AT DAGGERS DRAWN
The Birth of Weapons—All Flesh is Grass—Utility and Ornament—The
Fever of Love—The “Challenge” of the Deer—What it means—More about
“Hormones”—“Hummel” Stags—The Age of Deer—The “Courtship” of the
Moose—Types of Antlers—Antlered Females—Fighting Topi—The Lance of
the Oryx in the Lion’s Flanks—Happiness and Hartebeestes—Odoriferous
Suitors—The Bloody Sweat of the Hippopotamus—The Elephant in
Love—Concerning Tusks—Polygamy.
From Apes to Antelopes is a far cry, but contrasts are always helpful.
Antelopes and Deer, Zebras and Elephants, Rhinoceroses and Swine, are
types, taken at random, of that great and important group of animals
known as the “Ungulates,” or “Hoofed” animals. These illustrate in
a very striking manner what is meant by the term “Secondary Sexual
Characters.” They demonstrate no less forcibly what is meant by the
term “Sexual Selection.” They are valuable in this connection, because
of the often formidable weapons, in the shape of horns and tusks, which
so many species have developed during the struggle for mates.
But “Sexual Selection” will not explain their origin, and it is
difficult, in the present state of our knowledge, to discover any
clues which will reveal this. In seeking these there are certain broad
aspects of the problem which are not to be lost sight of. In the first
place, horns, at any rate, are confined to the hoofed animals. That the
various types of hoofed animals, living and extinct, have had a common
ancestry, no one at the present day will probably call in question. The
relationship, however, of the various living types, one to another, is
by no means always apparent: the missing links are to be sought in the
records of the rocks.
When the whole of the evidence comes to be surveyed, and not till
then, it becomes apparent that this wonderful diversity is the result
of complex factors. That the conditions of existence have controlled
the results is beyond question; but it is equally certain that these
conditions have been merely controlling and not causative. In other
words, we must regard each of these different groups or types—Deer,
Antelopes, Horses, Elephants, Swine, and so on—as witnesses of what
we call “Heredity.” They are so many “Diathetic types.” That is to
say, the forms, or individuals, belonging to each type have inherited
certain peculiarities in common; they display a “Diathesis” as the
doctors call it: an inherent, inborn tendency, or habit of growth, in
a definite direction: a tendency which, ever and anon, develops new
qualities, takes new directions. And thus it is that we get Oxen—using
this term in its widest sense and not in its special sense—Antelopes,
Goats and Sheep, for example. These have, among other things, a
“diathesis” in the direction of horn production, and each, too, of
a different type. What is meant by this apparently mystifying term
“diathesis” will perhaps be made clear by taking the case of the Ox and
the Sheep. While very different in appearance, these live on precisely
similar food; yet no one has any difficulty in discriminating between
the taste of beef and mutton. In the marvellous chemical laboratory of
the body the grass gathered in the same field is converted into flesh
which even in its uncooked state is easily distinguishable. Though for
the purposes of this illustration domesticated animals have been used,
the same is true of their wild relations. Sportsmen tell us that the
various types of Antelopes and the Zebra, which may be seen feeding
together, have yet flesh of very different qualities. These qualities
are to be attributed neither to “Natural” nor to “Sexual” selection;
they are “accidents.” Similarly, their horns are the witness of a
horn-producing “diathesis”: the various divergencies in curvature,
and in the form of their spirals, or the number of their encircling
rings—as in the horns of Antelopes—are to be interpreted in like
fashion. These twists and turns vary in the same way that the taste of
the flesh varies, and for the same reason; that is to say, they are not
the outcome of “Sexual Selection,” nor have they been brought about by
“Natural Selection” to serve the purpose of “Recognition marks,” as
Wallace would have us believe.
But horns, as horns, apart from their “accidents” of curvature
and ornament, must certainly be regarded as the product of Sexual
selection, for having once started into being those individuals had
the best chance of leaving descendants which were best armed. The
possession of horns was not necessary to the maintenance of the
species; but such armature was essential among the males in securing
possession of the females. Other things being equal, the male with the
biggest horns wins the prize. Since these are also used as weapons of
offence, or rather of defence, in warding off the attacks of beasts
of prey, it might be contended that they are as much the product of
Natural selection as of Sexual selection.
It soon becomes apparent that this interpretation must fail. In the
first place, if it were true, the females should be similarly armed.
In the second, in the presence of many of their enemies they are
useless. The Cape hunting-dog, for example, is more than a match for
any antelope. This ferocious animal kills his victim by running it
down, persistently tearing at its flanks, until at last the entrails
protrude and the horrid chase is ended. Furthermore, the horns are a
comparatively late acquirement of the species, as is shown in the case
of the Deer; for the earliest known fossil species were hornless. That
the females among the Oxen and many of the Antelopes possess horns is
an interesting fact, but it can only be regarded as another instance
of a character first acquired by the male and later, in successive
generations, transferred to the female. And it is to be noticed that
this transference is never found save in the cases where the character
in question has attained its maximum in the male. The transference
of weapons to the female is the more remarkable because there is no
evidence that they play any part in the struggle for existence, either
in securing mates or in warding off the attacks of enemies. Moreover,
these weapons in the female may exceed those of the male, in length,
though they are never so massive. They are to be regarded solely in
the light of ornaments. There are few more striking instances indeed
where the purely ornamental and the strictly utilitarian are so closely
associated.
