Skin and hair, correlation of colour of, i. 248.
Skull, variation of, in man, i. 108;
cubic contents of, no absolute test of intellect, i. 145;
Neanderthal, capacity of the, i. 146;
causes of modification of the, i. 147;
difference of, in form and capacity, in different races of men, i. 216;
variability of the shape of the, i. 226;
differences of, in the sexes in man, ii. 317;
artificial modifications of the shape of, ii. 340.
Skunk, odour emitted by the, ii. 279.
Slavery, prevalence of, i. 94;
of women, ii. 366.
Slaves, difference between those of field and house, i. 246.
Smell, sense of, in man and animals, i. 23.
Smith, Adam, on the basis of sympathy, i. 82.
Smith, Sir A., on the recognition of women by male Cynocephali, i. 13;
on an instance of memory in a baboon, i. 45;
on the retention of their colour by the Dutch in South Africa, i. 242;
on the polygamy of the South African antelopes, i. 267;
on the proportion of the sexes in Kobus ellipsiprymnus, i. 305;
on Bucephalus capensis, ii. 29;

Page 388

on South African lizards, ii. 37;
on fighting gnus, ii. 240;
on the horns of rhinoceroses, ii. 248;
on the fighting of lions, ii. 266;
on the colours of the Cape Eland, ii. 288;
on the colours of the gnu, ii. 289;
on Hottentot notions of beauty, ii. 345.
Smith, F., on the Cynipidæ and Tenthredinidæ, i. 314;
on the relative size of the sexes of Aculeate Hymenoptera, i. 347;
on the difference between the sexes of ants and bees, i. 365;
on the stridulation of Trox sabulosus, i. 380;
on the stridulation of Mononychus pseudacori, i. 382.
Smynthurus luteus, courtship of, i. 348.
Snakes, sexual differences of, ii. 29;
male, ardency of, ii. 30.
“Snarling muscles,” i. 127.
Snipe, drumming of the, ii. 63;
coloration of the, ii. 226.
Snipe, painted, sexes and young of, ii. 202.
Snipe, solitary, assemblies of, ii. 101.
Snipes, arrival of male before the female, i. 260;
pugnacity of male, ii. 43;
double moult in, ii. 80.
Snow-goose, whiteness of the, ii. 228.
Social animals, affection of, for each other, i. 76;
defence of, by the males, i. 83.
Sociability, the sense of duty connected with, i. 71;
impulse to, in animals, i. 79, 80;
manifestations of, in man, i. 84;
instinct of, in animals, i. 86.
Sociality, probable, of primeval men, i. 155;
influence of, on the development of the intellectual faculties, i. 160;
origin of, in man, i. 161.
Soldiers, American, measurements of, i. 114.
Soldiers and sailors, difference in the proportions of, i. 116.
Solenostoma, bright colours and marsupial sack of the females of, ii. 22.
Song of male birds appreciated by their females, i. 63;
want of, in brilliant plumaged birds, ii. 94;
of birds, ii. 163.
Sorex, odour of, ii. 279.
Sounds admired alike by man and animals, i. 64;
produced by fishes, ii. 23;
produced by male frogs and toads, ii. 27;
instrumentally produced by birds, ii. 63 et seqq.
Spain, decadence of, i. 178.
Sparassus smaragdulus, difference of colour in the sexes of, i. 337, 338.
Sparrow, pugnacity of the male, ii. 40;
acquisition of the Linnet’s song by a, ii. 55;
coloration of the, ii. 198;
immature plumage of the, ii. 188.

Page 389

Sparrow, white-crowned, young of the, ii. 217.
Sparrows, house- and tree-, ii. 170.
Sparrows, new mates found by, ii. 105.
Sparrows, sexes and young of, ii. 212;
learning to sing, ii. 334.
Spathura Underwoodi, ii. 77.
Spawning of fishes, ii. 15, 19.
Spear, origin of the, i. 234.
Species, causes of the advancement of, i. 172;
distinctive characters of, i. 214;
or races of man, i. 217;
sterility and fertility of, when crossed, i. 122;
supposed, of man, i. 226;
gradation of, i. 227;
difficulty of defining, i. 228;
representative, of birds, ii. 190, 191;
of birds, comparative differences between the sexes of distinct, ii. 192.
Spectre-insects, mimickry of leaves by, i. 414.
Spectrum femoratum, difference of colour in the sexes of, i. 361.
Speech, connection between the brain and the faculty of, i. 58.
“Spel” of the black-cock, ii. 60.
Spencer, Herbert, on the dawn of intelligence, i. 37;
on the origin of the belief in spiritual agencies, i. 66;
on the origin of the moral sense, i. 101;
on the influence of food on the size of the jaws, i. 118;
on the ratio between individuation and genesis, i. 318;
on music, ii. 336.
Sperm-whales, battles of male, ii. 240.
Sphingidæ, coloration of the, i. 396.
Sphinx, Humming-bird, i. 399.
Sphinx, Mr. Bates on the caterpillar of a, i. 416.
Spiders, i. 337;
male, more active than female, i. 272;
proportion of the sexes in, i. 314;
male, small size of, i. 338.
Spilosoma menthrasti, rejected by turkeys, i. 398.
Spine, alteration of, to suit the erect attitude of man, i. 143.
Spirits, fondness of monkeys for, i. 12.
Spiritual agencies, belief in, almost universal, i. 65.
Spoonbill, ii. 60;
Chinese, change of plumage in, ii. 179.
Spots, retained throughout groups of birds, i. 131;
disappearance of, in adult mammals, ii. 303.
Sprengel, C. K., on the sexuality of plants, i. 260.
Spring-boc, horns of the, ii. 251.
Sproat, Mr., on the extinction of savages in Vancouver Island, i. 239;
on the eradication of facial hair by the natives of Vancouver Island, ii. 348;
on the eradication of the beard by the Indians of Vancouver Island, ii. 380.
Spurs, occurrence of, in female fowls, i. 280, 284;
development of, in various species of Phasianidæ, i. 290;

