Descent of Man [ 1871]

[Bullseye]

With stags of many kinds the branches of the horns offer a curious
case of difficulty; for certainly a single straight point would
inflict a much more serious wound than several diverging ones. 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, one of
which bears the astonishing number of thirty-three branches and the
other twenty-seven, making altogether sixty branches. Richardson
figures a pair of antlers of the wild reindeer with twenty-nine
points.* 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,*(2) M. Bailly actually comes to the
conclusion that their horns are more injurious than useful to them.
But this author overlooks the pitched battles between rival males.
As I felt much perplexed about the use or advantage of the branches, I
applied to Mr. McNeill of Colonsay, 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 use, 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 both as to red-deer and fallow-deer that, in fighting,
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 in 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 ends 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.*(3)

  * On the horns of red-deer, Owen, British Fossil Mammals, 1846, p.
478; Richardson on the horns of the reindeer, Fauna Bor. Americana,
1829, p. 240. I am indebted to Prof. Victor Carus, for the
Moritzburg case.
  *(2) 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 Sciences
Nat., tom. ii., 1824, p. 371.
  *(3) See a most interesting account in the Appendix to Hon. J. D.
Caton's paper, as above quoted.

  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 in part as ornaments. That the
branched antlers of stags as well as the elegant lyrated horns of
certain antelopes, with their graceful double curvature (see fig. 64),
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
modified partly for this purpose, though mainly for actual service
in battle; but I have no evidence in favour of this belief.
  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 journal* 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 afterwards another, 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." A critic has well objected to this account
by asking, why, if the simple horns are now so advantageous, were
the branched antlers of the parent-form ever developed? To this I
can only answer by remarking, that a new mode of attack with new
weapons might be a great advantage, as shewn by the case of the Ovis
cycloceros, who thus conquered a domestic ram famous for his
fighting power. Though the branched antlers of a stag are well adapted
for fighting with his rivals, and though it might be an advantage to
the prong-horned variety slowly to acquire long and branched horns, if
he had to fight only with others of the same kind, yet it by no
means follows that branched horns would be the best fitted for
conquering a foe differently armed. In the foregoing case of the
Oryx leucoryx, it is almost certain that the victory would rest with
an antelope having short horns, and who therefore did not need to
kneel down, though an Oryx might profit by having still longer
horns, if he fought only with his proper rivals.

  * The American Naturalist, Dec., 1869, p. 552.

[Bullseye]

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 downwards with serious effect.* The walrus, though
having so short a neck and so unwieldy a body, "can strike either
upwards, or downwards, or sideways, with equal dexterity."*(2) I was
informed by the late Dr. Falconer, that the Indian elephant fights
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 howdah.*(3)

  * Pallas Spicilegia Zoologica, fasc. xiii., 1779, p. 18.
  *(2) Lamont, Seasons with the Sea-Horses, 1861, p. 141.
  *(3) See also Corse (Philosophical Transactions, 1799, p. 212) on
the manner in which the short-tusked Mooknah variety attacks other
elephants.

  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 we may infer from what follows that one
form of weapon has often been replaced in the course of ages by
another. With ruminants the development of horns generally stands in
an inverse relation with that of even moderately 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 males." The Camelidae have,
in addition to their true canines, a pair of canine-shaped incisors in
their upper jaws.* 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. In 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 of certain deer have been known occasionally to
exhibit rudiments of these teeth.*(2) 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 wide like camels and
guanacoes. Whenever the adult male possesses canines, now inefficient,
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 have 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 (but
not in the horse) caused by the development of new weapons.

  * Owen, Anatomy of Vertebrates, vol. iii., p. 349.
  *(2) See Ruppell (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 (Palaeont. 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.

  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 given by some authors.* 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
pounds and a quarter. Although the horns are not periodically
renewed in sheep, 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 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
horizontally back; and in this attitude cannot see the ground
distinctly."*(2) 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 red-deer for about twelve weeks, they are extremely
sensitive to a blow; so that in Germany the stags at this time
somewhat change their habits, and avoiding dense forests, frequent
young woods and low thickets.*(3) 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.

  * Emerson Tennent, Ceylon, 1859, vol. ii., p. 275; Owen, British
Fossil Mammals, 1846, p. 245.
  *(2) Richardson, Fauna Bor. Americana, on the moose, Alces
palmata, pp. 236, 237; on the expanse of the horns, Land and Water,
1869, p. 143. See also Owen, British Fossil Mammals, on the Irish elk,
pp. 447, 455.
  *(3) Forest Creatures, by C. Boner, 1861, p. 60.

  With mammals, when, as is often the case, the sexes differ in
size, the males are almost always larger and stronger. I am informed
by Mr. Gould that this holds good in a marked manner 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.* Dr. Gill remarks
that it is with the polygamous seals, the males of which are well
known to fight savagely together, that the sexes differ much in
size; the monogamous species differing but little. Whales also
afford evidence of the relation existing between the pugnacity of
the males and their large size compared with that of the female; the
males of the right-whales do not fight together, and they are not
larger, but rather smaller, than their females; on the other hand,
male sperm-whales fight much together, and their bodies are "often
found scarred with the imprint of their rival's teeth," and they are
double the size of the females. The greater strength of the male, as
Hunter long ago remarked,*(2) is invariably displayed 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 mere 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.

[Bullseye]

* See the very interesting paper by Mr. J. A. Allen in Bull. Mus.
Comp. Zoology of Cambridge, United States, vol. ii., No. 1, p. 82. The
weights were ascertained by a careful observer, Capt. Bryant. Dr. Gill
in The American Naturalist, January, 1871, Prof. Shaler on the
relative size of the sexes of whales, American Naturalist, January,
1873.
  *(2) Animal Economy, p. 45.

  From these considerations I was anxious to obtain information as
to the Scotch deer-hound, 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, well known for his success with
this breed, who has with great kindness collected for me the following
facts from various sources. Fine male dogs, measured at the
shoulder, range from 28 inches, which is low, to 33 or even 34
inches in height; and in weight from 80 pounds, which is light, to 120
pounds, or even more. The females range in height from 23 to 27, or
even to 28 inches; and in weight from 50 to 70, or even 80 pounds.*
Mr. Cupples concludes that from 95 to 100 pounds for the male, and
70 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 ****-puppy will occasionally be larger than any of
the dogs, but they are invariably beaten by them later." Mr.
McNeill, of Colonsay, 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
deer-hound is not acquired until rather late in life. The males almost
exclusively are 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 dimensions until rather late 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
deer-hound may probably be accounted for.

  * See also Richardson's Manual on the Dog, p. 59. Much valuable
information on the Scottish deer-hound 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.

  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. The same observer remarks that rhinoceroses in fighting,
parry each other's sidelong blows with their horns, which clatter
loudly together, as do the tusks of boars. Although wild boars fight
desperately, they seldom, according to Brehm, receive fatal wounds, as
the blows fall on each other's tusks, or on the layer of gristly
skin covering the shoulder, called by the German hunters, the
shield; and here we have a part specially modified for defence. With
boars in the prime of life (see fig. 65) 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 used in this way. They may, however, still serve, and even more
effectively, as a means of defence. In compensation for the loss of
the lower tusks as weapons of offence, those in the upper jaw, which
always project a little laterally, increase in old age so much in
length and curve so much upwards that they can be used for attack.
Nevertheless, an old boar is not so dangerous to man as one at the age
of six or seven years.*

  * Brehm, Thierleben, B. ii., ss. 729-32.