[Illustration: Plate 4.
_By the courtesy of Rowland, Ward, Ltd._
WEAPONS OF OFFENCE.
Horns of various types furnish the most conspicuous of the “Secondary
Sexual Characters” of the ruminants. In the Deer only are these
branched. In the “hollow-horned” ruminants they are either lance-like
or more or less spirally curved, or they may form more or less open
loops.
1. Black-tailed Deer. 2. Hangul or Kashmir Barasingha Deer.
3. Greater Kudu. 4. Black-buck. 5. Saiga Antelope, remarkable also for
its curiously swollen nose. 6. Marco-Polo’s Sheep.
[Face page 52.]
Attention may now profitably be turned to the behaviour of these
interesting tribes when under the alluring influences of love.
Tradition and the poets have contrived to persuade us that the fever
of Love becomes epidemic in the spring. This, however, is by no means
true, at any rate in so far as what we are pleased to call the “lower
animals” are concerned. For with many, as for example the Deer and the
Bats, this fever is not aroused till the time of autumn plenty. With
regard to the deer, we can find a reason for this. It is determined in
part by the period of gestation, and in part by the peculiar character
of the most conspicuous of the male secondary sexual characters—the
antlers. The deer, at any rate of the northern hemisphere, carry their
young about eight months. Now it is important that they should make
their entry into the world just as the food supply is increasing and
the temperature is rising. With the summer before them the young have
time to gather strength for the encounter with their first winter. We
have a striking witness to the truth of this contention in the fact
that when the Indian Spotted Deer, or Chital, was first introduced into
Europe, nearly all the fawns perished owing to having been born in
winter; later, the females took to calving in spring, and from thence
onwards the species has held its own among us.
As touching the stags. The antlers, as everybody knows, are shed
annually, and their renewal entails a very considerable strain on the
system. As a consequence, it is necessary that this period of stress
should fall after the trial of winter is overpast, and with the genial
summer before them. From the end of March, when the old weapons are
shed, till July, the masterful males of the community wander at large,
seeking seclusion and avoiding all occasion of quarrel; for they
are not only defenceless, but threatened with disaster should any
accident befall the growing horns, which, during their formation, are
exceedingly sensitive. Even a slight blow would not only spoil their
shapely proportions, but, further, might render them useless in the
warfare that is before them.
With some species this desire to go into retreat is more marked than in
others. The Red-deer, and the Wapiti, on the one hand, and the Moose on
the other, well illustrate this. The two first-named pass the winter in
herds, in the case of the Wapiti numbering many thousand individuals;
no other species, indeed, is so markedly gregarious. With the advance
of the spring, however, all is changed, for the males withdraw from
their companions to suffer humiliation in seclusion. As chill October
arrives, a striking alteration in their demeanour becomes apparent,
at any rate in the case of the older males. The new antlers are now
hardened, and the blood supply, which has hitherto been building up the
new weapons, is cut off. As a consequence, the “velvet,” which till now
has been directly concerned with the growth of the antlers, dies, and
peels off the underlying bone. To facilitate this work of cleaning,
the animal rubs them, first against the stems of saplings, and, later,
against larger trees, and even rocks, till at last they are ready for
“battle, murder and sudden death.” The “rutting” season, in short, has
commenced. And with the final completion of the antlers other signs
of that approaching frenzy, which is soon to establish itself, become
apparent. The most striking of these are the swelling of the neck, and
a marked increase in the mane thereof; while the voice enlarges its
compass enormously, whereby the females, so long neglected, are now
feverishly sought for.
[Illustration: Plate 5.
_Photo by G. W. Wilson Co. Ltd., from “The Living Animals of the World.”_
MANCHURIAN WAPITI “CALLING.”
The “stags” do not begin to call for mate’s until the horns have more
or less completely shed their velvet.
[Face page 54.]
The Red-deer, maddened with desire, scours the country, calling as he
travels with a loud musical roar, ever and anon impatiently listening
for the tremulous response of females hardly less anxious to mate
than himself. One after another is speedily added to his harem, but not
without conflict. For sooner or later he catches the call of another
stag in like case. A jealous fury at once takes possession of him, and
the call, intended as a message to mateless hinds, becomes translated
into a challenge to fight for the mates possessed. Each of the now
infuriated challengers makes all haste to come to blows, and speedily
they are rushing headlong on one another to meet in a crash of antlers.
Then follows a test of strength, a sort of tug-of-war reversed, for
each strives to push the other to his knees, and succeeding, to deal
a deadly sideways thrust at the kneeling adversary’s heart with the
spike-shaped brow-tines. This attempt, however, is rarely achieved. Yet
not seldom such encounters become a duel to the death, and one in which
both die, for in the remorseless tilt at one another the antlers of one
may spring apart, and then close in on those of the other. Once this
happens, it seems to be rare indeed that they can be extricated from
this close embrace. With heads thus locked, they sway, and twist, and
tug, not now for the mastery, but for life itself. But as the hours run
they become more and more exhausted by their efforts, weaker and weaker
from loss of food and rest, till finally death releases both.
