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Descent of Man [ 1871 ]

Charles Darwin [ 1809 - 1882 ]

 

Chapter XVII - Secondary Sexual Characters of Mammals


  WITH mammals the male appears to win the female much more through
the law of battle than through the display of his charms. The most
timid animals, not provided with any special weapons for fighting,
engage in desperate conflicts during the season of love. Two male
hares have been seen to fight together until one was killed; male
moles often fight, and sometimes with fatal results; male squirrels
engage in frequent contests, "and often wound each other severely"; as
do male beavers, so that "hardly a skin is without scars."* I observed
the same fact with the hides of the guanacoes in Patagonia; and on one
occasion several were so absorbed in fighting that they fearlessly
rushed close by me. Livingstone speaks of the males of the many
animals in southern Africa as almost invariably shewing the scars
received in former contests.

  * See Waterton's account of two hares fighting, Zoologist, vol.
i., 1843, p. 211. On moles, Bell, Hist. of British Quadrupeds, 1st
ed., p. 100. On squirrels, Audubon and Bachman, Viviparous
Quadrupeds of N. America, 1846, p. 269. On beavers, Mr. A. H. Green,
in Journal of Linnean Society, Zoology, vol. x., 1869, p. 362.

  The law of battle prevails with aquatic as with terrestrial mammals.
It is notorious how desperately male seals fight, both with their
teeth and claws, during the breeding-season; and their hides are
likewise often covered with scars. Male sperm-whales are very
jealous at this season; and in their battles "they often lock their
jaws together, and turn on their sides and twist about"; so that their
lower jaws often become distorted.*

  * On the battles of seals, see Capt. C. Abbott in Proc. Zool.
Soc., 1868, p. 191; Mr. R. Brown, ibid., 1868, p. 436; also L.
Lloyd, Game Birds of Sweden, 1867, p. 414; also Pennant. On the
sperm-whale see Mr. J. H. Thompson, in Proc. Zool. Soc., 1867, p. 246.

  All male animals which are furnished with special weapons for
fighting, are well known to engage in fierce battles. The courage
and the desperate conflicts of stags have often been described;
their skeletons have been found in various parts of the world, with
the horns inextricably locked together, shewing how miserably the
victor and vanquished had perished.* No animal in the world is so
dangerous as an elephant in "must". Lord Tankerville has given me a
graphic description of the battles between the wild bulls in
Chillingham Park, the descendants, degenerated in size but not in
courage, of the gigantic Bos primigenius. In 1861 several contended
for mastery; and it was observed that two of the younger bulls
attacked in concert the old leader of the herd, overthrew and disabled
him, so that he was believed by the keepers to be lying mortally
wounded in a neighbouring wood. But a few days afterwards one of the
young bulls approached the wood alone; and then the "monarch of the
chase," who had been lashing himself up for vengeance, came out and,
in a short time, killed his antagonist. He then quietly joined the
herd, and long held undisputed sway. Admiral Sir B. J. Sulivan informs
me that, when he lived in the Falkland Islands, he imported a young
English stallion, which frequented the hills near Port William with
eight mares. On these hills there were two wild stallions, each with a
small troop of mares; "and it is certain that these stallions would
never have approached each other without fighting. Both had tried
singly to fight the English horse and drive away his mares, but had
failed. One day they came in together and attacked him. This was
seen by the captain who had charge of the horses, and who, on riding
to the spot, found one of the two stallions engaged with the English
horse, whilst the other was driving away the mares, and had already
separated four from the rest. The captain settled the matter by
driving the whole party into a corral, for the wild stallions would
not leave the mares."

  * See Scrope (Art of Deer-stalking, p. 17) on the locking of the
horns with the Cervus elaphus. Richardson, in Fauna Bor. Americana,
1829, p. 252, says that the wapiti, moose, and reindeer have been
found thus locked together. Sir. A. Smith found at the Cape of Good
Hope the skeletons of two gnus in the same condition.

