CHAPTER XVII.
Secondary Sexual Characters of Mammals.
The law of battle—Special weapons, confined to the males—Cause of absence of weapons in the female—Weapons common to both sexes, yet primarily acquired by the male—Other uses of such weapons—Their high importance—Greater size of the male—Means of defence—On the preference shewn by either sex in the pairing of quadrupeds.
With mammals the male appears to win the female much more through the law of battle than through the display of his charms. The most timid animals, not provided with any special weapons for fighting, engage in desperate conflicts during the season of love. Two male hares have been seen to fight together until one was killed; male moles often fight, and sometimes with fatal results; male squirrels “engage in frequent contests, and often wound each other severely;” as do male beavers, so that “hardly a skin is without scars.”[288] I observed the same fact with the hides of the guanacoes in Patagonia; and on one occasion several were so absorbed in fighting that they fearlessly rushed close by me. Livingstone speaks of the males of the many animals in Southern Africa as almost invariably shewing the scars received in former contests.
The law of battle prevails with aquatic as with terrestrial mammals. It is notorious how desperately male seals fight, both with their teeth and claws, during the breeding-season; and their hides are likewise often covered with scars. Male sperm-whales are very jealous at this season; and in their battles “they often lock their jaws together, and turn on their sides and twist about;” so that it is believed by some naturalists that the frequently deformed state of their lower jaws is caused by these struggles.[289]
All male animals which are furnished with special weapons for fighting, are well known to engage in fierce battles. The courage and the desperate conflicts of stags have often been described; their skeletons have been found in various parts of the world, with the horns inextricably locked together, shewing how miserably the victor and vanquished had perished.[290] No animal in the world is so dangerous as an elephant in must. Lord Tankerville has given me a graphic description of the battles between the wild bulls in Chillingham Park, the descendants, degenerated in size but not in courage, of the gigantic Bos primigenius. In 1861 several contended for mastery; and it was observed that two of the younger bulls attacked in concert the old leader of the herd, overthrew and disabled him, so that he was believed by the keepers to be lying mortally wounded in a neighbouring wood. But a few days afterwards one of the young bulls singly approached the wood; and then the “monarch of the chase,” who had been lashing himself up for vengeance, came out and, in a short time killed his antagonist. He then quietly joined the herd, and long held undisputed sway. Admiral Sir B. J. Sulivan informs me that when he resided in the Falkland Islands he imported a young English stallion, which, with eight mares, frequented the hills near Port William. On these hills there were two wild stallions, each with a small troop of mares; “and it is certain that these stallions would never have approached each other without fighting. Both had tried singly to fight the English horse and drive away his mares, but had failed. One day they came in together and attacked him. This was seen by the capitan who had charge of the horses, and who, on riding to the spot, found one of the two stallions engaged with the English horse, whilst the other was driving away the mares, and had already separated four from the rest. The capitan settled the matter by driving the whole party into the corral, for the wild stallions would not leave the mares.”
Male animals already provided with efficient cutting or tearing teeth for the ordinary purposes of life, as in the carnivora, insectivora, and rodents, are seldom furnished with weapons especially adapted for fighting with their rivals. The case is very different with the males of many other animals. We see this in the horns of stags and of certain kinds of antelopes in which the females are hornless. With many animals the canine teeth in the upper or lower jaw, or in both, are much larger in the males than in the females; or are absent in the latter, with the exception sometimes of a hidden rudiment. Certain antelopes, the musk-deer, camel, horse, boar, various apes, seals, and the walrus, offer instances of these several cases. In the females of the walrus the tusks are sometimes quite absent.[291] In the male elephant of India and in the male dugong[292] the upper incisors form offensive weapons. In the male narwhal one alone of the upper teeth is developed into the well-known, spirally-twisted, so called horn, which is sometimes from nine to ten feet in length. It is believed that the males use these horns for fighting together; for “an unbroken one can rarely be got, and occasionally one may be found with the point of another jammed into the broken place.”[293] The tooth on the opposite side of the head in the male consists of a rudiment about ten inches in length, which is embedded in the jaw. It is not, however, very uncommon to find double-horned male narwhals in which both teeth are well developed. In the females both teeth are rudimentary. The male cachalot has a larger head than that of the female, and it no doubt aids these animals in their aquatic battles. Lastly, the adult male ornithorhynchus is provided with a remarkable apparatus, namely a spur on the foreleg, closely resembling the poison-fang of a venomous snake; its use is not known, but we may suspect that it serves as a weapon of offence.[294] It is represented by a mere rudiment in the female.