A male having once succeeded in forming a harem, will commonly contrive
to repeat his success year after year, withstanding all comers. But
sooner or later his vigour wanes and he is ousted by another and
younger male. Not else would the stamina of the race be preserved. It
is considered a moot point, however, whether physical strength and
sexual potency run at the same pace; for it is believed by some that
a stag will often contrive to hold a harem against all rivals after
his fertility has declined. This, however, is extremely improbable. A
lowering of fertility means a decline in the potency of the hormones,
and in the development of the secondary sexual characters, among which
are the antlers, which are by no means negligible factors. That they
are not all-important, however, seems to be shown by the fact that,
occasionally, stags appear in a herd which are congenitally unable to
produce antlers—a reversion to the ancestral condition—and such are
said, occasionally at any rate, to be able to oust their formidably
armed rivals. This may be so, but the fact that “hummel” stags, as they
are called, are so rare is surely to be regarded as eloquent testimony
of the disadvantages of their unarmoured state. They become speedily
eliminated, in short, by “Sexual Selection.”
After this outburst of sexual activity has spent itself, the various
harems, with their lords, amalgamate; all living in peace through the
winter. The stags retain their antlers at this season, partly as a
protection against predatory enemies, such as wolves, and other large
carnivores, which would otherwise play havoc in their ranks, and partly
because the cold of winter and scanty fodder would inhibit the growth
of new antlers or reduce their size. With the return of spring the
dangers of attack are lessened, temperature rises, and food becomes
once more plentiful. Then the inevitable disarmament takes place.
The Red-deer, though mature at six, does not reach his prime till his
eleventh year, and from thence till his fifteenth or sixteenth year is
at his best. The hinds mature earlier, and appear to be fertile for a
much longer period. At any rate, a wild hind in Jura, known by certain
peculiarities of its ears, during twenty-one years produced twenty
calves. She was killed at last with a calf at her side, but was thin
and haggard-looking. She was, therefore, not less than six-and-twenty
at her death. The calves, it may be mentioned, are born in May and June.
Old stags shed their antlers, it is remarked, earlier than young ones.
And this is an advantage to the species, since it prevents premature
breeding on the part of sexually precocious but immature males, and
limits competition to the adults.
What obtains in the case of the Red-deer obtains also with minor
variations due to environment, climate, and so on, in the case of all
other deer. The life-history of the Wapiti, as might be supposed,
differs only in detail from that of the Red-deer. But during the winter
they form vast herds, numbering thousands. It may be that in primitive
times the Red-deer was no less numerous. But in this country, at any
rate, conditions favourable to the maximum development, either in
bodily size, or in the massiveness of the antlers, have long since
passed away. Even in the Highlands of Scotland the conditions of
existence have entirely changed owing to disafforestation. Deer are
essentially forest dwellers. But the “deer forests” are such only in
name, and for the most part the wild stags of to-day must get what
shelter they can from rocks and inequalities of the ground. From
this cause, and from the very natural desire of the owners of such
“forests” to secure the finest heads in each year, the whole race
has deteriorated. How great a change has come over it may be seen by
comparing the heads of British Stags with those from German forests,
where the conditions of existence are more favourable. If we turn to
the records of the past we find that the antlers found in the fens,
turbaries, and caverns of our islands are vastly larger, heavier, and
carry a greater number of points on the sur-royals, than do those of
the existing Scotch stags.
Having regard to the fact that hundreds, and in the distant past
thousands, of antlers were shed annually, the comparative rarity of
these weapons in the haunts of deer excites comment. This is accounted
for by the fact that they are greedily eaten by their late owners,
apparently, though unconsciously, for the sake of their bone-producing
qualities.
By way of contrast with the Red-deer and Wapiti, we may take the Moose
(_Alces machlis_), which at no time, and nowhere, attains to large
herds. This is explained by the relatively restricted food supply which
obtains in the haunts of these creatures. For they frequent the margins
of streams, feeding largely on willows and birch. From the shortness
of their necks, and the great length of their legs, they cannot crop
grass and other short herbage, for unless they kneel they cannot reach
the ground. Hence it is obvious that though their geographical range
may be wide, their numbers are kept rigidly in check. They would be
fewer still but for the fact that, unlike other deer, they glean no
small amount of food from the water, wading out to feed upon aquatic
vegetation. The roots of water-lilies are especially sought for, and to
obtain these the animal will often disappear entirely under water.
As a consequence of the limited food supply the Moose lead solitary
lives. On the Eastern side of America, where the winter is severe, a
few individuals, generally a family party, will “yard up,” or make a
fortress for their mutual protection by trampling down the snow over a
restricted area. But in the Yukon district, my friend Mr. F. C. Selous
tells me this is never done.
The rutting season of the bulls begins as soon as the antlers begin
to “peel.” What follows is practically a repetition of what has
already been related in regard to the Red-deer and Wapiti. And in this
connection it is interesting to note that the natives take advantage of
the period of desire in the bull to entice him to his death. Generally
this is done by imitating the call of the cow in response to the bull’s
anxious bellowing. But in Southern Alaska the opposite side of his
nature is played upon. This is done by scraping or beating the bushes
with the shoulder-blade of a Moose in such a way as to reproduce the
sound of a bull cleaning his horns. The very suspicion of a rival
enrages him, and, rushing in a blind fury in the direction of the
tell-tale sounds, he speedily falls a victim to the trick which has
been played him.
That the mating period is the most critical, and most searching in
the whole life-history there can be no doubt. Every faculty during
this time is put to the test, and from the time of sexual maturity
until old age is at last attained it is an annual test. Alertness is
all important. Other things being equal, success falls most certainly
to those individuals with the keenest perception, and quickest
interpretation of sight, sound and smell.