Page 390

of Gallinaceous birds, ii. 44, 46;
development of, in female Gallinaceæ, ii. 162.
Squilla, different colours of the sexes of a species of, i. 335.
Squirrels, battles of male, ii. 239;
African, sexual differences in the colouring of, ii. 286;
black, ii. 294.
Stag, long hairs of the throat of, ii. 268;
horns of the, i. 279, 282;
battles of, ii. 240;
horns of the, with numerous branches, ii. 252;
bellowing of the, ii. 274;
crest of the, ii. 282.
Stag-beetle, large size of male, i. 347;
weapons of the male, i. 375;
numerical proportion of sexes of, i. 313.
Stainton, H. T., on the numerical proportion of the sexes in the smaller moths, i. 310;
habits of Elachista rufocinerea, i. 311;
on the coloration of moths, i. 397;
on the rejection of Spilosoma menthrasti, by turkeys, i. 398;
on the sexes of Agrotis exclamationis, i. 399.
Stallion, mane of the, ii. 268.
Stallions, two, attacking a third, i. 75;
fighting, ii. 241;
small canine teeth of, ii. 258.
Stansbury, Capt., observations on pelicans, i. 77.
Staphylinidæ, hornlike processes in male, i. 374.
Starfishes, bright colours of some, i. 322.
Stark, Dr., on the death-rate in towns and rural districts, i. 175;
on the influence of marriage on mortality, i. 176;
on the higher mortality of males in Scotland, i. 302.
Starling, American field, pugnacity of male, ii. 51.
Starling, red-winged, selection of a mate by the female, ii. 116.
Starlings, three, frequenting the same nest, i. 269, ii. 106;
new mates found by, ii. 105.
Statues, Greek, Egyptian, Assyrian, &c., contrasted, ii. 350.
Stature, dependence of, upon local influences, i. 114.
Staudinger, Dr., his list of Lepidoptera, i. 312;
on breeding Lepidoptera, i. 311.
Staunton, Sir G., hatred of indecency a modern virtue, i. 96.
Stealing of bright objects by birds, ii. 112.
Stebbing, T. R., on the nakedness of the human body, ii. 375.
Stemmatopus, ii. 278.
Stenobothrus pratorum, stridulating organs of, i. 357.
Sterility, general, of sole daughters, i. 170;
when crossed, a distinctive character of species, i. 214.
Sterna, seasonal change of plumage in, ii. 228.
Stickle-back, polygamous, i. 271;
male, courtship of the, ii. 2;
male, brilliant colouring of, during the breeding season, ii. 14;
nidification of the, ii. 20.

Page 391

Sticks used as implements and weapons by monkeys, i. 51.
Sting in bees, i. 254.
Stokes, Capt., on the habits of the great Bower-bird, ii. 70.
Stonechat, young of the, ii. 220.
Stone implements, difficulty of making, i. 138;
as traces of extinct tribes, i. 237.
Stones, used by monkeys for breaking hard fruits and as missiles, i. 140;
piles of, i. 233.
Stork, black, sexual differences in the bronchi of the, ii. 60;
red beak of the, ii. 227.
Storks, ii. 226, 230;
sexual difference in the colour of the eyes of, ii. 128.
Strange, Mr., on the Satin Bower-bird, ii. 69.
Stretch, Mr., on the numerical proportion in the sexes of chickens, i. 306.
Strepsiceros kudu, horns of, ii. 255;
markings of, ii. 300.
Stridulation, by males of Theridion, i. 339;
of the Orthoptera and Homoptera discussed, i. 360;
of beetles, i. 378.
Stripes, retained throughout groups of birds, ii. 131;
disappearance of, in adult mammals, ii. 303.
Strix flammea, ii. 105.
Structure, existence of unserviceable modifications of, i. 153.
Struggle for existence, in man, i. 180, 185.
Struthers, Dr., on the occurrence of the supra-condyloid foramen in the humerus of man, i. 28.
Sturnella ludoviciana, pugnacity of the male, ii. 51.
Sturnus vulgaris, ii. 105.
Sub-species, i. 227.
Suffering, in strangers, indifference of savages to, i. 94.
Suicide, i. 172;
formerly not regarded as a crime, i. 94;
rarely practised among the lowest savages, i. 94.
Suidæ, stripes of young, ii. 184.
Sumatra, compression of the nose by the Malays of, ii. 352.
Sumner, Archb., man alone capable of progressive improvement, i. 49.
Sun-birds, nidification of, ii. 169.
Superstitions, i. 182;
prevalence of, i. 99.
Superstitious customs, i. 68.
Superciliary ridge in man, ii. 316, 318.
Supernumerary digits, more frequent in men than in women, i. 276;
inheritance of, i. 285;
early development of, i. 292.
Supra-condyloid foramen in the early progenitors of man, i. 206.
Suspicion, prevalence of, among animals, i. 39.
Sulivan, Sir B. J., on two stallions attacking a third, ii. 241.
Swallow-tail Butterfly, i. 393.
Swallows deserting their young, i. 84, 90.
Swan, black, red beak of the, ii. 227;
black-necked, ii. 230;