When the males are provided with weapons which the females do not possess, there can hardly be a doubt that they are used for fighting with other males, and that they have been acquired through sexual selection.
It is not probable, at least in most cases, that the females have actually been saved from acquiring such weapons, owing to their being useless and superfluous, or in some way injurious. On the contrary, as they are often used by the males of many animals for various purposes, more especially as a defence against their enemies, it is a surprising fact that they are so poorly developed or quite absent in the females. No doubt with female deer the development during each recurrent season of great branching horns, and with female elephants the development of immense tusks, would have been a great waste of vital power, on the admission that they were of no use to the females. Consequently variations in the size of these organs, leading to their suppression, would have come under the control of natural selection, and if limited in their transmission to the female offspring would not have interfered with their development through sexual selection in the males. But how on this view can we explain the presence of horns in the females of certain antelopes, and of tusks in the females of many animals, which are only of slightly less size than in the males? The explanation in almost all cases must, I believe, be sought in the laws of transmission.
As the reindeer is the single species in the whole family of Deer in which the female is furnished with horns, though somewhat smaller, thinner, and less branched than in the male, it might naturally be thought that they must be of some special use to her. There is, however, some evidence opposed to this view. The female retains her horns from the time when they are fully developed, namely in September, throughout the winter, until May, when she brings forth her young; whilst the male casts his horns much earlier, towards the end of November. As both sexes have the same requirements and follow the same habits of life, and as the male sheds his horns during the winter, it is very improbable that they can be of any special service to the female at this season, which includes the larger proportion of the time during which she bears horns. Nor is it probable that she can have inherited horns from some ancient progenitor of the whole family of deer, for, from the fact of the males alone of so many species in all quarters of the globe possessing horns, we may conclude that this was the primordial character of the group. Hence it appears that horns must have been transferred from the male to the female at a period subsequent to the divergence of the various species from a common stock; but that this was not effected for the sake of giving her any special advantage.[295]
We know that the horns are developed at a most unusually early age in the reindeer; but what the cause of this may have been is not known. The effect, however, has apparently been the transference of the horns to both sexes. It is intelligible on the hypothesis of pangenesis, that a very slight change in the constitution of the male, either in the tissues of the forehead or in the gemmules of the horns, might lead to their early development; and as the young of both sexes have nearly the same constitution before the period of reproduction, the horns, if developed at an early age in the male, would tend to be developed equally in both sexes. In support of this view, we should bear in mind that the horns are always transmitted through the female, and that she has a latent capacity for their development, as we see in old or diseased females.[296] Moreover the females of some other species of deer either normally or occasionally exhibit rudiments of horns; thus the female of Cervulus moschatus has “bristly tufts, ending in a knob, instead of a horn;” and “in most specimens of the female Wapiti (Cervus Canadensis) there is a sharp bony protuberance in the place of the horn.”[297] From these several considerations we may conclude that the possession of fairly well-developed horns by the female reindeer, is due to the males having first acquired them as weapons for fighting with other males; and secondarily to their development from some unknown cause at an unusually early age in the males, and their consequent transmission to both sexes.