One is puzzled at what seems a concession of Darwin’s to the Lamarckian
theory of the inherited effects of use in this connection. For in
discussing the bellowing of the stag in “The Descent of Man,” he
remarks that it “does not seem to be of any special service to him,
either during 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?” All the evidence goes to show
that the production of sound, and the instant interpretation of its
significance, is a matter of the highest importance. In the case of the
Moose, for example, the noise occasioned by the cleaning of antlers
provokes the same frenzy as at another time is aroused by the voice.
Dullness of perception not only in these matters, but at all times, is
fatal.
As touching the less conspicuous secondary sexual characters of Deer
more must be said presently. For the moment the antlers must retain
our attention. Time was when the Deer lacked these appendages. When
they first appeared, in the now extinct species of the Middle Miocene
period, they were no more than short prongs. Later, one of the prongs
became elongated, and developed short branches or “tines,” which, in
succeeding species, became more numerous, while at the same time, with
the gradual evolution of more and more species, these antlers assumed
new features both in the matter of size and in the character and number
of the “tines,” a development which has reached its maximum to-day. But
apart from these specific variations, which have given us such types as
those of the Roe-deer, Red-deer, Wapiti, Caribou, Moose, Fallow-deer,
Sambar, Schombergk’s deer, the strange Milou-deer, Elds-deer and
Mule-deer, each species displays a quite remarkable range of variation
in regard to its particular type of antler. Nowhere, perhaps, is this
more strikingly marked than in the case of the Caribou and Moose. No
doubt this feature is due largely to the fact that the horns are shed
annually, and that the variations are due, in part at any rate, to
temporary environmental conditions, such as food and weather. But
these apart, individual peculiarities are constant, reappearing with
more or less exactness each year.
[Illustration: Plate 6.
_Photo by Lord Delamere, from “The Living Animals of the World.”_
GROUP OF BEISA ORYX.
The lance-like horns of these animals can be used with deadly effect,
even against lions.
[Face page 60.]
In contemplating these facts one asks: What are the underlying factors
of this variability? What is the significance of the branching? What
end is attained by the annual shedding? That the antlers constitute
very effective weapons of offence there can be no doubt, and one is
inclined to regard the branching as the outcome of natural selection,
on the assumption that branched antlers would be less deadly than
lance-like weapons. It would perhaps be tempting to accept this
interpretation as all sufficient were it not for the evidence afforded
by the hollow-horned ruminants. The Oryx and the Kudu, for example, are
lance-bearers, and therefore show conclusively that stags similarly
armed might well have continued to survive in spite of the foils which
the “tines” provide. Darwin, long since, guardedly suggested that
while these weapons primarily served for offensive purposes, their
elaborate systems of branching might have been brought about by sexual
selection. That is to say, the extreme beauty of the weapons may excite
the admiration of the females as well as our own. Granting this, he
inferred they might have played an important part in elaborating the
branching by constantly displaying a preference to mate with those
males possessed of the largest and most branched antlers. But there
are many and serious objections to this suggestion, and the most
important of all is the fact that the female is allowed no choice in
the selection of her lord and master. We can, then, only regard the
antlers of deer as another instance of the survival of a “fortuitous”
but inherent variation, which survived because, whatever the defects
thereof, they proved advantageous in the struggle for existence.
Having regard to the fact that so many of the females among the
hollow-horned ruminants have acquired horns, it is somewhat remarkable
that in the Reindeer alone among the deer are these weapons normally
possessed by the female. The gradual transference to the female of
features which were originally secondary sexual characters in the
male is an occurrence which is met with in every group of animals. In
writing “The Infancy of Animals” I gave a number of instances of this
kind. But the case of the Reindeer affords a more than usually striking
illustration of this curious sequence; and this because rudiments of
antlers are to be met with among the females in several different
species of Deer to-day. They have been found in the females of both
Roe- and Red-deer, though such cases are rarely met with. As a rule
this assumption of the male secondary sexual characters by the female
occurs only in very aged animals, or as one of the sequelæ of diseased
ovaries and consequent sterility. But at least one instance is on
record of a doe Roe-deer which possessed small antlers while pregnant.
Thus, then, we gain a further insight into the process by which the
female slowly assumes the outward attributes of the male; that is
to say, the secondary sexual characters appear first in the male,
and as seasonal characters. Sooner or later they become permanently
established. By the time they have become firmly fixed in the male, and
apparently not till then, they appear in a dilute form during senility,
or in consequence of ovarian disease, in the female. Having once
started, however, they appear earlier and earlier in the life-history
of succeeding generations of females, and at last in the juvenile
stages of both sexes.
The hollow-horned ruminants, which must now be considered, afford some
very striking facts in regard to these “secondary sexual characters,”
more especially in so far as horns are concerned. In the first place
these weapons are permanent structures, taking the form of a bony core
ensheathed in horn, with which we may compare the temporary covering
of velvet in the deer: in the second, they are unbranched. The only
exception to this rule is furnished by the Prong-horned Antelope,
wherein the sheath is both annually shed, and branched. The branching,
however, is very slight, taking the form of a short forwardly directed
prong about half-way up the sheath, which is borne on a long bony
pedicle recalling that of the Muntjac. The shedding is due to the
formation of new horn material at the base of the old sheath, which
is gradually forced off by the growth of the new tissue. Structurally
the horn of this remarkable Antelope differs somewhat from that of its
relatives.