Page 392

white, young of, ii. 211;
wild, trachea of the, ii. 59.
Swans, ii. 226, 230;
young, ii. 208.
Swaysland, Mr., on the arrival of migratory birds, i. 259.
Swinhoe, R., on the common rat in Formosa and China, i. 50;
on the sounds produced by the male Hoopoe, ii. 62;
on Dicrurus macrocercus and the Spoonbill, ii. 179;
on the young of Ardeola, ii. 190;
on the habits of Turnix, ii. 202;
on the habits of Rhynchæa bengalensis, ii. 203;
on Orioles breeding in immature plumage, ii. 214, 215.
Sylvia atricapilla, young of, ii. 219.
Sylvia cinerea, aerial love-dance of the male, ii. 68.
Sympathy, i. 168;
among animals, i. 77;
its supposed basis, i. 82.
Sympathies, gradual widening of, i. 100.
Syngnathous fishes, abdominal pouch in male, i. 210.
Sypheotides auritus, acuminated primaries of the male, ii. 64;
ear-tufts of, ii. 73.

T.
Tabanidæ, habits of, i. 254.
Tadorna variegata, sexes and young of, ii. 206.
Tadorna vulpanser, ii. 114.
Tahitians, i. 183;
compression of the nose by the, ii. 352.
Tail, rudimentary, occurrence of, in man, i. 29;
convoluted body in the extremity of the, i. 30;
absence of, in man and the higher apes, i. 150, 194;
variability of, in species of Macacus and in baboons, i. 150;
presence of, in the early progenitors of man, i. 206;
length of, in pheasants, ii. 156, 164, 166;
difference of length of the, in the two sexes of birds, ii. 164.
Tait, Lawson, on the effects of natural selection on civilised nations, i. 168.
Tanager, scarlet, variation in the male, ii. 126.
Tanagra æstiva, ii. 180;
age of mature plumage in, ii. 213.
Tanagra rubra, ii. 126;
young of, ii. 220.
Tanais, absence of mouth in the males of some species of, i. 255;
relations of the sexes in, i. 315;
dimorphic males of a species of, i. 328.
Tankerville, Earl, on the battles of wild bulls, ii. 240.
Tanysiptera, races of, determined from adult males, ii. 190.
Tanysiptera sylvia, long tail-feathers of, ii. 165.
Taphroderes distortus, enlarged left mandible of the male, i. 344.
Tapirs, longitudinal stripes of young, ii. 184, 303.
Tarsi, dilatation of front, in male beetles, i. 343.

Page 393

Tarsius, i. 200.
Tasmania, half-castes killed by the natives of, i. 220.
Tattooing, i. 232;
universality of, ii. 339.
Taste, in the Quadrumana, ii. 296.
Taylor, G. on Quiscalus major, i. 307.
Tea, fondness of monkeys for, i. 12.
Tear-sacks, of Ruminants, ii. 280.
Teebay, Mr., on changes of plumage in spangled Hamburgh fowls, i. 281.
Teeth, rudimentary incisor, in Ruminants, i. 17;
posterior molar, in man, i. 25;
wisdom, i. 26;
diversity of, i. 108;
canine, in the early progenitors of man, i. 206;
canine, of male mammals, ii. 241;
in man, reduced by correlation, ii. 325;
staining of the, ii. 339;
front, knocked out or filed by some savages, ii. 340.
Tegetmeier, Mr., on the abundance of male pigeons, i. 306;
on the wattles of game-cocks, ii. 98;
on the courtship of fowls, ii. 117;
on dyed pigeons, ii. 118.
Tembeta, ii. 341.
Temper, in dogs and horses, inherited, i. 40.
Tench, proportions of the sexes in the, i. 308, 309;
brightness of male, during breeding season, ii. 13.
Tenebrionidæ, stridulation of, i. 379.
Tennent, Sir J. E., on the tusks of the Ceylon Elephant, ii. 248, 258;
on the frequent absence of beard in the natives of Ceylon, ii. 321;
on the Chinese opinion of the aspect of the Cingalese, ii. 345.
Tennyson, A., on the control of thought, i. 101.
Tenthredinidæ, proportions of the sexes in, i. 314;
fighting habits of male, i. 364;
difference of the sexes in, i. 365.
Tephrodornis, young of, ii. 190.
Terai, i. 237.
Termites, habits of, i. 364.
Terns, white, ii. 228;
and black, ii. 230.
Terns, seasonal change of plumage in, ii. 228.
Terror, common action of, upon the lower animals and man, i. 39.
Testudo nigra, ii. 28.
Tetrao cupido, battles of, ii. 50;
sexual difference in the vocal organs of, ii. 56.
Tetrao phasianellus, dances of, ii. 68;
duration of dances of, ii. 100.
Tetrao scoticus, ii. 170, 185, 194.
Tetrao tetrix, ii. 170, 185, 194;
pugnacity of the male. ii. 45.
Tetrao umbellus, pairing of, ii. 49;