Turning to the sheath-horned ruminants: with antelopes a graduated series can be formed, beginning with the species, the females of which are completely destitute of horns—passing to those which have horns so small as to be almost rudimentary, as in Antilocapra Americana—to those which have fairly well-developed horns, but manifestly smaller and thinner than in the male, and sometimes of a different shape,[298] and ending with those in which both sexes have horns of equal size. As with the reindeer, so with antelopes there exists a relation between the period of the development of the horns and their transmission to one or both sexes; it is therefore probable that their presence or absence in the females of some species, and their more or less perfect condition in the females of other species, depend, not on their being of some special use, but simply on the form of inheritance which has prevailed. It accords with this view that even in the same restricted genus both sexes of some species, and the males alone of other species, are thus provided. It is a remarkable fact that, although the females of Antilope bezoartica are normally destitute of horns, Mr. Blyth has seen no less than three females thus furnished; and there was no reason to suppose that they were old or diseased. The males of this species have long straight spirated horns, nearly parallel to each other, and directed backwards. Those of the female, when present, are very different in shape, for they are not spirated, and spreading widely bend round, so that their points are directed forwards. It is a still more remarkable fact that in the castrated male, as Mr. Blyth informs me, the horns are of the same peculiar shape as in the female, but longer and thicker. In all cases the differences between the horns of the males and females, and of castrated and entire males, probably depend on various causes,—on the more or less complete transference of male characters to the females,—on the former state of the progenitors of the species,—and partly perhaps on the horns being differently nourished, in nearly the same manner as the spurs of the domestic cock, when inserted into the comb or other parts of the body, assume various abnormal forms from being differently nourished.
In all the wild species of goats and sheep the horns are larger in the male than in the female, and are sometimes quite absent in the latter.[299] In several domestic breeds of the sheep and goat, the males alone are furnished with horns; and it is a significant fact, that in one such breed of sheep on the Guinea coast, the horns are not developed, as Mr. Winwood Reade informs me, in the castrated male; so that they are affected in this respect like the horns of stags. In some breeds, as in that of N. Wales, in which both sexes are properly horned, the ewes are very liable to be hornless. In these same sheep, as I have been informed by a trustworthy witness who purposely inspected a flock during the lambing-season, the horns at birth are generally more fully developed in the male than in the female. With the adult musk-ox (Ovibos moschatus) the horns of the male are larger than those of the female, and in the latter the bases do not touch.[300] In regard to ordinary cattle Mr. Blyth remarks: “In most of the wild bovine animals the horns are both longer and thicker in the bull than in the cow, and in the cow-banteng (Bos sondaicus) the horns are remarkably small, and inclined much backwards. In the domestic races of cattle, both of the humped and humpless types, the horns are short and thick in the bull, longer and more slender in the cow and ox; and in the Indian buffalo, they are shorter and thicker in the bull, longer and more slender in the cow. In the wild gaour (B. gaurus) the horns are mostly both longer and thicker in the bull than in the cow.”[301] Hence with most sheath-horned ruminants the horns of the male are either longer or stronger than those of the female. With the Rhinoceros simus, as I may here add, the horns of the female are generally longer but less powerful than in the male; and in some other species of rhinoceros they are said to be shorter in the female.[302] From these various facts we may conclude that horns of all kinds, even when they are equally developed in both sexes, were primarily acquired by the males in order to conquer other males, and have been transferred more or less completely to the female, in relation to the force of the equal form of inheritance.
The tusks of the elephant, in the different species or races, differ according to sex, in nearly the same manner as the horns of ruminants. In India and Malacca the males alone are provided with well-developed tusks. The elephant of Ceylon is considered by most naturalists as a distinct race, but by some as a distinct species, and here “not one in a hundred is found with tusks, the few that possess them being exclusively males.”[303] The African elephant is undoubtedly distinct, and the female has large, well-developed tusks, though not so large as those of the male. These differences in the tusks of the several races and species of elephants—the great variability of the horns of deer, as notably in the wild reindeer—the occasional presence of horns in the female Antilope bezoartica—the presence of two tusks in some few male narwhals—the complete absence of tusks in some female walruses;—are all instances of the extreme variability of secondary sexual characters, and of their extreme liability to differ in closely-allied forms.