As may be seen in Plate 4, in the form of the horns the typical
hollow-horned ruminants present an exceedingly varied range, and one
often of great beauty in the matter of curvature. That they serve as
formidable weapons of offence was demonstrated during 1912, when,
according to the Annual Report of the Government Game Reserves,
published by the Pretoria Government, the game warden, Major Stevenson
Hamilton, reported of the Antelopes that “many carcases of males of
almost all species, killed in single combat with rivals, were found
during the mating season, untouched by anything except vultures.” As
a rule, however, these animals, like the Sheep and Goats, and their
larger relatives the Cattle, seem to avoid a duel to the death. One
or two instances as to the general character of these combats for
the possession of mates must suffice. Thus the late A. H. Neumann,
a hunter of experience, remarks that he once or twice saw conflicts
between the Topi (_Damaliscus jimela_), an ally of the Hartebeestes.
The two rivals would stand a little apart, affecting, apparently, to be
unaware of one another’s presence. Suddenly they would rush headlong at
one another, bringing their heads together with a clash, each, at the
same moment, falling on his knees.
Major Powell Cotton, again, once witnessed an affray between two Beisa
Oryx. Here the master bull of the herd was infuriated by the advent of
an intruder in his harem. Time after time they dashed at each other,
their foreheads meeting with a thud; then, with horns interlocked,
they wrestled fiercely; then, separating, they charged again. Yet
neither, he remarks, tried to use his lance-points, as they do when
cornered by man or beasts of prey. Nevertheless, encounters of a more
sanguinary character appear to be by no means rare, for it is no
uncommon experience of hunters to kill bulls of this species in which
one eye has been burst by a horn-thrust. Another peculiarity of these
animals is the extreme thickness of the hide of the neck and withers,
which seems to afford a shield against such spear-thrusts during
these battles. How powerful is the thrust of these weapons, and how
efficiently they can be used, is shown by the fact that lions in making
an attack on an old bull are often severely wounded, or even killed.
And there are many instances on record of cases where both the lion and
his intended victim have died together, the Antelope having been unable
to withdraw his horns from his adversary’s body. The beautiful Pala
Antelope fights furiously with rival rams, and the vanquished, as with
so many of the Antelopes, form herds by themselves, till one by one
they gather strength and skill enough to establish their right to mate.
[Illustration: Plate 7.
_Photo by courtesy of the Duchess of Bedford, Woburn Abbey._
ELAND COWS.
Among antelopes the females commonly bear horns, which may be even
longer than in the males, though less massive.
[Face page 64.]
[Illustration: Plate 8.
_Photo by courtesy of the Duchess of Bedford, Woburn Abbey._
AMERICAN BISON
The “Secondary Sexual Characters” of the male are here conspicuously
developed, and are seen in the massive fore-quarters and enormous
head.]
The Elands present some puzzling features, for both sexes bear large
horns, and they are very massive in the bulls. Yet these animals
are generally described as the most inoffensive of all the horned
ruminants. That the horns are used to any extent in conflicts between
rival males seems doubtful, inasmuch as this species is remarkable for
the development of an enormous “dewlap,” a thin pendulous fold of skin
which runs from the throat to the chest. Such a form of “ornament”—for
in this light we must regard it—would be dangerous, indeed, when much
fighting was to be done. Nevertheless, it would be contrary to all
our experience to conclude that weapons so well developed as are the
horns of the bull Eland were entirely useless. This is a matter which
decidedly calls for further investigation.
That our knowledge of that most important period of life of the larger
mammals, the period of sexual exaltation, is lamentably incomplete will
be realized by anyone who seeks enlightenment on this subject. Most
of the meagre information we possess has been collected by travellers
and sportsmen, neither of whom have the time to devote to the long and
laborious watches that a fuller history demands. Every now and then a
glimpse is afforded of this period of the life-history which brings
home in a very convincing fashion, how little is really known. It
seems certain that the fighting hitherto described is to be regarded
as but a phase of a cycle of events which takes place at this time.
Thus, for example, the old naturalist and traveller Schweinfurth tells
how he once encountered a herd of Hartebeest which were apparently
effervescing with animal spirits, for they kept running around in
couples, like horses in a circus, using a clump of trees as a pivot.
Others, in groups of three or four, stood by, interested spectators.
After a time these, in turn, took their places and ran round, two at a
time, in their own circuit, and in the same fashion. Their evolutions,
he says, were so regular as to suggest the guidance of some invisible
ring-master. These gyrations may be regarded as an erotic dance. The
Sambar, under like excitement, will stalk about with erected tail,
outstretched muzzle and everted face glands, and the Black-buck, among
the antelopes, behaves in like fashion.
It cannot be supposed that these quaint performances are peculiar to
the species in which they have been observed, but rather it may be
inferred that similar antics, besides others yet to be discovered,
are performed by all. Their purpose seems plain enough, for they must
be regarded surely as aphrodisiacs, excitants to pairing. They recall
the erotic dances of savages, or the ceremonial orgies of ancient
civilizations. Such performances, on an even more elaborate scale, are
to be met with among the birds.