Page 394

battles of, ii. 50;
drumming of the male, ii. 61.
Tetrao urogalloides, dances of, ii. 100.
Tetrao urogallus, pugnacity of the male, ii. 45.
Tetrao urophasianus, inflation of the œsophagus in the male, ii. 57.
Thamnobia, young of, ii. 190.
Thaumalea picta, display of plumage by the male, ii. 89.
Thecla, sexual differences of colouring in species of, i. 389.
Thecla rubi, protective colouring of, i. 392.
Theridion, i. 337;
stridulation of males of, i. 339.
Theridion lineatum, variability of, i. 338.
Thomisus citreus, and T. floricolens, difference of colour in the sexes of, i. 337.
Thompson, J. H., on the battles of sperm-whales, ii. 240.
Thompson, W., on the colouring of the male char during the breeding season, ii. 14;
on the finding of new mates by magpies, ii. 103;
on the finding of new mates by Peregrine falcons, ii. 104.
Thorax, processes of, in male beetles, i. 370.
Thorell, T., on the proportion of the sexes in spiders, i. 315.
Thornback, difference in the teeth of the two sexes of the, ii. 6.
Thoughts, control of, i. 101.
Thrush, pairing with a blackbird, ii. 113;
colours and nidification of the, ii. 170.
Thrushes, characters of young, ii. 185, 269.
Thug, his regrets, i. 94.
Thumb, absence of, in Ateles and Hylobates, i. 140.
Thury, M., on the numerical proportion of male and female births among the Jews, i. 301.
Thylacinus, possession of the marsupial sack by the male, i. 208.
Thysanura, i. 348.
Tibia, dilated, of the male Crabro cribrarius, i. 343.
Tibia and femur, proportions of, in the Aymara Indians, i. 119.
Tierra del Fuego, marriage-customs of, ii. 373.
Tiger, colours and markings of the, ii. 302.
Tigers, depopulation of districts by, in India, i. 134.
Tillus elongatus, difference of colour in the sexes of, i. 368.
Timidity, variability of, in the same species, i. 40.
Tineina, proportion of the sexes in, i. 310.
Tipula, pugnacity of male, i. 349.
Tits, sexual difference of colour in, ii. 174.
Toads, ii. 25;
male, treatment of ova by some, i. 210;
male, ready to breed before the female, i. 260.
Toe, great, condition of, in the human embryo, i. 17.
Tomtit, blue, sexual difference of colour in the, ii. 174.
Tonga Islands, beardlessness of the natives of, ii. 322, 349.
Tooke, Horne, on language, i. 55.
Tools, flint, i. 183;
used by monkeys, i. 51;
use of, i. 137.
Top-knots in birds, ii. 74.

Page 395

Tomicus villosus, proportion of the sexes in, i. 314.
Tortoise, voice of the male, ii. 331.
Tortures, submitted to by American savages, i. 95.
Totanus, double moult in, ii. 81.
Toucans, colours and nidification of the, ii. 171;
beaks and ceres of the, ii. 227.
Towns, residence in, a cause of diminished stature, i. 115.
Toynbee, J., on the external shell of the ear in man, i. 21.
Trachea, convoluted and imbedded in the sternum, in some birds, ii. 59;
structure of the, in Rhynchæa, ii. 203.
Trades, affecting the form of the skull, i. 147.
Tragelaphus, sexual differences of colour in, ii. 288.
Tragelaphus scriptus, dorsal crest of, ii. 282;
markings of, ii. 299, 300.
Tragopan, i. 270;
swelling of the wattles of the male, during courtship, ii. 72;
display of plumage by the male, ii. 91;
markings of the sexes of the, ii. 134.
Tragops dispar, sexual difference in the colour of, ii. 30.
Training, effect of, on the mental difference between the sexes of man, ii. 329.
Transfer of male characters to female birds, ii. 193.
Transmission, equal, of ornamental characters, to both sexes in mammals, ii. 297.
Traps, avoidance of, by animals, i. 49;
use of, i. 137.
Treachery, to comrades, avoidance of, by savages, i. 88.
Tremex columbæ, i. 365.
Tribes, extinct, i. 160;
extinction of, i. 236.
Trichius, difference of colour in the sexes of a species of, i. 368.
Trimen, R., on the proportion of the sexes in South African butterflies, i. 310;
on the attraction of males by the female of Lasiocampa quercus, i. 312;
on Pneumora, i. 358;
on difference of colour in the sexes of beetles, i. 367;
on moths brilliantly coloured beneath, i. 397;
on mimickry in butterflies, i. 412;
on Gynanisa Isis, and on the ocellated spots of Lepidoptera, ii. 132;
on Cyllo Leda, ii. 133.
Tringa, sexes and young of, ii. 216.
Tringa canutus, ii. 82.
Triphæna, coloration of the species of, i. 395.
Tristram, H. B., on unhealthy districts in North Africa, i. 244;
on the habits of the chaffinch in Palestine, i. 307;
on the birds of the Sahara, ii. 172;
on the animals inhabiting the Sahara, ii. 224.
Triton cristatus, ii. 24.
Triton palmipes, ii. 24.
Triton punctatus, ii. 24, 25.
Troglodytes vulgaris, ii. 198.
Trogons, colours and nidification of the, ii. 171, 173.
Tropic-birds, white only when mature, ii. 228.