  In the full-grown male Babirusa pig of Celebes (see fig. 66), 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, however, if the head were held a
little laterally, would serve as an excellent guard; and hence,
perhaps, it is that in old animals they "are generally broken off,
as if by fighting."* 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 tusks assume in a less degree
and only during old age nearly the same form, and then serve in like
manner solely for defence.

  * See Mr. Wallace's interesting account of this animal, The Malay
Archipelago, 1869, vol. i., p. 435.

  In the wart-hog (see Phacochoerus aethiopicus, fig. 67) 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 to a certain extent used
for this purpose. But the wart-hog is not destitute of other special
means of protection, for it has, on each side of the face, beneath the
eyes, a rather stiff, yet flexible, cartilaginous, oblong pad (see
fig. 67), 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 admirably protect the somewhat
prominent eyes. I may add, on the authority of Mr. Bartlett, that
these boars when fighting stand directly face to face.
  Lastly, the African river-hog (Potomochoerus 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 long, and were found in the
morning much exhausted, but not seriously wounded. It is a significant
fact, as shewing the purposes of the above-described projections and
excrescences, that these were covered with blood, and were scored
and abraded in an extraordinary manner.
  Although the males of so many members of the pig family are provided
with weapons, and as we have just seen with means of defence, these
weapons seem to have been acquired within a rather late geological
period. Dr. Forsyth Major specifies* several miocene species, in
none of which do the tusks appear to have been largely developed in
the males; and Professor Rutimeyer was formerly struck with this
same fact.

[Bullseye]

* Atti della Soc. Italiana di Sc. Nat., 1873, vol. xv. fasc. iv.

  The mane of the lion forms a good defence against the attacks of
rival lions, the one danger to which he is liable; 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."* 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)*(2) 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, except 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.

  * The Times, Nov. 10, 1857. In regard to the Canada lynx, see
Audubon and Bachman, Quadrupeds of North America, 1846, p. 139.
  *(2) 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.

  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 of the stag (Cervus elaphus) 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 have been equally protected.

  Choice in Pairing by either Sex of Quadrupeds.- Before describing in
the next chapter, the differences between the sexes in voice, odours
emitted, and ornaments, 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 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 by several males, as
commonly occurs, she would often have the opportunity, whilst they
were fighting together, of escaping with some one male, or at least of
temporarily pairing with him. This latter contingency has often been
observed in Scotland with female red-deer, as I am informed by Sir
Philip Egerton and others.*

  * 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.

  It is scarcely possible that much should be known about female
quadrupeds in a state of nature making any choice in their marriage
unions. The following curious details on the courtship of one of the
eared seals (Callorhinus ursinus) are given* 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 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."

  * Mr. J. A. Allen in Bull. Mus. Comp. Zoolog. of Cambridge, United
States, vol. ii., No. 1, p. 99.

  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 a large size.* The
well-known veterinary Blaine states*(2) that his own female pug dog
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.

  * Dogs: their Management, by E. Mayhew, M. R. C. V. S., 2nd ed.,
1864, pp. 187-192.
  *(2) Quoted by Alex. Walker, On Intermarriage, 1838, p. 276; see
also p. 244.

  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
showing 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 in 1868, a female deerhound in his
kennel 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, as I could, other instances, and I will only
add that Mr. Barr, who has carefully bred many bloodhounds, states
that in almost every instance particular individuals of opposite sexes
shew a decided preference for each other. Finally, Mr. Cupples,
after attending to this subject for another year, has 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 characters, 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 as to be exhausted, should be so
particular in their choice. Mr. Blenkiron has never known a mare to
reject a horse; but this has occured in Mr. Wright's stable, so that
the mare had to be cheated. Prosper Lucas* quotes various statements
from French authorities, and remarks, "On voit des etalons qui
s'eprennent d'une jument, et negligent toutes les autres." He gives,
on the authority of Baelen, similar facts in regard to bulls; and
Mr. H. Reeks assures me that a famous short-horn bull belonging to his
father "invariably refused to be matched with a black cow."
Hoffberg, in describing the domesticated reindeer of Lapland says,
"Foeminae majores et fortiores mares prae, caeteris admittunt, ad
eos confugiunt, a junioribus agitatae, qui hos in fugam
conjiciunt."*(2) A clergyman, who has bred many pigs, asserts that
sows often reject one boar and immediately accept another.

  * Traite de l'Hered. Nat., tom. ii., 1850, p. 296.
  *(2) Amaenitates Acad., vol. iv., 1788, p. 160.

  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.



--------------------------------------------------------------------------------

[Bullseye]

Chapter XVIII - Secondary Sexual Characters of Mammals- Continued

  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 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,* are said to be completely mute excepting at this season.
As the throats (i.e. the larynx and thyroid bodies*(2)) of stags
periodically become enlarged at the beginning of the
breeding-season, it might be thought that their powerful voices must
be somehow 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 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 fists in rage or agony. No doubt
stags challenge each other to mortal combat by bellowing; but those
with the more powerful voices, unless at the same time the stronger,
better-armed, and more courageous, would not gain any advantage over
their rivals.

  * Owen, Anatomy of Vertebrates, vol. iii., p. 585.
  *(2) Ibid., p. 595.

  It is possible that the roaring of the lion may be of some service
to him by 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 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,* 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 be intelligible on the principle
of sexual selection, together with inheritance limited to the same sex
and season; 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, in our
present state of knowledge, the most probable view.

  * See, for instance, Major W. Ross King (The Sportsman in Canada,
1866, pp. 53, 131) on the habits of the moose and wild reindeer.

  The voice of the adult male gorilla is tremendous, and he is
furnished with a laryngeal sack, as is the adult male orang.* The
gibbons rank among the noisiest of monkeys, and the Sumatra species
(Hylobates syndactylus) is also furnished with an air sack; but Mr.
Blyth, who has had opportunities for observation, does not believe
that the male is noisier 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 the
beaver.*(2) Another gibbon, the H. agilis, is remarkable, from
having the power of giving a complete and correct octave of musical
notes,*(3) 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 in warm weather make the forests resound at morning and
evening with their overwhelming voices. The males begin the dreadful
concert, and often continue it during many hours, the females
sometimes joining in with their less powerful voices. An excellent
observer, Rengger,*(4) could not perceive that they were excited to
begin 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 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 thus
gained- I will not pretend to say; but the former view, at least in
the case of the Hylobates agilis, seems the most probable.

  * Owen Anatomy of Vertebrates, vol. iii., p. 600.
  *(2) Mr. Green, in Journal of Linnean Society, vol. x., Zoology,
1869, note 362.
  *(3) C. L. Martin, General Introduction to the Natural History of
Mamm. Animals, 1841, p. 431.
  *(4) Naturgeschichte der Saugethiere von Paraguay, 1830, ss. 15, 21.