Although tusks and horns appear in all cases to have been primarily developed as sexual weapons, they often serve for other purposes. The elephant uses his tusks in attacking the tiger; according to Bruce, he scores the trunks of trees until they can be easily thrown down, and he likewise thus extracts the farinaceous cores of palms; in Africa he often uses one tusk, this being always the same, to probe the ground and thus to ascertain whether it will bear his weight. The common bull defends the herd with his horns; and the elk in Sweden has been known, according to Lloyd, to strike a wolf dead with a single blow of his great horns. Many similar facts could be given. One of the most curious secondary uses to which the horns of any animal are occasionally put, is that observed by Captain Hutton[304] with the wild goat (Capra ægagrus) of the Himalayas, and as it is said with the ibex, namely, that when the male accidentally falls from a height he bends inwards his head, and, by alighting on his massive horns, breaks the shock. The female cannot thus use her horns, which are smaller, but from her more quiet disposition she does not so much need this strange kind of shield.
Each male animal uses his weapons in his own peculiar fashion. The common ram makes a charge and butts with such force with the bases of his horns, that I have seen a powerful man knocked over as easily as a child. Goats and certain species of sheep, for instance the Ovis cycloceros of Afghanistan,[305] rear on their hind legs, and then not only butt, but “make a cut down and a jerk up, with the ribbed front of their scimitar-shaped horn, as with a sabre. When the O. cycloceros attacked a large domestic ram, who was a noted bruiser, he conquered him by the sheer novelty of his mode of fighting, always closing at once with his adversary, and catching him across the face and nose with a sharp drawing jerk of his head, and then bounding out of the way before the blow could be returned.” In Pembrokeshire a male goat, the master of a flock which during several generations had run wild, was known to have killed several other males in single combat; this goat possessed enormous horns, measuring 39 inches in a straight line from tip to tip. The common bull, as every one knows, gores and tosses his opponent; but the Italian buffalo is said never to use his horns, he gives a tremendous blow with his convex forehead, and then tramples on his fallen enemy with his knees—an instinct which the common bull does not possess.[306] Hence a dog who pins a buffalo by the nose is immediately crushed. We must, however, remember that the Italian buffalo has long been domesticated, and it is by no means certain that the wild parent-form had similarly shaped horns. Mr. Bartlett informs me that when a female Cape buffalo (Bubalus caffer) was turned into an enclosure with a bull of the same species, she attacked him, and he in return pushed her about with great violence. But it was manifest to Mr. Bartlett that had not the bull shewn dignified forbearance, he could easily have killed her by a single lateral thrust with his immense horns. The giraffe uses his short hair-covered horns, which are rather longer in the male than in the female, in a curious manner; for with his long neck he swings his head to either side, almost upside down, with such force, that I have seen a hard plank deeply indented by a single blow.
Fig. 61. Oryx leucoryx, male (from the Knowsley Menagerie).
With antelopes it is sometimes difficult to imagine how they can possibly use their curiously-shaped horns; thus the spring-boc (Ant. euchore) has rather short upright horns, with the sharp points bent inwards almost at a right angle, so as to face each other; Mr. Bartlett does not know how they are used, but suggests that they would inflict a fearful wound down each side of the face of an antagonist. The slightly-curved horns of the Oryx leucoryx (fig. [61]) are directed backwards, and are of such length that their points reach beyond the middle of the back, over which they stand in an almost parallel line. Thus they seem singularly ill-fitted for fighting; but Mr. Bartlett informs me that when two of these animals prepare for battle, they kneel down, with their heads between their front legs, and in this attitude the horns stand nearly parallel and close to the ground, with the points directed forwards and a little upwards. The combatants then gradually approach each other and endeavour to get the upturned points under each other’s bodies; if one succeeds in doing this, he suddenly springs up, throwing up his head at the same time, and can thus wound or perhaps even transfix his antagonist. Both animals always kneel down so as to guard as far as possible against this manœuvre. It has been recorded that one of these antelopes has used his horns with effect even against a lion; yet from being forced to place his head between the forelegs in order to bring the points of the horns forward, he would generally be under a great disadvantage when attacked by any other animal. It is, therefore, not probable that the horns have been modified into their present great length and peculiar position, as a protection against beasts of prey. We can, however, see that as soon as some ancient male progenitor of the Oryx acquired moderately long horns, directed a little backwards, he would be compelled in his battles with rival males to bend his head somewhat inwards or downwards, as is now done by certain stags; and it is not improbable that he might have acquired the habit of at first occasionally and afterwards of regularly kneeling down. In this case it is almost certain that the males which possessed the longest horns would have had a great advantage over others with shorter horns; and then the horns would gradually have been rendered longer and longer, through sexual selection, until they acquired their present extraordinary length and position.