  Male animals which are provided with efficient cutting or tearing
teeth for the ordinary purposes of life, such as the Carnivora,
Insectivora, and rodents, are seldom furnished with weapons especially
adapted for fighting with their rivals. The case is very different
with the males of many other animals. We see this in the horns of
stags and of certain kinds of antelopes in which the females are
hornless. With many animals the canine teeth in the upper or lower
jaw, or in both, are much larger in the males than in the females,
or are absent in the latter, with the exception sometimes of a
hidden rudiment. Certain antelopes, the musk-deer, camel, horse, boar,
various apes, seals, and the walrus, offer instances. In the females
of the walrus the tusks are sometimes quite absent.* In the male
elephant of India and in the male dugong*(2) the upper incisors form
offensive weapons. In the male narwhal the left canine alone is
developed into the well-known, spirally-twisted, so-called horn, which
is sometimes from nine to ten feet in length. It is believed that
the males use these horns for fighting together; for "an unbroken
one can rarely be got, and occasionally one may be found with the
point of another jammed into the broken place."*(3) The tooth on the
opposite side of the head in the male consists of a rudiment about ten
inches in length, which is embedded in the jaw; but sometimes,
though rarely, both are equally developed on the two sides. In the
female both are always rudimentary. The male cachalot has a larger
head than that of the female, and it no doubt aids him in his
aquatic battles. Lastly, the adult male Ornithorhynchus is provided
with a remarkable apparatus, namely a spur on the foreleg, closely
resembling the poison-fang of a venomous snake; but according to
Harting, the secretion from the gland is not poisonous; and on the leg
of the female there is a hollow, apparently for the reception of the
spur.*(4)

  * Mr. Lamont (Seasons with the Sea-Horses, 1861, p. 143) says that a
good tusk of the male walrus weighs 4 pounds, and is longer than
that of the female, which weighs about 3 pounds. The males are
described as fighting ferociously. On the occasional absence of the
tusks in the female, see Mr. R. Brown, Proceedings, Zoological
Society, 1868, p. 429.
  *(2) Owen, Anatomy of Vertebrates, vol. iii., p. 283.
  *(3) Mr. R. Brown, in Proc. Zool. Soc., 1869, p. 553. See Prof.
Turner, in Journal of Anat. and Phys., 1872, p. 76, on the homological
nature of these tusks. Also Mr. J W. Clarke on two tusks being
developed in the males, in Proceedings of the Zoological Society,
1871, p. 42.
  *(4) Owen on the cachalot and Ornithorhynchus, ibid., vol. iii., pp.
638, 641. Harting is quoted by Dr. Zouteveen in the Dutch
translation of this work, vol. ii., p. 292.

  When the males are provided with weapons which in the females are
absent, there can be hardly a doubt that these serve for fighting with
other males; and that they were acquired through sexual selection, and
were transmitted to the male sex alone. It is not probable, at least
in most cases, that the females have been prevented from acquiring
such weapons, on account of their being useless, superfluous, or in
some way injurious. On the contrary, as they are often used by the
males for various purposes, more especially as a defence against their
enemies, it is a surprising fact that they are so poorly developed, or
quite absent, in the females of so many animals. With female deer
the development during each recurrent season of great branching horns,
and with female elephants the development of immense tusks, would be a
great waste of vital power, supposing that they were of no use to
the females. Consequently, they would have tended to be eliminated
in the female through natural selection; that is, if the successive
variations were limited in their transmission to the female sex, for
otherwise the weapons of the males would have been injuriously
affected, and this would have been a greater evil. On the whole, and
from the consideration of the following facts, it seems probable
that when the various weapons differ in the two sexes, this has
generally depended on the kind of transmission which has prevailed.
  As the reindeer is the one species in the whole family of deer, in
which the female is furnished with horns, though they are somewhat
smaller, thinner, and less branched than in the male, it might
naturally be thought that, at least in this case, they must be of some
special service to her. The female retains her horns from the time
when they are fully developed, namely, in September, throughout the
winter until April or May, when she brings forth her young. Mr. Crotch
made particular enquiries for me in Norway, and it appears that the
females at this season conceal themselves for about a fortnight in
order to bring forth their young, and then reappear, generally
hornless. In Nova Scotia, however, as I hear from Mr. H. Reeks, the
female sometimes retains her horns longer. The male on the other
hand casts his horns much earlier, towards the end of November. As
both sexes have the same requirements and follow the same habits of
life, and as the male is destitute of horns during the winter, it is
improbable that they can be of any special service to the female
during this season, which includes the larger part of the time
during which she is horned. Nor is it probable that she can have
inherited horns from some ancient progenitor of the family of deer,
for, from the fact of the females of so many species in all quarters
of the globe not having horns, we may conclude that this was the
primordial character of the group.*