Page 396

Tropics, freshwater fishes of the, ii. 17.
Trout, proportion of the sexes in, i. 308;
male, pugnacity of the, ii. 3.
Trox sabulosus, stridulation of, i. 380.
Truth, not rare between members of the same tribe, i. 95;
more highly appreciated by certain tribes, i. 100.
Tulloch, Major, on the immunity of the negro from certain fevers, i. 243.
Tumbler, almond, change of plumage in the, i. 294.
Turdus merula, ii. 170;
young of, ii. 219.
Turdus migratorius, ii. 185.
Turdus musicus, ii. 170.
Turdus polyglottus, young of, ii. 219.
Turdus torquatus, ii. 170.
Turkey, swelling of the wattles of the male, ii. 72;
variety of, with a top-knot, ii. 74;
recognition of a dog by a, ii. 110;
wild, pugnacity of young male, ii. 48;
wild, notes of the, ii. 60;
male, wild, acceptable to domesticated females, ii. 119;
wild, first advances made by older females, ii. 121;
wild, breast-tuft of bristles of the, ii. 179.
Turkey-cock, scraping of the wings of, upon the ground, ii. 61;
wild, display of plumage by, ii. 87;
fighting habits of, ii. 98.
Turner, Prof. W., on muscular fasciculi in man referable to the panniculus carnosus, i. 19;
on the occurrence of the supra-condyloid foramen in the human humerus, i. 28;
on muscles attached to the coccyx in man, i. 29;
on the filum terminale in man, i. 30;
on the variability of the muscles, i. 109;
on abnormal conditions of the human uterus, i. 123;
on the development of the mammary glands, i. 209;
on male fishes hatching ova in their mouths, i. 210.
Turnix, sexes of some species of, ii. 201, 207.
Turtle-dove, cooing of the, ii. 60.
Tuttle, H., on the number of species of man, i. 226.
Tylor, E. B., on emotional cries, gestures, &c., of man, i. 54;
on the origin of the belief in spiritual agencies, i. 66;
on the primitive barbarism of civilised nations, i. 181;
on the origin of counting, i. 181;
on resemblances of the mental characters in different races of man, i. 232.
Type of structure, prevalence of, i. 211.
Typhœus, stridulating organs of, i. 378;
stridulation of, i. 380.
Twins, tendency to produce, hereditary, i. 133.
Twite, proportion of the sexes in the, i. 307.

U.
Ugliness, said to consist in an approach to the lower animals, ii. 354.
Umbrella-bird, ii. 58, 59.

Page 397

Umbrina, sounds produced by, ii. 23.
United States, rate of increase in, i. 131;
influence of natural selection on the progress of, i. 179;
change undergone by Europeans in the, i. 246.
Upupa epops, sounds produced by the male, ii. 62.
Uraniidæ, coloration of the, i. 396.
Uria troile, variety of, (= U. lacrymans), ii. 127.
Urodela, ii. 24.
Urosticte Benjamini, sexual differences in, ii. 151.
Use and disuse of parts, effects of, i. 116;
influence of, on the races of man, i. 247.
Uterus, reversion in the, i. 123;
more or less divided, in the human subject, i. 123, 130;
double, in the early progenitors of man, i. 206.

V.
Vaccination, influence of, i. 168.
Vancouver Island, Mr. Sproat on the savages of, i. 239;
natives of, eradication of facial hair by the, ii. 348.
Vanellus cristatus, wing tubercles of the male, ii. 48.
Vanessæ, i. 387;
resemblance of lower surface of, to bark of trees, i. 392.
Variability, causes of, i. 111;
in man, analogous to that in the lower animals, i. 112;
of the races of man, i. 225;
greater in men than in women, i. 275;
period of, relation of the, to sexual selection, i. 296;
of birds, ii. 124;
of secondary sexual characters in man, ii. 320.
Variation, correlated, i. 30;
laws of, i. 113;
in man, i. 185;
analogous, i. 194;
analogous, in plumage of birds, ii. 74.
Variations, spontaneous, i. 131.
Varieties, absence of, between two species, evidence of their distinctness, i. 215.
Variety, an object in nature, ii. 230.
Variola, communicable between man and the lower animals, i. 11.
Vauréal, i. 29.
Veddahs, monogamous habits of, ii. 363.
Veitch, Mr., on the aversion of Japanese ladies to whiskers, ii. 349.
Vengeance, instinct of, i. 89.
Venus Erycina, priestesses of, ii. 357.
Vermes, i. 327.
Vermiform appendage, i. 27.
Verreaux, M., on the attraction of numerous males by the female of an Australian Bombyx, i. 312.
Vertebræ, caudal, number of, in macaques and baboons, i. 150;
of monkeys, partly imbedded in the body, i. 151.
Vertebrata, ii. 1;
common origin of the, i. 203;

Page 398

most ancient progenitors of, i. 212;
origin of the voice in air-breathing, ii. 331.
Vesicula prostatica, the homologue of the uterus, i. 31, 208.
Vibrissæ, represented by long hairs in the eyebrows, i. 25.
Vidua, ii. 181.
Vidua axillaris, i. 269.
Villerme, M., on the influence of plenty upon stature, i. 115.
Vinson, Aug., on the male of Epeira nigra, i. 338.
Viper, difference of the sexes in the, ii. 29.
Virey, on the number of species of man, i. 226.
Virtues, originally social only, i. 93;
gradual appreciation of, i. 165.
Viscera, variability of, in man, i. 109.
Viti Archipelago, population of the, i. 225.
Vlacovich, Prof., on the ischio-pubic muscle, i. 127.
Vocal music of birds, ii. 51.
Vocal organs of man, i. 58;
of birds, i. 59; ii. 163;
of frogs, ii. 28;
of the Insessores, ii. 55;
difference of, in the sexes of birds, ii. 56;
primarily used in relation to the propagation of the species, ii. 330.
Vogt, Carl, on the origin of species, i. 1;
on the origin of man, i. 4;
on the semilunar fold in man, i. 23;
on the imitative faculties of microcephalous idiots, i. 57;
on microcephalous idiots, i. 121;
on skulls from Brazilian caves, i. 218;
on the evolution of the races of man, i. 230;
on the formation of the skull in women, ii. 317;
on the Ainos and negroes, ii. 321;
on the increased cranial difference of the sexes in man with race-development, ii. 329;
on the obliquity of the eye in the Chinese and Japanese, ii. 344.
Voice in mammals, ii. 274;
in monkeys and man, ii. 319;
in man, ii. 330;
origin of, in air-breathing vertebrates, ii. 331.
Von Baer, definition of advancement in the organic scale, i. 211.
Vulpian, Prof., on the resemblance between the brains of man and of the higher apes, i. 11.
Vultures, selection of a mate by the female, ii. 116;
colours of, ii. 229.