So far, in describing the horned ruminants, the horns only have been
considered; but these animals display yet other secondary sexual
characters, which, while less conspicuous, are yet no less important
during this critical period of life. Some, as for instance the canine
teeth possessed by some of the deer, are decidedly puzzling. While
absent, or vestigial, in most, in a few they are greatly developed, and
this, too, in species which possess relatively large horns, as in the
Muntjac. It seems difficult to believe that the co-existence of these
very different kinds of weapons can be of vital importance to their
possessors; yet unless this be so, one or other would surely have
degenerated. It is significant that in the hornless Musk-deer these
teeth attain to a very considerable length, at their maximum as much
as three inches. That they are used by rival males, and with effect,
is shown by the fact that the hides of these animals are often found
scored by deep lines cut by these tusks. In those aberrant ruminants,
the Camels, quite formidable tusks are present both in the upper and
lower jaws, and these are used with effect whenever occasion demands,
and often when it does not.
The armoury necessary for successful love-making contains yet other
weapons, evolved to supplement physical force, and more subtle in their
effect. Such are certain skin glands which, at the rutting season,
secrete a copious flow of a creamy, or semi-fluid matter, and pungent
odour. In the deer the more important of these are found in the deep
pit, or “larmier,” which opens in front of the eye. In the Musk-deer,
however, this secretion has a most powerful odour of musk, and is
formed in a pouch, or “pod,” of about the size of a small orange, under
the skin of the abdomen. The secretion, which is formed by the male
only, is of a chocolate colour, and of about the consistence of moist
gingerbread. It has a most pungent scent, and when diluted forms the
basis of many of our most powerful and most highly-prized perfumes, on
which account, it may be mentioned, this animal has for generations
been submitted to a most unrelenting persecution. But that is another
story.
In most of the antelopes the principal scent gland is seated in a pit
in front of the eye, as in the deer. In some, as in the Gnu, it forms
instead a swollen, tumid area, oblong in shape, instead of lying in a
pit. In the Reedbuck it is placed around the bases of the horns; and
in the Rocky-Mountain Goat it forms a great bare cushion behind the
horns. All have more or less well-developed glands seated in the skin
between the toes. But, wherever placed, the secretions thereof are more
or less completely suspended save during the breeding season, when they
are poured forth abundantly. The precise rôle they play is by no means
certainly known. It seems reasonable to suppose that, in the first
place, the odour they disperse enables the males to announce their
whereabouts to the females seeking mates, should they fail to hear
their bellowing. But the antelopes, for the most part, unlike deer, do
not, the year round, lose touch with one another; so that it must be
concluded that these odours serve as excitants to the act of pairing,
and we know that the sense of smell plays a very important part at
this time, which, so far as these animals are concerned, is the only
period which comes more or less exactly within the meaning of the term
“courtship.”
That scent among the antelopes holds a really important place is shown
by the fact that the bull of the common Eland intensifies his natural
odours by micturating upon the mass of long hair which grows upon the
forehead. To do this the head is bent down and turned tailwards, in
order that the tuft should receive its due urinary spray! And goats in
captivity exhibit the same curious habit. In them, indeed, it is often
pushed to such an excess that blindness results, so that the animal has
to be slaughtered.
While in many cases these odours are imperceptible to human nostrils,
in others this is far from being the case. Among the ruminants the goat
is particularly odorous. So also are the giraffe and the water-buck,
both of which may be detected by their smell at considerable
distances. And these emanations are most noticeable in the males and at
the breeding season. The bull elephant, both in the Indian and African
species, during the breeding season produces a copious flow of aromatic
matter from a gland which opens above the eye in the form of a tubular
aperture large enough to admit a pencil. This aperture in the African
elephant is remarkable for the fact that it is invariably found to be
“plugged” with numerous spines of the acacia, which have from time to
time found their way in as the animal was forcing its way through the
dense undergrowth. This extraordinary fact was first noticed by Mr. F.
C. Selous, and has since been confirmed by Dr. Einar Lonnberg.
It is probable that the “bloody sweat,” which at times covers the hide
of the Hippopotamus just after leaving the water, is associated with
the period of rut. This mysterious exudation is accompanied by small
crystals; but though red in colour, it contains no blood. So far no
reasonable explanation for this remarkable phenomenon has ever been
given, but probably it will be found to be associated with the sexual
activities and is possibly odoriferous. A precisely similar exudation
occurs in the neck of the male of the Red Kangaroo.
That these secretions play an important and perhaps variable part
in the selection of mates seems demonstrated in the case of an
incident related to me by my friend Mr. John Cooke, who some time
ago was watching a flock of some three hundred sheep while it was
being driven by the shepherd and his dogs into a field. As soon as
they were securely shut in, and the shepherd had gone, three rams
who were included in the flock at once began a three-cornered fight.
One, presumably the youngest, was soon vanquished. The other two
soon settled their differences, and the clashing of horns was at once
followed by a very different performance. The master ram began to run
in and out among the ewes, sniffing at each, and driving out those
whose odour most pleased him. Having at last satisfied himself with a
harem of about one hundred, the second ram was allowed to make a like
choice, and behaved in a like manner, leaving the remainder to the
ram which was first vanquished. May we take it that the strongest and
oldest rams selected the youngest ewes, and the oldest were left to the
youngest, and first conquered ram? By some such rough and ready method
of selection Nature may contrive that the immature male shall do as
little harm to the race as possible by mating with the oldest, and in
many cases barren females.