  * On the structure and shedding of the horns of the reindeer,
Hoffberg, Amaenitates Acad., vol. iv., 1788, p. 149. See Richardson,
Fauna Bor. Americana,. p. 241, in regard to the American variety or
species: also Major W. Ross King, The Sportsman in Canada, 1866, p.
80.

  The horns of the reindeer are developed at a most unusually early
age; but what the cause of this may be is not known. The effect has
apparently been the transference of the horns to both sexes. We should
bear in mind that horns are always transmitted through the female, and
that she has a latent capacity for their development, as we see in old
or diseased females.* Moreover the females of some other species of
deer exhibit, either normally or occasionally, rudiments of horns;
thus the female of Cervulus moschatus has "bristly tufts, ending in
a knob, instead of a horn"; and "in most specimens of the female
wapiti (Cervus canadensis) there is a sharp bony protuberance in the
place of the horn."*(2) From these several considerations we may
conclude that the possession of fairly well-developed horns by the
female reindeer, is due to the males having first acquired them as
weapons for fighting with other males; and secondarily to their
development from some unknown cause at an unusually early age in the
males, and their consequent transference to both sexes.

  * Isidore Geoffroy St-Hilaire, Essais de Zoolog. Generale, 1841,
p. 513. Other masculine characters, besides the horns, are sometimes
similarly transferred to the female; thus Mr. Boner, in speaking of an
old female chamois (Chamois Hunting in the Mountains of Bavaria, 1860,
2nd ed., p. 363), says, "not only was the head very male-looking,
but along the back there was a ridge of long hair, usually to be found
only in bucks."
  *(2) On the Cervulus, Dr. Gray, Catalogue of Mammalia in the British
Museum, part iii., p. 220. On the Cervus canadensis or wapiti, see
Hon. J. D. Caton, Ottawa Academy of Nat. Sciences, May, 1868, p. 9.

  Turning to the sheath-horned ruminants: with antelopes a graduated
series can be formed, beginning with species, the females of which are
completely destitute of horns- passing on to those which have horns so
small as to be almost rudimentary (as with the Antilocapra
americana, in which species they are present in only one out of four
or five females*)- to those which have fairly developed horns, but
manifestly smaller and thinner than in the male and sometimes of a
different shape,*(2)- ending with those in which both sexes have horns
of equal size. As with the reindeer, so with antelopes, there
exists, as previously shewn, a relation between the period of the
development of the horns and their transmission to one or both
sexes; it is therefore probable that their presence or absence in
the females of some species, and their more or less perfect
condition in the females of other species, depends, not on their being
of any special use, but simply on inheritance. It accords with this
view that even in the same restricted genus both sexes of some
species, and the males alone of others, are thus provided. It is
also a remarkable fact that, although the females of Antilope
bezoartica are normally destitute of horns, Mr. Blyth has seen no less
than three females thus furnished; and there was no reason to
suppose that they were old or diseased.