  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) becomes greatly elongated during the breeding-season,
and can then be erected. In this state it is sometimes a foot in
length. The female is not thus provided at any period of life. The
male makes a wild, hoarse, gurgling noise, which is audible at a great
distance and is believed to be strengthened by the proboscis; the
voice of the female being different. Lesson compares the **** of
the proboscis, with the swelling of the wattles of male gallinaceous
birds whilst courting the females. In another allied kind of seal, the
bladder-nose (Cystophora cristata), the head is covered by a great
hood or bladder. This supported by the septum of the nose, which is
produced far backwards and rises into an internal crest seven inches
in height. The hood is clothed with short hair, and is muscular; 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 they likewise roar or bellow; and whenever irritated the
bladder is inflated and quivers. 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; but this
is not probable, for, as I am assured by Mr. Lamont who killed 600
of these animals, the hood is rudimentary in the females, and it is
not developed in the males during youth.*

  * 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, in Proc.
Zoolog. Soc., 1868, p. 435.

  Odour.- With some animals, as with the notorious skunk of America,
the overwhelming odour which they emit appears to serve exclusively as
a defence. With shrew-mice (Sorex) both sexes possess abdominal
scent-glands and there can be little doubt, from the rejection of
their bodies by birds and beasts of prey, that the odour is
protective; nevertheless, the glands become enlarged in the males
during the breeding-season. In many other quadrupeds the glands are of
the same size in both sexes,* but their uses are not known. In other
species the glands are confined to the males, or are more developed
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. The males, and rarely the females, of many kinds
of bats have glands and protrudable sacks situated in various parts;
and it is believed that these are odoriferous.

  * 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. On bats, Mr.
Dobson, Proceedings of the Zoological Society. 1873, p. 241.

  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 perceived the air tainted with the odour of the male
Cervus campestris, at half a mile to leeward of a herd; and a silk
handkerchief, in which I carried home a skin, though often 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.* Besides the general odour, permeating the whole body of
certain ruminants (for instance Bos moschatus) in 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 myself seen in an antelope. They
are "usually larger in the male than in the female, and their
development is checked by castration."*(2) According to Desmarest they
are altogether absent in the female of Antilope subgutturosa. Hence,
there can be no doubt that they stand in 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 of this deer is from its position 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.*(3)

  * Rengger, Naturgeschichte der Saugethiere von Paraguay, 1830, s.
355. This observer also gives some curious particulars in regard to
the odour.
  *(2) Owen, Anatomy of Vertebrates, vol. iii., p. 632. See also Dr.
Murie's observations on those glands in the Proc. Zoolog. Soc.,
1870, p. 340. Desmarest, "On the Antilope subgutturosa," Mammalogie,
1820, p. 455.
  *(3) Pallas, Spicilegia Zoolog., fasc. xiii., 1779, p. 24;
Desmoulins, Dict. Class. d'Hist. Nat., tom. iii., p. 586.

  In most cases, when only the male emits a strong odour during the
breeding-season, it 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 far from agreeable to us; and that dogs, though they will
not eat carrion, sniff and roll on 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 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 most 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
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 only a thin and narrow crest runs along the back; for a
crest of this kind would afford scarcely any protection, and the ridge
of the back is not a place likely 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
scriptus* (see fig. 70) and Portax picta may be given as instances.
When stags, and the males of the wild goat, are enraged or
terrified, these crests stand erect;*(2) but it cannot be supposed
that they have been developed merely 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 fore-legs are almost concealed by an extraordinary
growth of hair, which depends 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.

[Bullseye]

* Dr. Gray, Gleanings from the Menagerie at Knowsley, pl. 28.
  *(2) Judge Caton on the wapiti, Transact. Ottawa Acad. Nat.
Sciences, 1868, pp. 36, 40; Blyth, Land and Water, on Capra aegagrus
1867, p. 37.

 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. Thus the bull alone has curled hair on the forehead.* In
three closely-allied
sub-genera of the goat family, only the males possess beards sometimes
of large size; in two other sub-genera both sexes have a beard, but it
disappears in 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 it is so small at other times
that it may be called rudimentary.*(2) 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
(see fig. 68). So it is with the whiskers of some species of
Macacus,*(3) 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.

  * Hunter's Essays and Observations, edited by Owen, 1861. vol. i.,
p. 236.
  *(2) See Dr. Gray's Catalogue of Mammalia in the British Museum,
part iii., 1852, p. 144.
  *(3) Rengger, Saugthiere, &c., s. 14; Desmarest, Mammalogie, p. 86.

  The males of various members of the ox family (Bovidae), 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 dewlaps of the bull, or the crests of hair along the backs of
certain male antelopes, are of any use to them in their ordinary
habits. 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 served by the whiskers, moustache, and other tufts of hair on the
face; and no one will suppose that these are useful as a protection.
Must we attribute all these appendages of hair or skin to mere
purposeless variability in the male? It cannot be denied that this
is possible; for in many domesticated quadrupeds, certain
characters, apparently not derived through reversion from any wild
parent form, are confined to the males, or are more developed in
them than in the females- for instance, the hump on the male
zebu-cattle of India, the tail of fat-tailed rams, the arched
outline of the forehead in the males of several breeds of sheep, and
lastly, the mane, the long hairs on the hind legs, and the dewlap of
the male of the Berbura goat.* The mane, which occurs only in the rams
of an African breed of sheep, is a true secondary sexual character,
for, as I hear from Mr. Winwood Reade, it is not developed 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
above African ram is a descendant of the same primitive stock as the
other breeds of sheep, and if the Berbura male-goat with his mane,
dewlap, &c., is descended from the same stock as other goats, then,
assuming that selection has not been applied to these characters, they
must be due to simple variability, together with sexually-limited
inheritance.

  * See the chapters on these several animals in vol. i. of my
Variation of Animals under Domestication; also vol. ii., p. 73; also
chap. xx. on the practice of selection by semi-civilised people. For
the Berbuar goat, see Dr. Gray, Catalogue, ibid., p. 157.

  Hence it appears reasonable to extend this same view to all
analogous cases with animals in a state of nature. Nevertheless I
cannot persuade myself that it generally holds good, as in the case of
the extraordinary development of hair on the throat and fore-legs of
the male Ammotragus, or in that of the immense beard of the male
Pithecia. Such study as I have been able to give to nature makes me
believe that parts or organs which are highly developed, were acquired
at some period for a special purpose. With those antelopes in which
the adult male is more strongly-coloured than the female, and with
those monkeys in which the hair on the face is elegantly arranged
and coloured in a diversified manner, it seems probable that the
crests and tufts of hair were gained as ornaments; and this I know
is the opinion of some naturalists. If this be correct, there can be
little doubt that they were gained or at least modified through sexual
selection; but how far the same view may be extended to other
mammals is doubtful.