Our survey of the “hoofed” animals has so far been confined to the
ruminants. Space must now be found for a brief review of what obtains
under like circumstances in the case of the great pachyderms—the
Elephant, Rhinoceros and Hippopotamus; the Pig and the Camel.
[Illustration: Plate 9.
_Photo by Lord Delamere, from “The Living Animals of the World”_
ELEPHANTS.
The sexes differ but little in general appearance: and chiefly in the
superior size of the male and his more massive tusks.
[Face page 70.]
As to actual “courtship” among these animals practically nothing
is known; but the varied and formidable weapons which they possess
are enough to show that the secondary sexual characters play a very
important part in the preliminary capture of mates. That they may
also be used for the more prosaic purpose of securing food is nothing
to the point. In the Elephant, for example, the tusks are sometimes
of enormous size and weight, specimens of eleven feet in length and
weighing as much as two hundred and fifty pounds are on record. They
are used for cutting through the bark of machabel trees, which is then
seized by the trunk and torn off, for elephants are extremely fond of
this bark; and they are also turned to account in breaking up roots
which have been exposed by digging with the fore-feet. But this is
certainly not the main purpose of such weapons. On the contrary, their
use is primarily as weapons of offence between rival bulls. As one
would expect, they never attain to a very large size in the female, but
that they are large enough to serve her at need is shown by the fact
that a portion of a tusk, evidently of a cow-elephant, was once found
embedded in the jaw of a bull. There can be little doubt but that this
was broken off in an endeavour to repel the advances of a too amorous
male, for, as with all animals, pairing is impossible without the
consent of the female, and this is never accorded until she is desirous
that it should take place. As a preliminary to this, an amorous
dalliance is perhaps the invariable rule among animals, and this takes
many and often strange forms. The Elephant affords a case in point. For
the late A. H. Neumann once came upon a pair which were evidently, as
he says, “love-making.” Creeping upon them noiselessly, he found the
male fondling his mate with his trunk, and then, standing side by side,
they crossed their trunks, and put the tips thereof into each other’s
mouths, the elephantine form of kissing. Deer, cattle and horses, cats
and dogs, constantly lick one another under like circumstances.
Superficial secondary sexual characters are wanting both in the
Hippopotamus and the Camel. Both, however, possess a formidable
armature of teeth which are capable of inflicting very severe wounds.
In the Hippopotamus the canines are of enormous size, and their
punishing power is further strengthened by the fact that they work
in opposition to a pair of similar teeth in the lower jaw; they cut
like a pair of shears, the upper closing upon the lower pair with
the precision of scissors-blades. In addition, the lower jaw develops
two long, blunt-pointed, ivory spikes, which are scarcely less to
be dreaded. With these weapons the bulls fight furiously, and it is
no uncommon thing to find vanquished males frightfully mauled, the
hide being lacerated from head to tail. Protection, in a measure, is
afforded by its enormous thickness, but the great folds and pleats of
skin seen in the Rhinoceros are never developed. The females, however,
are similarly armed, and the teeth are nearly as large as in the males,
which is a rather unusual occurrence.
The Swine, which are near relations of the Hippopotamus, in like manner
develop huge pointed canines, and these reach their maximum in the
great Wart-hogs of Africa. But in the swine the mechanism differs,
for although the canines are closely opposed, the shaft of the upper
teeth curves upwards, and the lower teeth are much smaller than the
upper. In fighting, these animals do not bite, like the Hippopotamus,
but use the upper canines to rip up their antagonist with a sudden,
swift upward and sideways movement of the head. How dangerous is the
wound thus inflicted those who have hunted the wild-boar know well. A
curious exaggeration of this arrangement of the teeth is seen in the
Babiroussa. Herein the upper canines grow directly upwards, actually
piercing the upper lip as in the case of the downwardly growing tusks
of the elephant. That these teeth, however, are of any service in
fighting is doubtful, for the upper tooth curves upwards and backwards
in a semicircle so that the points are harmless. The tusks of the lower
jaw, however, are extremely long and pointed, though their wounding
power is limited by reason of the upper teeth. This may account for
the fact that the head, the part mostly attacked by enraged
boars, presents no sort of armature designed for defence; while in the
Wart-hog, on the other hand, great solid bucklers of hide stand out
on either side of the head below the eyes, giving the animal a most
repulsive appearance, but affording him a very present help in time
of trouble. In the wild-boar, where the tusks are shorter, no such
protective armature is needed.
[Illustration: Plate 10.
HEAD OF MALE WART-HOG.
In the “Swine” family the canine teeth are always greatly developed,
but they attain to their maximum, relatively, in the Wart-hog.]
[Illustration:
_Photos by Scholastic Photo Co., from “The Living Animals of the World.”_
MALE AND FEMALE BABIRUSA.
A characteristic of this pig is the peculiar development of the tusks
in the male, the upper pair of which grow through the lips and curve
upwards.
[Face page 72.]
[Illustration: Plate 11.
_Photo by Lord Delamere, from “The Living Animals of the World.”_
SOMALI ZEBRAS.
The Zebras, unlike their cloven-hoofed relations, have no weapons, save
for inter-tribal conflicts. Yet they have been as successful in holding
their own against lions and other predatory animals as species provided
with horns.
[Face page 72]
[Illustration: Plate 12.
_Photo copyright by A. H. Bishop._
GIRAFFE.