  * I am indebted to Dr. Canfield for this information; see also his
paper in the Proceedings of the Zoological Society, 1866, p. 105.
  *(2) For instance the horns of the female Ant. euchore resemble
those of a distinct species, viz. the Ant. dorcas var. corine, see
Desmarest, Mammalogie, p. 455.

  In all the wild species of goats and sheep the horns are larger in
the male than in the female, and are sometimes quite absent in the
latter.* In several domestic breeds of these two animals, the males
alone are furnished with horns; and in some breeds, for instance, in
the sheep of North Wales, though both sexes are properly horned, the
ewes are very liable to be hornless. I have been informed by a
trustworthy witness, who purposely inspected a flock of these same
sheep during the lambing season, that the horns at birth are generally
more fully developed in the male than in the female. Mr. J. Peel
crossed his Lonk sheep, both sexes of which always bear horns, with
hornless Leicesters and hornless Shropshire Downs; and the result
was that the male offspring had their horns considerably reduced,
whilst the females were wholly destitute of them. These several
facts indicate that, with sheep, the horns are a much less firmly
fixed character in the females than in the males; and this leads us to
look at the horns as properly of masculine origin.

  * Gray, Catalogue of Mammalia, the British Museum, part iii.,
1852, p. 160.

  With the adult musk-ox (Ovibos moschatus) the horns of the male
are larger than those of the female, and in the latter the bases do
not touch.* In regard to ordinary cattle Mr. Blyth remarks: "In most
of the wild bovine animals the horns are both longer and thicker in
the bull than in the cow, and in the cow-banteng (Bos sondaicus) the
horns are remarkably small, and inclined much backwards. In the
domestic races of cattle, both of the humped and humpless types, the
horns are short and thick in the bull, longer and more slender in
the cow and ox; and in the Indian buffalo, they are shorter and
thicker in the bull, longer and more slender in the cow. In the wild
gaour (B. gaurus) the horns are mostly both longer and thicker in
the bull than in the cow."*(2) Dr. Forsyth Major also informs me
that a fossil skull, believed to be that of the female Bos
estruscus, has been found in Val d'Arno, which is wholly without
horns. In the Rhinoceros simus, as I may add, the horns of the
female are generally longer but less powerful than in the male; and in
some other species of rhinoceros they are said to be shorter in the
female.*(3) From these various facts we may infer as probable that
horns of all kinds, even when they are equally developed in the two
sexes, were primarily acquired by the male in order to conquer other
males, and have been transferred more or less completely to the
female.

  * Richardson, Fauna Bor. Americana, p. 278.
  *(2) Land and Water, 1867, p. 346.
  *(3) Sir Andrew Smith, Zoology of S. Africa, pl. xix. Owen,
Anatomy of Vertebrates, vol. iii., p. 624.

  The effects of castration deserve notice, as throwing light on
this same point. Stags after the operation never renew their horns.
The male reindeer, however, must be excepted, as after castration he
does renew them. This fact, as well as the possession of horns by both
sexes, seems at first to prove that the horns in this species do not
constitute a sexual character;* but as they are developed at a very
early age, before the sexes differ in constitution, it is not
surprising that they should be unaffected by castration, even if
they were aboriginally acquired by the male. With sheep both sexes
properly bear horns; and I am informed that with Welch sheep the horns
of the males are considerably reduced by castration; but the degree
depends much on the age at which the operation is performed, as is
likewise the case with other animals. Merino rams have large horns,
whilst the ewes "generally speaking are without horns"; and in this
breed castration seems to produce a somewhat greater effect, so that
if performed at an early age the horns "remain almost
undeveloped."*(2)

  * This is the conclusion of Seidlitz, Die Darwin'sche Theorie, 1871,
p. 47.
  *(2) I am much obliged to Prof. Victor Carus, for having made
enquiries for me in Saxony on this subject. H. von Nathusius
(Viehzucht, 1872, p. 64) says that the horns of sheep castrated at
an early period, either altogether disappear or remain as mere
rudiments; but I do not know whether he refers to merinos or to
ordinary breeds.