  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."* In
the Didelphis opossum of Cayenne the female is said to be a little
more red than the male. Of the 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."*(2) 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 a large number of bats
the fur of the male is lighter than in the female.*(3) Mr. Dobson also
remarks, with respect to these animals: "Differences, depending partly
or entirely on the possession by the male of fur of a much more
brilliant hue, or distinguished by different markings or by the
greater length of certain portions, are met only, to any appreciable
extent, in the frugivorous bats in which the sense of sight is well
developed." This last remark deserves attention, as bearing on the
question whether bright colours are serviceable to male animals from
being ornamental. In one genus of sloths, it is now established, as
Dr. Gray states, "that the males are ornamented differently from the
females- that is to say, that they have a patch of soft short hair
between the shoulders, which is generally of a more or less orange
colour, and in one species pure white. The females, on the contrary,
are destitute of this mark."

[Bullseye]

* Osphranter rufus, Gould, Mammals of Australia, 1863, vol. ii. On
the Didelphis, Desmarest, Mammalogie, p. 256.
  *(2) Annals and Magazine of Natural History, Nov., 1867, p. 325.
On the Mus minutus, Desmarest, Mammalogie, p. 304.
  *(3) J. A. Allen, in Bulletin of Mus. Comp. Zoolog. of Cambridge,
United States, 1869, p. 207. Mr. Dobson on sexual characters in the
Chiroptera, Proceedings of the Zoological Society, 1873, p. 241. Dr.
Gray on sloths, ibid., 1871, p. 436.

  The terrestrial Carnivora and Insectivora rarely exhibit sexual
differences of any kind, including colour. The ocelot (Felis
pardalis), however, is exceptional, for the colours of the female,
compared with those of the male, are "moins apparentes, le fauve,
etant plus terne, le blanc moins pur, les raies ayant moins de largeur
et les taches moins de diametre."* The sexes of the allied Felis mitis
also differ, but 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 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."*(2)

  * Desmarest Mammalogie, 1820, p. 220. On Felis mitis, Rengger,
ibid., s. 194.
  *(2) Dr. Murie on the Otaria, Proceedings Zoological Society,
1869, p. 108. Mr. R. Brown on the P. groenlandica, ibid., 1868, p.
417. See also on the colours of seals, Desmarest, ibid., pp. 243, 249.

  With ruminants sexual differences of colour occur more commonly than
in any other order. A difference of this kind is general in the
strepsicerene 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. I am informed by Mr. Blyth
that the male, without shedding his hair, periodically becomes
darker during the breeding-season. Young males cannot be distinguished
from young females until about 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
evidence that the colouring of the Portax is of sexual origin, becomes
obvious, when we hear* 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.*(2)

  * Judge Caton, in Transactions of the Ottawa Academy of Natural
Sciences, 1868, p. 4.
  *(2) Dr. Gray, Cat. of Mamm. in Brit. Mus., part iii., 1852, pp.
134-142; also Dr. Gray, Gleanings from the Menagerie of Knowsley, in
which there is a splendid drawing of the Oreas derbianus: see the text
on Tragelaphus. For the Cape eland (Oreas canna), see Andrew Smith,
Zoology of S. Africa, pls. 41 and 42. There are also many of these
antelopes in the Zoological Gardens.

  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 meet in this species, as Mr.
Blyth informs me, with an exactly similar series of facts, as in 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 indistinguishable from
each other. In the Antilope niger the male is black, the female, as
well as the young of both sexes, 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 brown as in
the female; 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."* Other analogous cases could be added.

  * 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. 627. 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.

  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 kemas goat is paler, and both it and the
female Capra aegagrus are said to be more uniformly tinted than
their males. Deer rarely present any sexual differences in colour.
Judge Caton, however, informs me that in the males of the wapiti
deer (Cervus canadensis) the neck, belly, and legs are much darker
than 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 differs 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 breeding plumage.* 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 are characteristic of the adult males.*(2) Lastly, as I am
informed by Mr. Blyth, the mature male of the beautifully coloured and
spotted axis deer is considerably darker than the female: and this hue
the castrated male never acquires.

[Bullseye]

* Ottawa Academy of Sciences, May 21, 1868, pp. 3,5.
  *(2) S. Muller, 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, Catalogue, of the
British Museum, p. 146; Desmarest, Mammalogie, p. 482. On the Cervus
paludosus, Rengger, ibid., s. 345.

  The last Order which we need consider is that of the primates. The
male of the Lemur macaco is generally coal-black, whilst the female is
brown.* Of the Quadrumana of the New World, the females and young of
Mycetes caraya are greyish-yellow and like each other; in the second
year the young male becomes reddish-brown; in the third, 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 of 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.*(2) 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 the thighs is
of an elegant fawn colour, in the latter it is paler. In the beautiful
and curious moustache monkey (Cercopithecus cephus) the only
difference between the sexes is that the tail of the male is
chestnut and that of the female grey; but Mr. Bartlett informs me that
all the hues become more 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 Muller, the male of
Semnopithecus chrysomelas is nearly black, the female being pale
brown. In the Cercopithecus cynosurus and griseoviridis 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.

  * Sclater, Proc. Zool. Soc., 1866, p. i. The same fact has also been
fully ascertained by M. M. Pollen and van Dam. See, also, Dr. Gray
in Annals and Magazine of Natural History, May, 1871, p. 340.
  *(2) On Mycetes, Rengger, ibid., s. 14; and Brehm, Illustriertes
Thierleben, B. i., ss. 96, 107. On Ateles Desmarest, Mammalogie, p.
75. On Hylobates, Blyth, Land and Water, 1867, p. 135. On the
Semnopithecus, S. Muller, Zoog. Indischen Archipel., tab. x.

  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 (C. leucophaeus) the females and young are much
paler-coloured, with less green, than the adult males. No other member
in the whole class of mammals is coloured in so extraordinary a manner
as the adult male mandrill (C. 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 superieures de leurs
cuisses et le grand espace nu de leurs fesses sont egalement colores
du rouge le plus vif, avec un melange de bleu qui ne manque reellement
pas d'elegance."* 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
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. (See fig. 69.) In the
adult females and in the young of both sexes these protuberances are
scarcely perceptible; and the naked parts are much less bright
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.

  * Gervais, Hist., Nat. des Mammiferes, 1854, p. 103. Figures are
given of the skull of the male. Also Desmarest, Mammalogie, p. 70.
Geoffroy St-Hilaire and F. Cuvier, Hist. Nat. des Mammiferes, 1824,
tom. i.

[Bullseye]

In all the cases hitherto given the male is more strongly or
brighter coloured than the female, and differs from the young of
both sexes. But as with some few birds it is the female which is
brighter coloured than the male, 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, as I was assured by the
keepers in the Zoological Gardens, periodically becomes even yet
more vivid, and her face also is pale red. On the other hand, in the
adult male and in 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 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. Some of these may be the result of
variations confined to one sex and transmitted to the same sex,
without any good being gained, and therefore without the aid of
selection. We have instances of this with our domesticated animals, as
in the males of certain cats being rusty-red, whilst the females are
tortoise-shell coloured. Amalogous cases occur in 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 these
animals, were males. On the other hand, with wolves, foxes, and
apparently American squirrels, both sexes are occasionally born black.
Hence it is quite possible that with some mammals a difference in
colour between the sexes, especially when this is congenital, may
simply be the result, without the aid of selection, of the
occurrence of one or more variations, which from the first were
sexually limited in their transmission. Nevertheless it is
improbable that the diversified, vivid, and contrasted colours of
certain quadrupeds, for instance, of the above monkeys and
antelopes, can thus be accounted for. We should bear in mind that
these colours do not appear in the male at birth, but only at or
near maturity; and that unlike ordinary variations, they are lost if
the male be emasculated. It is on the whole probable 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. This
view is strengthened by the differences in colour between the sexes
occurring almost exclusively, as may be collected from the previous
details, in those groups and sub-groups of mammals which present other
and strongly-marked secondary sexual characters; these being
likewise due to sexual selection.
  Quadrupeds manifestly take notice of colour. Sir S. Baker repeatedly
observed that the African elephant and rhinoceros attacked white or
grey horses with special fury. I have elsewhere shewn* that
half-wild horses apparently prefer to pair with those of the same
colour, and that herds of fallow-deer of different colours, 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."*(2)

  * The Variation of Animals and Plants under Domestication, 1868,
vol. ii., pp. 102, 103.
  *(2) Essays and Observations, by J. Hunter, edited by Owen, 1861,
i., p. 194.