The horns of this animal can prove formidable weapons of offence on
occasion, though they are useless against predatory animals.]
While the ungulates, or hoofed animals, are peculiar in the development
of horns as weapons of offence, they are by no means singular in the
use of teeth for this purpose. In some cases, as in the Muntjac, both
forms of armature are present. The only other instances where teeth
in this group of animals are used for offensive purposes are those
furnished by the Camel and the Horse. But here they do not exhibit
that excessive size which is met with in the Elephant, and some of
the Swine. In both the Camel and the Horse it is the canine which is
used, and both jaws are similarly armed. Since the camel has no upper
incisors, the part played by the teeth is beyond dispute; but it has
been contended that the horse uses his incisor or “front-teeth” alone
when fighting. But this is not so; the canines can, and do, inflict
ugly wounds, as is shown by the necks of zebras.
A further method of defence among the larger Ungulates, at any rate,
is resorted to when hard pressed: and this is the use of the hoof in
kicking. Giraffes kick both after the usual fashion and in striking
downwards with the fore-foot. And an interesting demonstration of this
has been furnished by Mr. F. C. Selous in his delightful “African
Nature Notes.” He relates that on one occasion he came across a calf
only a day or two old, with its back broken. From scratches on the
calf, and the footprints on the ground in its vicinity, he was at once
enabled to gather the cause of its terrible plight. In a word, it had
been attacked by two leopards, and the mother, in an endeavour to beat
off the assailants with a blow of her fore-foot had accidentally struck
her offspring. Horses, Cattle, Antelopes, Camels and Elephants can all
kick with precision and effect. So far as the evidence goes, however,
this is a method of defence used against beasts of prey, and is rarely,
if ever, employed in conflicts between rival males. Females persecuted
by the undesired attentions of amorous males, however, do, as we know
from the case of domesticated animals, use this device to defend
themselves.
It is not difficult to account for the origin of such secondary sexual
characters as manes, beards, tusks, and brightly-coloured areas of
skin, though whether our interpretations are really correct is another
matter. But no attempt to explain the origin of horns has yet achieved
a like degree of persuasiveness. These weapons appear only in the
Ungulates, a group which has, in past times, given birth to some very
extraordinary types of head armature of this kind. These must be
excluded from the present discussion; suffice it to say that, as usual,
they were the adjuncts of the males. According to current theories it
is supposed that these weapons arose as the result of the action of
sexual selection. It is assumed that the hornless ancestors of now
horned ruminants fought for their mates by “butting” with the forehead.
Naturally, other things being equal, the thickest skulled combatants
obtained the mastery. Any tendency to develop frontal “bosses” of bone
would further enhance the chances of success, and would, indeed, soon
become necessary for survival. And from such “bosses” the passage to
horns and antlers forms an easy transition. Just such incipient horns
or “bosses” actually make their appearance in the domesticated horse:
but these animals never butt at one another. If, however, we regard
horn-production as an inherent diathesis of the ungulate somatoplasm,
we have an intelligible basis for the explanation of horn development.
The formidable horns of the Rhinoceros are of a totally different
character, being solid structures formed by hairlike agglomerations,
firmly fixed upon a roughened area of the nasal region. These weapons
play a very important part in settling disputes between rival males,
but on other occasions demanding offensive tactics the Indian
Rhinoceros at any rate seems to depend rather on his power of wounding
by means of the chisel-shaped lower incisors. These, by means of a
swift lateral movement of the head can be made to inflict most terrible
gashes, as those who hunt with elephants well know. It is quite
possible, however, that the teeth are also thus used during struggles
for supremacy. And this may perhaps account for the enormous bucklers
of skin developed by the Indian Rhinoceros, but only indicated in the
case of the African species.
All the larger Ungulates, and many of the smaller species, are
polygamous. The Rhinoceros, and all of the swine-group save the
Hippopotamus, among the larger species are exceptions to the rule. The
preponderance of females which this implies is generally supposed to
be due to the losses sustained among the males by fighting during the
struggle for mates. The case of horses, however, seems to militate
against this view, for though they undoubtedly fight furiously, no
evidence is forthcoming to show that such conflicts terminate fatally.
Were it possible to secure the necessary data it would probably be
found that polygamy, and polyandry, are determined solely by the
numerical proportions of the sexes: the excess of males or females
being due neither to “Natural” nor “Sexual” Selection, but to inherent
peculiarities of the germ-plasm tending to produce an excess of males,
or females, as the case may be.
Finally, all the evidence goes to show that it is a mistake to
suppose that polygamy is due to the excessive sexual avidness of the
males, which impels them to first essay the overthrow of all possible
rivals, and then to appropriate every female within their sphere of
influence, holding them by force. On the contrary, this plurality of
mates is thrust upon them. And this because the females, impelled by
“mate-hunger,” attach themselves to the nearest male within call: the
size of the harem depending on the number of available males. The
battles which are fought between rival males are no more sanguinary
than in the case of monogamous species. This contention is well
illustrated by the African Wydah-birds (_Vidua_), which are markedly
polygamous, though they have no special weapons of offence. In
districts where males are numerous the harem will not exceed eight, or
ten, females; where males are scarce this number may be increased to
fifty. In like manner the varying number of hinds accompanying a stag
are to be regarded, not as an index of his prowess, but of the scarcity
or abundance of males in the neighbourhood.