  On the Guinea coast there is a breed in which the females never bear
horns, and, as Mr. Winwood Reade informs me, the rams after castration
are quite destitute of them. With cattle, the horns of the males are
much altered by castration; for instead of being short and thick, they
become longer than those of the cow, but otherwise resemble them.
The Antilope bezoartica offers a somewhat analogous case: the males
have long straight spiral horns, nearly parallel to each other, and
directed backwards; the females occasionally bear horns, but these
when present are of a very different shape, for they are not spiral,
and spreading widely, bend round with the points forwards. Now it is a
remarkable fact that, in the castrated male, as Mr. Blyth informs
me, the horns are of the same peculiar shape as in the female, but
longer and thicker. If we may judge from analogy, the female
probably shews us, in these two cases of cattle and the antelope,
the former condition of the horns in some early progenitor of each
species. But why castration should lead to the reappearance of an
early condition of the horns cannot be explained with any certainty.
Nevertheless, it seems probable, that in nearly the same manner as the
constitutional disturbance in the offspring, caused by a cross between
two distinct species or races, often leads to the reappearance of
long-lost characters;* so here, the disturbance in the constitution of
the individual, resulting from castration, produces the same effect.

  * I have given various experiments and other evidence proving that
this is the case, in my Variation of Animals and Plants under
Domestication, vol. ii., 1868, pp. 39-47.

  The tusks of the elephant, in the different species or races, differ
according to sex, nearly as do the horns of ruminants. In India and
Malacca the males alone are provided with well-developed tusks. The
elephant of Ceylon is considered by most naturalists as a distinct
race, but by some as a distinct species, and here "not one in a
hundred is found with tusks, the few that possess them being
exclusively males."* The African elephant is undoubtedly distinct, and
the female has large well-developed tusks, though not so large as
those of the male.

  * Sir J. Emerson Tennent, Ceylon, 1859, vol. ii., p. 274. For
Malacca, Journal of Indian Archipelago, vol. iv., p. 357.

  These differences in the tusks of the several races and species of
elephants- the great variability of the horns of deer, as notably in
the wild reindeer- the occasional presence of horns in the female
Antilope bezoartica, and their frequent absence in the female of
Antilocapra americana- the presence of two tusks in some few male
narwhals- the complete absence of tusks in some female walruses- are
all instances of the extreme variability of secondary sexual
characters, and of their liability to differ in closely-allied forms.
  Although tusks and horns appear in all cases to have been
primarily developed as sexual weapons, they often serve other
purposes. The elephant uses his tusks in attacking the tiger;
according to Bruce, he scores the trunks of trees until they can be
thrown down easily, and he likewise thus extracts the farinaceous
cores of palms; in Africa he often uses one tusk, always the same,
to probe the ground and thus ascertain whether it will bear his
weight. The common bull defends the herd with his horns; and the elk
in Sweden has been known, according to Lloyd, to strike a wolf dead
with a single blow of his great horns. Many similar facts could be
given. One of the most curious secondary uses to which the horns of an
animal may be occasionally put is that observed by Captain Hutton*
with the wild goat (Capra aegagrus) of the Himalayas and, as it is
also said with the ibex, namely that when the male accidentally
falls from a height he bends inwards his head, and by alighting on his
massive horns, breaks the shock. The female cannot thus use her horns,
which are smaller, but from her more quiet disposition she does not
need this strange kind of shield so much.

  * Calcutta Journal of Natural History, vol. ii, 1843, p. 526.