W.
Waders, young of, ii. 217.
Wagner, R., on the occurrence of the diastema in a Kafir skull, i. 126;
on the bronchi of the black stork, ii. 60.
Wagtail, Ray’s, arrival of the male before the female, i. 260.
Wagtails, Indian, young of, ii. 190.
Waist, proportions of, in soldiers and sailors, i. 117.
Waitz, Prof., on the number of species of man, i. 226;

Page 399

on the colour of Australian infants, ii. 318;
on the beardlessness of negroes, ii. 321;
on the fondness of mankind for ornaments, ii. 338;
on the liability of negroes to tropical fevers after residence in a cold climate, i. 243;
on negro ideas of female beauty, ii. 346;
on Javanese and Cochin Chinese ideas of beauty, ii. 347.
Walckenaer and Gervais, on the Myriapoda, i. 340.
Waldeyer, M., on the hermaphroditism of the vertebrate embryo, i. 207.
Wales, North, numerical proportion of male and female births in, i. 301.
Walker, Alex., on the large size of the hands of labourers’ children, i. 117.
Walker, F., on sexual differences in the diptera, i. 348.
Wallace, Dr. A., on the prehensile
use of the tarsi in male moths, i. 256;
on the rearing of the Ailanthus silk-moth, i. 311;
on breeding Lepidoptera, i. 311;
proportion of sexes of Bombyx cynthia, B. yamamai, and B. Pernyi, reared by, i. 313;
on the development of Bombyx cynthia and B. yamamai, i. 346;
on the pairing of Bombyx cynthia, i. 401;
on the fertilisation of moths, i. 406.
Wallace, A. R., on the origin of man, i. 4;
on the power of imitation in man, i. 39;
on the use of missiles by the orang, i. 52;
on the varying appreciation of truth among different tribes, i. 100;
on the limits of natural selection in man, i. 137, 158;
on the occurrence of remorse among savages, i. 165;
on the effects of natural selection on civilised nations, i. 168;
on the use of the convergence of the hair at the elbow in the orang, i. 193;
on the contrast in the characters of the Malays and Papuans, i. 216;
on the line of separation between the Papuans and Malays, i. 218;
on the sexes of Ornithoptera Crœsus, i. 310;
on protective resemblances, i. 322;
on the relative sizes of the sexes of insects, i. 346;
on Elaphomyia, i. 349;
on the Birds of Paradise, i. 269;
on the pugnacity of the males of Leptorhynchus angustatus, i. 375;
on sounds produced by Euchirus longimanus, i. 381;
on the colours of Diadema, i. 388;
on Kallima, i. 392;
on the protective colouring of moths, i. 394;
on bright coloration as protective in butterflies, i. 395;
on variability in the Papilionidæ, i. 402;
on male and female butterflies inhabiting different stations, i. 403;
on the protective nature of the dull colouring of female butterflies, i. 403, 405, 414;
on mimickry in butterflies, i. 412;
on the mimickry of leaves by Phasmidæ, i. 414;
on the bright colours of caterpillars, i. 416;
on brightly-coloured fishes frequenting reefs, ii. 17;
on the coral snakes, ii. 31;
on Paradisea apoda, ii. 74, 78;
on the display of plumage by male Birds of Paradise, ii. 88;

Page 400

on assemblies of Birds of Paradise, ii. 101;
on the instability of the ocellated spots in Hipparchia Janira, ii. 132;
on sexually limited inheritance, ii. 155;
on the sexual coloration of birds, ii. 166, 196, 197, 200, 206;
on the relation between the colours and nidification of birds, ii. 166, 171;
on the coloration of the Cotingidæ, ii. 177;
on the females of Paradisea apoda and papuana, ii. 193;
on the incubation of the cassowary, ii. 204;
on protective coloration in birds, ii. 223;
on the hair of the Papuans, ii. 340;
on the Babirusa, ii. 264;
on the markings of the tiger, ii. 302;
on the beards of the Papuans, ii. 322;
on the distribution of hair on the human body, ii. 375.
Walrus, development of the nictitating membrane in the, i. 23;
tusks of the, ii. 241, 248;
use of the tusks by the, ii. 257.
Walsh, B. D., on the proportion of the sexes in Papilio Turnus, i. 310;
on the Cynipidæ and Cecidomyidæ, i. 314;
on the jaws of Ammophila, i. 342;
on Corydalis cornutus, i. 342;
on the prehensile organs of male insects, i. 342;
on the antennæ of Penthe, i. 343;
on the caudal appendages of dragon-flies, i. 344;
on Platyphyllum concavum, i. 356;
on the sexes of the Ephemeridæ, i. 361;
on the difference of colour in the sexes of Spectrum femoratum, i. 361;
on sexes of dragon-flies, i. 361;
on the difference of the sexes in the Ichneumonidæ, i. 365;
on the sexes of Orsodacna atra, i. 368;
on the variation of the horns of the male Phanæus carnifex, i. 370;
on the coloration of the species of Anthocharis, i. 393.
Wapiti, battles of, ii. 240;
traces of horns in the female, ii. 245;
attacking a man, ii. 253;
crest of the male, ii. 282;
sexual difference in the colour of the, ii. 289.
Warbler, Hedge-, ii. 198;
young of the, ii. 209.
Warblers, Superb, nidification of, ii. 169.
Wariness, acquired by animals, i. 50.
Warington, R., on the habits of the sticklebacks, ii. 2, 20;
on the brilliant colours of the male stickleback during the breeding season, ii. 14.
Wart-hog, tusks and pads of the, ii. 265.
Watchmakers, short-sighted, i. 118.
Water-hen, ii. 40.
Waterhouse, C. O., on blind beetles, i. 367;
on difference of colour in the sexes of beetles, i. 367.
Waterhouse, G. R., on the voice of Hylobates agilis, ii. 332.
Water-ouzel, autumn song of the, ii. 54.