  In an earlier chapter we have seen that the mental powers of the
higher animals do not differ in kind, though 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";*- as negroes
and 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 be coloured for the sake of ornament even more brilliantly than
the face; but this is not more strange than that the tails of many
birds should be especially decorated.

  * Sir S. Baker, The Nile Tributaries of Abyssinia, 1867.

  With mammals we 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 and other
animals 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 majority of cases resemble the adult female. In both
classes the male assumes the characters proper to his sex shortly
before the age of reproduction; and if emasculated at an early period,
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 of hair or feathers, or other such 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
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 applies to mammals. With a considerable number of species,
especially of 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 so striking a
manner, as in most of the lower classes. Audubon remarks that he often
mistook the musk-rat,* 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, the
rabbit, for when running to its burrow, it is made conspicuous to
the sportsman, and no doubt to all beasts of prey, by its upturned
white tail. No one doubts that the quadrupeds inhabiting 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 for long, a white coat would be injurious; consequently,
species of this colour are 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. Pallas*(2)
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, 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,
until they became as white as snow.

  * Fiber zibethicus, Audubon and Bachman, The Quadrupeds of North
America, 1846, p. 109.
  *(2) Novae species Quadrupedum e Glirium ordine, 1778, p. 7. What
I have called the roe is the Capreolus sibricus subecaudatus of
Pallas.

  Mr. Reeks has given me a curious instance of an animal profiting
by being peculiarly coloured. He raised from fifty to sixty white
and brown piebald rabbits in a large walled orchard; and he had at the
same time some similarly coloured cats in his house. Such cats, as I
have often noticed, are very conspicuous during day; but as they
used to lie in watch during the dusk at the mouths of the burrows, the
rabbits apparently did not distinguish them from their
parti-coloured brethren. The result was that, within eighteen
months, every one of these parti-coloured rabbits was destroyed; and
there was evidence that this was effected by the cats. Colour seems to
be advantageous to another animal, the skunk, in a manner of which
we have had many instances in other classes. No animal will
voluntarily attack one of these creatures on account of the dreadful
odour which it emits when irritated; but during the dusk it would
not easily be recognized and might be attacked by a beast of prey.
Hence it is, as Mr. Belt believes,* that the skunk is provided with
a great white bushy tail, which serves as a conspicuous warning.

  * The Naturalist in Nicaragua, p. 249.

  Although we must admit that many quadrupeds have received their
present tints either as a protection, or as an aid in procuring
prey, 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 these purposes. We may take as an illustration
certain antelopes; when we see the square white patch on the throat,
the white marks on the fetlocks, and the round black spots on the
ears, 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 (see fig. 70),- we cannot
believe that differences of this kind are of any service to either sex
in their daily habits of life. It seems a much more probable
conclusion that the various marks were first acquired by the males and
their colours intensified through sexual selection, 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) (see fig. 64) have narrow white vertical
lines on their hind 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 these colours are abruptly separated from the white
belly and from 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 (see fig. 71); 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.* 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.

[Bullseye]

* See the fine plates in A. Smith's Zoology of South Africa, and Dr.
Gray's Gleanings from the Menagerie of Knowsley.

  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 believes* 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 the
analogous marks and colours are rather brighter in the male than in
the female. The zebra is conspicuously striped, and stripes cannot
afford any protection in the open plains of South Africa. Burchell*(2)
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." But as throughout the whole group
of the Equidae the sexes are identical in colour, we have here no
evidence of sexual selection. Nevertheless he who attributes the white
and dark vertical stripes on the flanks of various antelopes to this
process, will probably extend the same view to the royal tiger and
beautiful zebra.

  * Westminster Review, July 1, 1867, p. 5.
  *(2) Travels in South Africa, 1824, vol. ii., p. 315.

  We have seen in a former chapter that when young animals belonging
to any class follow nearly the same habits of life as 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 tapirs, the young are
marked with longitudinal stripes, and thus differ from all the
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 Manchurian deer (Cervus mantchuricus) is spotted during the
whole year, but, as I have seen in the Zoological Gardens, the spots
are much plainer 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.* 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 in the adults at all
seasons; and, lastly, to their absence at all ages and seasons, 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 somewhat resembled 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.
Hyomoschus, also, offers one of those interesting cases of a form
linking together two groups, for it is intermediate in certain
osteological characters between the pachyderms and ruminants, which
were formerly thought to be quite distinct.*(2)

  * 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.
  *(2) Falconer and Cautley, Proc. Geolog. Soc., 1843; and
Falconer's Pal. Memoirs, vol. i., p. 196.

  A curious difficulty here arises. If we admit that coloured spots
and stripes were first 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 almost sure that the spots and stripes disappeared at or near
maturity in the progenitors of our existing species, so that they were
still retained by the young; and owing to the law of inheritance at
corresponding ages, were transmitted to the young of all succeeding
generations. It may have been a great advantage to the lion and
puma, from the open nature of their usual haunts, 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 is now the case. As to deer, pigs, and tapirs, Fritz
Muller has suggested to me that these animals, by the removal of their
spots or stripes through natural selection, would have been less
easily seen by their enemies; and that 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 thus protected, and still more so that the adults of some species
should have retained their spots, either partially or completely,
during part of the year. We know 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, though we cannot
explain the cause. Very few horses, except dun-coloured kinds, have
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.* 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 to some other
unknown cause, it is impossible to decide. An observation made by
Mr. Sclater well 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. taeniopus, 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.*(2)

[Bullseye]

* The Variation of Animals and Plants under Domestication, 1868,
vol. i., pp. 61-64.
  *(2) Proc. Zool. Soc., 1862, p. 164. See, also, Dr. Hartmann, Ann.
d. Landw., Dd. xliii., s. 222.

  Quadrumana.- Before we conclude, it will be well to add a few
remarks on the ornaments 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 so
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 the sake of ornament. The accompanying
figures (see figs. 72 to 76) 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 use in
any ordinary way 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.
  A few instances will suffice of the strange manner in which both
sexes of some species are coloured, and of the beauty of others. The
face of the Cercopithecus petaurista (see fig. 77) 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 aethiops is grotesque, with its black face,
white whiskers and collar, chestnut 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,*
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.*(2) 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 reason to believe that the 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.