  Each male animal uses his weapons in his own peculiar fashion. The
common ram makes a charge and butts with such force with the bases
of his horns, that I have seen a powerful man knocked over like a
child. Goats and certain species of sheep, for instance the Ovis
cycloceros of Afghanistan,* rear on their hind legs, and then not only
butt, but "make a cut down and a jerk up, with the ribbed front of
their scimitar-shaped horn, as with a sabre. When the O. cycloceros
attacked a large domestic ram, who was a noted bruiser, he conquered
him by the sheer novelty of his mode of fighting, always closing at
once with his adversary, and catching him across the face and nose
with a sharp drawing jerk of the head, and then bounding out of the
way before the blow could be returned." In Pembrokeshire a male
goat, the master of a flock which during several generations had run
wild, was known to have killed several males in single combat; this
goat possessed enormous horns, measuring thirty-nine inches in a
straight line from tip to tip. The common bull, as every one knows,
gores and tosses his opponent; but the Italian buffalo is said never
to use his horns: he gives a tremendous blow with his convex forehead,
and then tramples on his fallen enemy with his knees- an instinct
which the common bull does not possess.*(2) Hence a dog who pins a
buffalo by the nose is immediately crushed. We must, however, remember
that the Italian buffalo has been long domesticated, and it is by no
means certain that the wild parent-form had similar horns. Mr.
Bartlett informs me that when a female Cape buffalo (Bubalus caffer)
was turned into an enclosure with a bull of the same species, she
attacked him, and he in return pushed her about with great violence.
But it was manifest to Mr. Bartlett that, had not the bull shewn
dignified forbearance, he could easily have killed her by a single
lateral thrust with his immense horns. The giraffe uses his short,
hair-covered horns, which are rather longer in the male than in the
female, in a curious manner; for, with his long neck he swings his
head to either side, almost upside down, with such force that I have
seen a hard plank deeply indented by a single blow.

  * Mr. Blyth, in Land and Water, March, 1867, p. 134, on the
authority of Capt. Hutton and others. For the wild Pembrokeshire
goats, see the Field, 1869, p. 150.
  *(2) M. E. M. Bailly, "Sur l'Usage des cornes," &c., Annal des
Sciences Nat., tom. ii., 1824, p. 369.

  With antelopes it is sometimes difficult to imagine how they can
possibly use their curiously-shaped horns; thus the springboc (Ant.
euchore) has rather short upright horns, with the sharp points bent
inwards almost at right angles, so as to face each other; Mr. Bartlett
does not know how they are used, but suggests that they would
inflict a fearful wound down each side of the face of an antagonist.
The slightly-curved horns of the Oryx leucoryx (see fig. 63) are
directed backwards, and are of such length that their points reach
beyond the middle of the back, over which they extend in almost
parallel lines. Thus they seem singularly ill-fitted for fighting; but
Mr. Bartlett informs me that when two of these animals prepare for
battle, they kneel down, with their beads between their forelegs,
and in this attitude the horns stand nearly parallel and close to
the ground, with the points directed forwards and a little upwards.
The combatants then gradually approach each other, and each endeavours
to get the upturned points under the body of the other; if one
succeeds in doing this, he suddenly springs up, throwing up his head
at the same time, and can thus wound or perhaps even transfix his
antagonist. Both animals always kneel down, so as to guard as far as
possible against this manoeuvre. It has been recorded that one of
these antelopes has used his horn with effect even against a lion; yet
from being forced to place his head between the fore legs in order
to bring the points of the horns forward, he would generally be
under a great disadvantage when attacked by any other animal. It is,
therefore, not probable that the horns have been modified into their
present great length and peculiar position, as a protection against
beasts of prey. We can however see that, as soon as some ancient
male progenitor of the Oryx acquired moderately long horns, directed a
little backwards, he would be compelled, in his battles with rival
males, to bend his head somewhat inwards or downwards, as is now
done by certain stags; and it is not improbable that he might have
acquired the habit of at first occasionally and afterwards of
regularly kneeling down. In this case it is almost certain that the
males which possessed the longest horns would have had a great
advantage over others with shorter horns; and then the horns would
gradually have been rendered longer and longer, through sexual
selection, until they acquired their present extraordinary length
and position.
  With stags of many kinds the 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.

  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.

  * 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 bitch-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.

  * 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.

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