Page 401

Waterton, C., on the pairing of a Canada goose with a Bernicle gander, ii. 114;
on hares fighting, ii. 239;
on the Bell-bird, ii. 79.
Wattles, disadvantageous to male birds in fighting, ii. 98.
Wealth, influence of, i. 169.
Weale, J. Mansel, on a South African caterpillar, i. 416.
Weapons, employed by monkeys, i. 51;
use of, i. 137;
offensive, of males, i. 257;
of mammals, ii. 241 et seq.
Weaver-bird, ii. 54.
Weaver-birds, rattling of the wings of, ii. 62;
assemblies of, ii. 101.
Webb, Dr., on the wisdom teeth, i. 25.
Wedgwood, Hensleigh, on the origin of language, i. 56.
Weevils, sexual difference in length of snout in some, i. 255.
Weir, Harrison, on the numerical proportion of the sexes in pigs and rabbits, i. 305;
on the sexes of young pigeons, i. 306;
on the songs of birds, ii. 52;
on pigeons, ii. 109;
on the dislike of blue pigeons to other coloured varieties, ii. 118;
on the desertion of their mates by female pigeons, ii. 119.
Weir, J. Jenner, on the nightingale and blackcap, i. 259;
on the relative sexual maturity of male birds, i. 261;
on female pigeons deserting a feeble mate, i. 262;
on three starlings frequenting the same nest, i. 269;
on the proportion of the sexes in Machetes pugnax and other birds, i. 306, 307;
on the coloration of the Triphænæ, i. 395;
on the rejection of certain caterpillars by birds, i. 417;
on sexual differences of the beak in the goldfinch, ii. 40;
on a piping bullfinch, ii. 52;
on the object of the nightingale’s song, ii. 52;
on song-birds, ii. 53;
on the pugnacity of male fine-plumaged birds, ii. 93;
on the courtship of birds, ii. 94;
on the finding of new mates by Peregrine-falcons and Kestrels, ii. 104;
on the bullfinch and starling, ii. 105;
on the cause of birds remaining unpaired, ii. 107;
on starlings and parrots living in triplets, ii. 107;
on recognition of colour by birds, ii. 110;
on hybrid birds, ii. 113;
on the selection of a greenfinch by a female canary, ii. 115;
on a case of rivalry of female bullfinches, ii. 121;
on the maturity of the Golden pheasant, ii. 213.
Weisbach, Dr., measurement of men of different races, i. 216;
on the greater variability of men than of women, i. 275;
on the relative proportions of the body in the sexes of different races of man, ii. 320.
Welcker, M., on Brachycephaly and Dolichocephaly, i. 148;
on sexual differences in the skull in man, ii. 317.
Wells, Dr., on the immunity of coloured races from certain poisons, i. 243.

Page 402

Westring, on the stridulation of Reduvius personatus, i. 350;
on the stridulating organs of the Coleoptera, i. 382;
on sounds produced by Cychrus, i. 382;
on the stridulation of males of Theridion, i. 339;
on the stridulation of beetles, i. 379;
on the stridulation of Omaloplia brunnea, i. 381.
Westphalia, greater proportion of female illegitimate children in, i. 301.
Westropp, H. M., on the prevalence of certain forms of ornamentation, i. 233.
Westwood, J. O., on the classification of the Hymenoptera, i. 188;
on the Culicidæ and Tabanidæ, i. 254;
on a Hymenopterous parasite with a sedentary male, i. 272;
on the proportions of the sexes in Lucanus cervus and Siagonium, i. 313;
on the absence of ocelli in female mutillidæ, i. 341;
on the jaws of Ammophila, i. 342;
on the copulation of insects of distinct species, i. 342;
on the male of Crabro cribrarius, i. 343;
on the pugnacity of male Tipulæ i. 349;
on the stridulation of Pirates stridulus, i. 350;
on the Cicadæ, i. 351;
on the stridulating organs of the crickets, i. 354;
on Pneumora, i. 357;
on Ephippiger vitium, i. 355, 358;
on the pugnacity of the Mantides, i. 360;
on Platyblemnus, i. 361;
on difference in the sexes of the Agrionidæ, i. 362;
on the pugnacity of the males of a species of Tenthredinæ, i. 364;
on the pugnacity of the male stag-beetle, i. 375;
on Bledius taurus and Siagonium, i. 375;
on lamellicorn beetles, i. 378;
on the coloration of Lithosia, i. 396.
Whale, Sperm-, battles of male, ii. 240.
Whales, nakedness of, i. 148.
Whately, Archb., language not peculiar to man, i. 53;
on the primitive civilisation of man, i. 181.
Whewell, Prof., on maternal affection, i. 40.
Whiskers, in monkeys, i. 192.
White, Gilbert, on the proportion of the sexes in the partridge, i. 306;
on the house-cricket, i. 352;
on the object of the song of birds, ii. 52;
on the finding of new mates by white owls, ii. 105;
on spring coveys of male partridges, ii. 107.
Whiteness, a sexual ornament in some birds, ii. 232;
of mammals inhabiting snowy countries, ii. 298.
White-throat, aerial love-dance of the male, ii. 68.
Widow-bird, polygamous, i. 269;
breeding plumage of the male, ii. 84, 97;
female, rejecting the unadorned male, ii. 120.
Widows and widowers, mortality of, i. 176.
Wigeon, pairing with a pintail duck, ii. 114.
Wilckens, Dr., on the modification of domestic animals in mountainous regions, i. 120;