  * I observed this fact in the Zoological Gardens; and many cases may
be seen in the coloured plates in Geoffroy St-Hilaire and F. Cuvier,
Histoire Nat. des Mammiferes, tom. i., 1824.
  *(2) Bates, The Naturalist on the Amazons, 1863, vol. ii., p. 310.

  Although many kinds of monkeys are far from beautiful according to
our taste, other species are universally admired for their elegant
appearance and bright colours. The Semnopithecus nemaeus, though
peculiarly coloured, is described as extremely pretty; the
orange-tinted face is surrounded by long whiskers of glossy whiteness,
with a line of chestnut-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 being of a pure white; a gorget of chestnut
surmounts the chest; the thighs are black, with the legs chestnut-red.
I will mention only two other monkeys for their beauty; and I have
selected these as presenting 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 chestnut, 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 Zoological
Society's Gardens I have often overheard visitors admiring the
beauty of another monkey, deservedly called Cercopithecus diana (see
fig. 78); the general colour of the fur is grey; the chest and inner
surface of the fore legs are white; a large triangular defined space
on the hinder part of the back is rich chestnut; 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,
contrasting finely with a white transverse crest over the eyebrows and
a long white peaked beard, of which the basal portion is black.*

  * I have seen most of the above monkeys in the Zoological
Society's Gardens. The description of the Semnopithecus nemaeus is
taken from Mr. W. C. Martin's Natural History of Mammalia, 1841, p.
460; see also pp. 475, 523.

 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 been acquired or modified through
that form of selection which I have called sexual. This does not
depend on any superiority in the general struggle for life, but on
certain individuals of one sex, generally the male, being successful
in conquering other males, and leaving a larger number of offspring to
inherit their superiority than do 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 is probably carried on in some cases 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 must have been strengthened by use during maturity, under
the powerful excitements of love, jealousy or rage, and will
consequently have been 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 female, 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 defence,
have been partly modified for 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
tints, 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 brilliantly coloured in some species.
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 same groups and sub-groups 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 weapons, such as
horns and tusks, have often been transmitted either exclusively or
much more perfectly to the males than to the females. This is
surprising, for, as the males generally use their weapons for
defence against enemies of all kinds, their weapons would have been of
service to the females. As far as we can see, their absence in this
sex can be accounted for 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 the latter have
been modified through sexual selection, far more commonly than the
females, either for fighting with each other or for alluring the
opposite sex.




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Part Three - Sexual Selection in Relation to Man and Conclusion
Chapter XIX - Secondary Sexual Characters of Man

  WITH mankind the differences between the sexes are greater than in
most of the 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,* the
superciliary ridge is generally more 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 races the women are
said to differ slightly in tint from the men. For instance,
Schweinfurth, in speaking of a negress belonging to the Monbuttoos,
who inhabit the interior of Africa a few degrees north of the equator,
says, "Like all her race, she had a skin several shades lighter than
her husband's, being something of the colour of half-roasted
coffee."*(2) As the women labour in the fields and are quite
unclothed, it is not likely that they differ in colour from the men
owing to less exposure to the weather. European women are perhaps
the brighter coloured of the two sexes, as may be seen when both
have been equally exposed.

  * Schaaffhausen, translation, in Anthropological Review, Oct., 1868,
pp. 419, 420, 427.
  *(2) The Heart of Africa, English transl., 1873, vol i., p. 544.

  Man is more courageous, pugnacious and energetic than woman, and has
a more inventive genius. His brain is absolutely larger, but whether
or not proportionately to his larger body, 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 the body rounder, in parts
more prominent; and her pelvis is broader than in man;* 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.

  * Ecker, translation, in Anthropological Review, Oct., 1868, pp.
351-356. The comparison of the form of the skull in men and women
has been followed out with much care by Welcker.

  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 the head.
It is probably due to the rather late appearance in life of the
successive variations whereby man has 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 widely; 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.* 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.*(2) In
regard to colour, the new-born negro child is reddish nut-brown, which
soon becomes slaty-grey; the black colour being fully developed within
a year in the Soudan, but not until three years in Egypt. The eyes
of the negro are at first blue, and the hair chestnut-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.*(3)

  * Ecker and Welcker, ibid., pp. 352, 355; Vogt, Lectures on Man,
Eng. translat., p. 81.
  *(2) Schaaffhausen, Anthropolog. Review, ibid., p. 429.
  *(3) 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, Saugethiere, &c., s. 3. See also Godron, De l'Espece, tom.
ii., 1859, p. 253. For the Australians, Waitz, Introduction to
Anthropology, Eng. translat., 1863, p. 99.

  I have specified the foregoing differences between the male and
female sex in mankind, because they are curiously like those of the
Quadrumana. With these animals the female is mature at an earlier
age than the male; at least this is certainly the case in Cebus
azarae.* The males of most species are larger and stronger than the
females, of which fact the gorilla affords 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,*(2) 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.*(3) 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 the
female than in the male monkey. Even in the colour of the beard
there is a curious parallelism between man and the Quadrumana, for
with man when the beard differs in colour from the hair of the head,
as is commonly the case, it is, I believe, almost always of a
lighter tint, being often reddish. I have repeatedly observed this
fact in England; but two gentlemen have lately written to me, saying
that they form an exception to the rule. One of these gentlemen
accounts for the fact by the wide difference in colour of the hair
on the paternal and maternal sides of his family. Both had been long
aware of this peculiarity (one of them having often been accused of
dyeing his beard), and had been thus led to observe other men, and
were convinced that the exceptions were very rare. Dr. **** attended
to this little point for me in Russia, and found no exception to the
rule. In Calcutta, Mr. J. Scott, of the Botanic Gardens, was so kind
as to observe the many races of men to be seen there, as well as in
some other parts of India, namely, two races of Sikhim, the Bhoteas,
Hindoos, Burmese, and Chinese, most of which races have very little
hair on the face; and he always found that when there was any
difference in colour between the hair of the head and the beard, the
latter was invariably lighter. Now with monkeys, as has already been
stated, the beard frequently differs strikingly in colour from the
hair of the head, and in such cases it is always of a lighter hue,
being often pure white, sometimes yellow or reddish.*(4)

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* Rengger, Saugethiere, &c., 1830, s. 49.
  *(2) As in Macacus cynomolgus (Desmarest, Mammalogie, p. 65), and in
Hylobates agilis (Geoffroy St-Hilaire and F. Cuvier, Histoire Nat. des
Mammiferes, 1824, tom. i., p. 2).
  *(3) Anthropological Review, Oct., 1868, p. 353.
  *(4) Mr. Blyth informs me that he has only seen 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
Macacus cynomolgus, kept in confinement, 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 nicknamed. In certain races of man the
hair on the head hardly ever becomes grey; thus Mr. D. Forbes has
never, as he informs me, seen an instance with the Aymaras and
Quechuas of South America.

 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.* 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 the parallelism between the sexual
differences of man and the Quadrumana. With some few species, however,
as with certain baboons, the orang and the gorilla, there is a
considerably greater difference between the sexes, as 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
mankind.

  * 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 Cyclopedia, vol. ii., pp. 149, 150.