With stags of many kinds the branching of the horns offers a curious case of difficulty; for certainly a single straight point would inflict a much more serious wound than several diverging points. In Sir Philip Egerton’s museum there is a horn of the red-deer (Cervus elaphus) thirty inches in length, with “not fewer than fifteen snags or branches;” and at Moritzburg there is still preserved a pair of antlers of a red-deer, shot in 1699 by Frederick I., each of which bears the astonishing number of thirty-three branches. Richardson figures a pair of antlers of the wild reindeer with twenty-nine points.[307] From the manner in which the horns are branched, and more especially from deer being known occasionally to fight together by kicking with their fore-feet,[308] M. Bailly actually came to the conclusion that their horns were more injurious than useful to them! But this author overlooks the pitched battles between rival males. As I felt much perplexed about the use or advantage of the branches, I applied to Mr. McNeill of Colinsay, who has long and carefully observed the habits of red-deer, and he informs me that he has never seen some of the branches brought into action, but that the brow-antlers, from inclining downwards, are a great protection to the forehead, and their points are likewise used in attack. Sir Philip Egerton also informs me in regard both to red-deer and fallow-deer, that when they fight they suddenly dash together, and getting their horns fixed against each other’s bodies a desperate struggle ensues. When one is at last forced to yield and turn round, the victor endeavours to plunge his brow-antlers into his defeated foe. It thus appears that the upper branches are used chiefly or exclusively for pushing and fencing. Nevertheless with some species the upper branches are used as weapons of offence; when a man was attacked by a Wapiti deer (Cervus Canadensis) in Judge Caton’s park in Ottawa, and several men tried to rescue him, the stag “never raised his head from the ground; in fact he kept his face almost flat on the ground, with his nose nearly between his fore-feet, except when he rolled his head to one side to take a new observation preparatory to a plunge.” In this position the terminal points of the horns were directed against his adversaries. “In rolling his head he necessarily raised it somewhat, because his antlers were so long that he could not roll his head without raising them on one side, while on the other side they touched the ground.” The stag by this procedure gradually drove the party of rescuers backwards, to a distance of 150 or 200 feet; and the attacked man was killed.[309]
Although the horns of stags are efficient weapons, there can, I think, be no doubt that a single point would have been much more dangerous than a branched antler; and Judge Caton, who has had large experience with deer, fully concurs in this conclusion. Nor do the branching horns, though highly important as a means of defence against rival stags, appear perfectly well adapted for this purpose, as they are liable to become interlocked. The suspicion has therefore crossed my mind that they may serve partly as ornaments. That the branched antlers of stags, as well as the elegant lyrated horns of certain antelopes, with their graceful double curvature, (fig. [62]), are ornamental in our eyes, no one will dispute. If, then, the horns, like the splendid accoutrements of the knights of old, add to the noble appearance of stags and antelopes, they may have been partly modified for this purpose, though mainly for actual service in battle; but I have no evidence in favour of this belief.
Fig. 62. Strepsiceros Kudu (from Andrew Smith’s ‘Zoology of South Africa’).