Page 403

on a numerical relation between the hairs and excretory pores in sheep, i. 248.
Wilder, Dr. Burt, on the greater frequency of supernumerary digits in men than in women, i. 276.
Williams, on the marriage-customs of the Fijians, ii. 374.
Wilson, Dr., on the conical heads of the natives of North-Western America, ii. 351;
on the Fijians, ii. 352;
on the persistence of the fashion of compressing the skull, ii. 353.
Wing-spurs, ii. 162.
Wings, differences of, in the two sexes of butterflies and Hymenoptera, i. 345;
play of, in the courtship of birds, ii. 95.
Winter, change of colour of mammals in, ii. 298.
Witchcraft, i. 68.
Wives, traces of the forcible capture of, i. 182.
Wolf, winter change of the, ii. 298.
Wolff, on the variability of the viscera in man, i. 109.
Wollaston, T. V., on Eurygnathus, i. 344;
on musical curculionidæ, i. 378;
on the stridulation of Acalles, i. 384.
Wolves learning to bark from dogs, i. 44;
hunting in packs, i. 75.
Wolves, black, ii. 294.
Wombat, black varieties of the, ii. 294.
Women distinguished from men by male monkeys, i. 13;
preponderance of, in numbers, i. 302;
effects of selection of, in accordance with different standards of beauty, ii. 355;
practice of capturing, ii. 360, 364;
early betrothals and slavery of, ii. 366;
selection of, for beauty, ii. 372;
freedom of selection by, in savage tribes, ii. 372.
Wonder, manifestations of, by animals, i. 42.
Wonfor, Mr., on sexual peculiarities in the wings of butterflies, i. 345.
Woolner, Mr., observations on the ear in man, i. 22.
Wood, J., on muscular variations in man, i. 109, 128, 129;
on the greater variability of the muscles in men than in women, i. 275.
Wood, T. W., on the colouring of the orange-tip butterfly, i. 394;
on the habits of the Saturniidæ, i. 398;
on the habits of Menura Alberti, ii. 56;
on Tetrao cupido, ii. 56;
on the display of plumage by male pheasants, ii. 89;
on the ocellated spots of the Argus pheasant, ii. 144;
on the habits of the female Cassowary, ii. 204.
Woodcock, coloration of the, ii. 226.
Woodpecker, selection of a mate by the female, ii. 116.
Woodpeckers, ii. 56;
tapping of, ii. 62;
colours and nidification of the, ii. 171, 174, 223;
characters of young, ii. 185, 199, 209.
Wormald, Mr., on the coloration of Hypopyra, i. 397.
Wounds, healing of, i. 13.
Wren, ii. 198;
young of the, ii. 209.

Page 404

Wright, C. A., on the young of Orocetes and Petrocincla, ii. 220.
Wright, Chauncey, on correlative acquisition, ii. 335;
on the enlargement of the brain in man, ii. 391.
Wright, Mr., on the Scotch deerhound, ii. 261;
on sexual preference in dogs, ii. 271;
on the rejection of a horse by a mare, ii. 272.
Wright, W. von, on the protective plumage of the Ptarmigan, ii. 81.
Writing, i. 182.
Wyman, Prof., on the prolongation of the coccyx in the human embryo, i. 16;
on the condition of the great toe in the human embryo, i. 17;
on variation in the skulls of the natives of the Sandwich Islands, i. 108;
on the hatching of the eggs in the mouths and branchial cavities of male fishes, i. 210, ii. 20.

X.
Xenarchus, on the Cicadæ, i. 350.
Xenorhynchus, sexual difference in the colour of the eyes in, ii. 129.
Xiphophorus Hellerii, peculiar anal fin of the male, ii. 9, 10.
Xylocopa, difference of the sexes in, i. 366.

Y.
Yarrell, W., on the habits of the Cyprinidæ, i. 309;
on Raia clavata, ii. 2;
on the characters of the male salmon during the breeding season, ii. 4, 14;
on the characters of the rays, ii. 6;
on the gemmeous dragonet, ii. 8;
on the spawning of the salmon, ii. 19;
on the incubation of the Lophobranchii, ii. 21;
on rivalry in song-birds, ii. 53;
on the trachea of the swan, ii. 60;
on the moulting of the anatidæ, ii. 85;
on an instance of reasoning in a gull, ii. 108;
on the young of the waders, ii. 217.
Yellow fever, immunity of negroes and mulattoes from, i. 243.
Youatt, Mr., on the development of the horns in cattle, i. 284.
Yura-caras, their notions of beauty, ii. 347.

Z.
Zebra, rejection of an ass by a female, ii. 295;
stripes of the, ii. 302.
Zebus, humps of, i. 284.
Zigzags, prevalence of, as ornaments, i. 233.
Zincke, Mr., on European emigration to America, i. 179.
Zootoca vivipara, sexual difference in the colour of, ii. 36.
Zygænidæ, coloration of the, i. 396.

Page 405

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