  All the secondary sexual characters of man are highly variable, even
within the limits of the same race; and they differ much in the
several races. These two rules hold good generally throughout the
animal kingdom. In the excellent observations made on board the
Novara,* the male Australians were found to exceed the females by only
65 millim. in height, whilst with the Javans 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. Numerous measurements were carefully made of the stature,
the circumference of the neck and chest, the length of the back-bone
and of the arms, in various races; and nearly all these measurements
shew that the males differ much more from one another than do the
females. This fact indicates that, as far as these characters are
concerned, it is the male which has been chiefly modified, since the
several races diverged from their common stock.

  * The results were deduced by Dr. Weisbach from the measurements
made by Drs. K. Scherzer and Schwarz, see Reise der Novara:
Anthropolog. Theil, 1867, ss. 216, 231, 234, 236, 239, 269.

  The development of the beard and the hairiness of the body differ
remarkably in the men of distinct races, and even in different
tribes or families of the same race. We Europeans see this amongst
ourselves. In the Island of St. Kilda, according to Martin,* the men
do not acquire beards until the age of thirty or upwards, and even
then the beards are very thin. On the Europaeo-Asiatic continent,
beards prevail until we pass beyond India; though with the natives
of Ceylon they are often absent, as was noticed in ancient times by
Diodorus.*(2) Eastward of India beards disappear, as with the Siamese,
Malays, Kalmucks, Chinese, and Japanese; nevertheless, the
Ainos,*(3) who inhabit the northernmost islands of the Japan
Archipelago, are the hairiest men in the world. With negroes the beard
is scanty or wanting, and they rarely have whiskers; in both sexes the
body is frequently almost destitute of fine down.*(4) On the other
hand, the Papuans of the Malay Archipelago, who are nearly as black as
negroes, possess well-developed beards.*(5) 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 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."*(6)

  * Voyage to St. Kilda (3rd ed., 1753), p. 37.
  *(2) Sir J. E. Tennent, Ceylon, vol. ii., 1859, p. 107.
  *(3) Quatrefages, Revue des Cours Scientifiques, Aug. 29, 1868, p.
630; Vogt, Lectures on Man, Eng. trans., p. 127.
  *(4) On the beards of negroes, Vogt, Lectures, &c., 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 Europeans.
  *(5) Wallace, The Malay Arch., vol. ii., 1869, p. 178.
  *(6) Dr. J. Barnard Davis on Oceanic Races, in Anthropological
Review, April, 1870, pp. 185, 191.

[Bullseye]

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 in 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.* I am informed
by Mr. D. Forbes, who particularly attended to this point, 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 so much
from each other, as in most other races.*(2) This fact is analogous
with what occurs with some closely allied monkeys; thus the sexes of
the chimpanzee are not as different as those of the orang or
gorilla.*(3)

  * Catlin, North American Indians, 3rd. ed., 1842, vol. ii., p.
227. On the Guaranys, see Azara, Voyages dans l'Amerique Merid.,
tom. ii., 1809, p. 85; also Rengger, Saugethiere von Paraguay, s. 3.
  *(2) 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.
  *(3) Rutimeyer, Die Grenzen der Thierwelt; eine Betrachtung zu
Darwin's Lehre, 1868, s. 54.

  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, have been transferred to the other. As this same form of
transmission has apparently prevailed much with mankind, it will
save useless repetition if we discuss the origin of characters
peculiar to the male sex together with certain other characters common
to both sexes.

  Law of Battle.- With savages, for instance, the Australians, the
women are the constant cause of war both between members of the same
tribe and between distinct tribes. So no doubt it was in ancient
times; "nam fuit ante Helenam mulier teterrima belli causa." With some
of the North American Indians, the contest is reduced to a system.
That excellent observer, Hearne,* 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 years old or
more, as before that age they cannot conquer their rivals.

  * A Journey from Prince of Wales Fort, 8vo ed., 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 South
America see Azara, Voyages, &c., tom. ii., p. 94.

  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,* 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 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 been reduced through disuse, as would the teeth through the not
well understood principles of correlation and 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 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 orang and gorilla 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 the
early male progenitors of man must have led to a most striking and
favourable change in his appearance.

  * On the fighting of the male gorillas, see Dr. Savage, in Boston
Journal of Natural History, vol. v., 1847, p. 423. On Presbytis
entellus, see the Indian Field, 1859, p. 146.

  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 his
half-human male ancestors. These characters would, however, have
been preserved or even augmented during the long ages of man's
savagery, by the success of the strongest and boldest men, both in the
general struggle for life and in their contests for wives; a success
which would have ensured their leaving a more numerous progeny than
their less favoured brethren. 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 joint 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 highly important part. I am aware
that some writers doubt whether there is any such inherent difference;
but this is at least probable from the analogy of the lower animals
which present other secondary sexual characters. No one disputes
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 would
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 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's attaining to a higher eminence, in whatever he takes
up, than can woman- 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 (inclusive both of 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 from averages, so well illustrated by
Mr. Galton, in his work on Hereditary Genius, that if men are
capable of a decided pre-eminence over women in many subjects, the
average of mental power in man must be above that of woman.
  Amongst the half-human progenitors of man, and amongst savages,
there have been struggles between the males 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 mankind, 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 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,
both possessed of every mental quality in equal perfection, save
that one has higher energy, perseverance, and courage, the latter will
generally become more eminent in every pursuit, and will gain the
ascendancy.* 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 can be gained in many
subjects. These latter faculties, as well as the former, 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 gained will have been transmitted more fully to the male
than to the female offspring. It accords in a striking manner with
this view of the modification and re-inforcement of many of our mental
faculties by sexual selection, that, firstly, they notoriously undergo
a considerable change at puberty,*(2) and, secondly, that eunuchs
remain throughout life inferior in these same qualities. Thus, man has
ultimately become superior to woman. It is, indeed, fortunate that the
law of the equal transmission of characters to both sexes prevails
with 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.

  * 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?
  *(2) Maudsley, Mind and Body, p. 31.

  It must be borne in mind that the tendency in characters acquired by
either sex late in life, to be transmitted to the same sex at the same
age, and of early acquired characters to be transmitted to both sexes,
are rules which, though general, do not always hold. If they always
held good, we might conclude (but I here exceed 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
in mental power between the sexes would 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. All women, however, could
not be thus raised, unless during many generations those who
excelled in the above robust virtues were married, and produced
offspring in larger numbers than other women. As before remarked of
bodily strength, although men do not now fight for their wives, and
this form of selection has passed away, yet during manhood, they
generally undergo 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.*

  * 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.

  Voice and Musical Powers.- In some species of Quadrumana there is
a great difference between the adult sexes, in the power of their
voices 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."* With respect to the
cause of this difference between the sexes, I have nothing to add to
the remarks 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,*(2) the voice and the form of the larynx differ in the different
races of mankind; but with the Tartars, Chinese, &c., the voice of the
male is said not to differ so much from that of the female, as in most
other races.

  * Owen, Anatomy of Vertebrates, vol. iii., p. 603.
  *(2) Journal of the Anthropological Society, April, 1869, pp.
lvii. and lxvi.

  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. The sounds
thus produced consist, I believe in all cases, of the same note,
repeated rhythmically;* and this is sometimes pleasing even to the
ears of man. The chief and, in some cases, exclusive purpose appears
to be either to call or charm the opposite sex.

  * Dr. Scudder, "Notes on Stridulation," in Proc. Boston Soc. of Nat.
Hist., vol. xi., April, 1868.

  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 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 lowest vertebrates
which breathe air are amphibians; and of these, 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 much 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. With other species both
sexes, or only the females, 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 the Hylobates agilis, an ape allied to man. This
gibbon has an extremely loud but musical voice. Mr. Waterhouse
states,* "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 a musician, confirms the
foregoing statement, and remarks, though erroneously, 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
other animals, it is probable that it uses its musical powers more
especially during the season of courtship.

  * Given in W. C. L. Martin's General Introduction to Natural History
of Mamm. Animals, 1841, p. 432; Owen, Anatomy of Vertebrates, vol.
iii, p. 600.

[Bullseye]

This gibbon is not the only species in the genus which sings, for my
son, Francis Darwin, attentively listened in the Zoological Gardens to
H. leuciscus whilst singing a cadence of three notes, in true
musical intervals and with a clear musical tone. It is a more
surprising fact that certain rodents utter musical sounds. Singing
mice have often been mentioned and exhibited, but imposture has
commonly been suspected. We have, however, at last a clear account
by a well-known observer, the Rev. S. Lockwood,* of the musical powers
of an American species, the Hesperomys cognatus, belonging to a
genus distinct from that of the English mouse. This little animal
was kept in confinement, and the performance was repeatedly heard.
In one of the two chief songs, "the last bar would frequently be
prolonged to two or three; and she would sometimes change from C sharp
and D, to C natural and D, then warble on these two notes awhile,
and wind up with a quick chirp on C sharp and D. The distinctness
between the semitones was very marked, and easily appreciable to a
good ear." Mr. Lockwood gives both songs in musical notation; and adds
that though this little mouse "had no ear for time, yet she would keep
to the key of B (two flats) and strictly in a major key." ... "Her
soft clear voice falls an octave with all the precision possible; then
at the wind up, it rises again into a very quick trill on C sharp
and D."

  * American Naturalist, 1871, p. 761.

  A critic has asked how the ears of man, and he ought to have added
of other animals, could have been adapted by selection so as to
distinguish musical notes. But this question shows some confusion on
the subject; a noise is the sensation resulting from the
co-existence of several aerial "simple vibrations" of various periods,
each of which intermits so frequently that its separate existence
cannot be perceived. It is only in the want of continuity of such
vibrations, and in their want of harmony inter se, that a noise
differs from a musical note. Thus, an ear to be capable of
discriminating noises- and the high importance of this power to all
animals is admitted by every one- must be sensitive to musical
notes. We have evidence of this capacity even low down in the animal
scale; thus, crustaceans are provided with auditory hairs of different
lengths, which have been seen to vibrate when the proper musical notes
are struck.* As stated in a previous chapter, similar observations
have been made on the hairs of the antennae of gnats. It has been
positively asserted by good observers that spiders are attracted by
music. It is also well known that some dogs howl when hearing
particular tones.*(2) 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."*(3)

  * Helmholtz, Theorie Phys. de la Musique, 1868, p. 187.
  *(2) Several accounts have been published to this effect. Mr.
Peach writes to me that an old dog of his howls when B flat is sounded
on the flute, and to no other note. I may add another instance of a
dog always whining, when one note on a concertina, which was out of
tune, was played.
  *(3) Mr. R. Brown, in Proc. Zool. Soc., 1868, p. 410.

  Therefore, as far as the mere perception of musical notes is
concerned, there seems no special difficulty in the case of man or
of any other animal. Helmholtz has explained on physiological
principles why concords are agreeable, and discords disagreeable to
the human ear; but we are little concerned with these, as music in
harmony is a late invention. We are more concerned with melody, and
here again, according to Helmholtz, it is intelligible why the notes
of our musical scale are used. The ear analyses all sounds into
their component "simple vibrations," although we are not conscious
of this analysis. In a musical note the lowest in pitch of these is
generally predominant, and the others which are less marked are the
octave, the twelfth, the second octave, &c., all harmonies of the
fundamental predominant note; any two notes of our scale have many
of these harmonic over-tones in common. It seems pretty clear then,
that if an animal always wished to sing precisely the same song, he
would guide himself by sounding those notes in succession, which
possess many overtones in common- that is, he would choose for his
song, notes which belong to our musical scale.
  But if it be further asked why musical tones in a certain order
and rhythm give man and other animals pleasure, we can no more give
the reason than for the pleasantness of certain tastes and smells.
That they do give pleasure of some kind to animals, we may infer
from their being produced during the season of courtship by many
insects, spiders, fishes, amphibians, and birds; for unless the
females were able to appreciate such sounds and were excited or
charmed by them, the persevering efforts of the males, and the complex
structures often possessed by them alone, would be useless; and this
it is impossible to believe.
  Human 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 use to man in
reference to his daily habits of life, they must be ranked amongst the
most mysterious with which he is endowed. They are present, though
in a very rude condition, in men of all races, even the most savage;
but so different is the taste of the several races, that our music
gives no pleasure to savages, and their music is to us in most cases
hideous and unmeaning. Dr. Seemann, in some interesting remarks on
this subject,* "doubt 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 singing gibbons, the capacity of producing, and therefore
no doubt of appreciating, musical notes, we know that man possessed
these faculties at a very remote period. M. Lartet has described two
flutes made out of the bones and horns of the reindeer, found in caves
together with flint tools and the remains of extinct animals. The arts
of singing and of dancing are also very ancient, and are now practised
by all or nearly all the lowest races of man. Poetry, which may be
considered as the offspring of song, is likewise so ancient, that many
persons have felt astonished that it should have arisen during the
earliest ages of which we have any record.

  * Journal of Anthropological Society, Oct., 1870, p. clv. See also
the several later chapters in Sir John Lubbock's Prehistoric Times,
2nd ed., 1869, which contain an admirable account of the habits of
savages.

  We see that the musical faculties, which are not wholly deficient in
any race, are capable of prompt and high development, for Hottentots
and Negroes have become excellent musicians, although in their
native countries they rarely practise anything that we should consider
music. Schweinfurth, however, was pleased with some of the simple
melodies which he heard in the interior of Africa. But there is
nothing anomalous in the musical faculties lying dormant in man:
some species of birds which never naturally sing, can without much
difficulty be taught to do so; thus a 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 possible that a progenitor of the
sparrow may have been a songster. It is more remarkable that
parrots, belonging to a group distinct from the Insessores, and having
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 very rash to assume
that parrots are descended from some ancient form which was a
songster. Many cases could be advanced of organs and instincts
originally adapted for one purpose, having been utilised for some
distinct purpose.* Hence the capacity for high musical development
which the savage races of man possess, may be due either to the
practice by our semi-human progenitors of some rude form of music,
or simply to their having acquired the proper vocal organs for a
different purpose. But in this latter ease we must assume, as in the
above instance of parrots, and as seems to occur with many animals,
that they already possessed some sense of melody.