An interesting case has lately been published, from which it appears that the horns of a deer in one district in the United States are now being modified through sexual and natural selection. A writer in an excellent American Journal[310] says, that he has hunted for the last twenty-one years in the Adirondacks, where the Cervus Virginianus abounds. About fourteen years ago he first heard of spike-horn bucks. These became from year to year more common; about five years ago he shot one, and subsequently another, and now they are frequently killed. “The spike-horn differs greatly from the common antler of the C. Virginianus. It consists of a single spike, more slender than the antler, and scarcely half so long, projecting forward from the brow, and terminating in a very sharp point. It gives a considerable advantage to its possessor over the common buck. Besides enabling him to run more swiftly through the thick woods and underbrush (every hunter knows that does and yearling bucks run much more rapidly than the large bucks when armed with their cumbrous antlers), the spike-horn is a more effective weapon than the common antler. With this advantage the spike-horn bucks are gaining upon the common bucks, and may, in time, entirely supersede them in the Adirondacks. Undoubtedly the first spike-horn buck was merely an accidental freak of nature. But his spike-horns gave him an advantage, and enabled him to propagate his peculiarity. His descendants, having a like advantage, have propagated the peculiarity in a constantly increasing ratio, till they are slowly crowding the antlered deer from the region they inhabit.”
Male quadrupeds which are furnished with tusks use them in various ways, as in the case of horns. The boar strikes laterally and upwards; the musk-deer with serious effect downwards.[311] The walrus, though having so short a neck and so unwieldy a body, “can strike either upwards, or downwards, or sideways, with equal dexterity.”[312] The Indian elephant fights, as I was informed by the late Dr. Falconer, in a different manner according to the position and curvature of his tusks. When they are directed forwards and upwards he is able to fling a tiger to a great distance—it is said to even thirty feet; when they are short and turned downwards he endeavours suddenly to pin the tiger to the ground, and in consequence is dangerous to the rider, who is liable to be jerked off the hoodah.[313]
Very few male quadrupeds possess weapons of two distinct kinds specially adapted for fighting with rival males. The male muntjac-deer (Cervulus), however, offers an exception, as he is provided with horns and exserted canine teeth. But one form of weapon, has often been replaced in the course of ages by another form, as we may infer from what follows. With ruminants the development of horns generally stands in an inverse relation with that of even moderately well-developed canine teeth. Thus camels, guanacoes, chevrotains and musk-deer, are hornless, and they have efficient canines; these teeth being “always of smaller size in the females than in the males.” The Camelidæ have in their upper jaws, in addition to their true canines, a pair of canine-shaped incisors.[314] Male deer and antelopes, on the other hand, possess horns, and they rarely have canine teeth; and these when present are always of small size, so that it is doubtful whether they are of any service in their battles. With Antilope montana they exist only as rudiments in the young male, disappearing as he grows old; and they are absent in the female at all ages; but the females of certain other antelopes and deer have been known occasionally to exhibit rudiments of these teeth.[315] Stallions have small canine teeth, which are either quite absent or rudimentary in the mare; but they do not appear to be used in fighting, for stallions bite with their incisors, and do not open their mouths widely like camels and guanacoes. Whenever the adult male possesses canines now in an inefficient state, whilst the female has either none or mere rudiments, we may conclude that the early male progenitor of the species was provided with efficient canines, which had been partially transferred to the females. The reduction of these teeth in the males seems to have followed from some change in their manner of fighting, often caused (but not in the case of the horse) by the development of new weapons.
Tusks and horns are manifestly of high importance to their possessors, for their development consumes much organised matter. A single tusk of the Asiatic elephant,—one of the extinct woolly species,—and of the African elephant, have been known to weigh respectively 150, 160, and 180 pounds; and even greater weights have been assigned by some authors.[316] With deer, in which the horns are periodically renewed, the drain on the constitution must be greater; the horns, for instance, of the moose weigh from fifty to sixty pounds, and those of the extinct Irish elk from sixty to seventy pounds,—the skull of the latter weighing on an average only five and a quarter pounds. With sheep, although the horns are not periodically renewed, yet their development, in the opinion of many agriculturists, entails a sensible loss to the breeder. Stags, moreover, in escaping from beasts of prey are loaded with an additional weight for the race, and are greatly retarded in passing through a woody country. The moose, for instance, with horns extending five and a half feet from tip to tip, although so skilful in their use that he will not touch or break a dead twig when walking quietly, cannot act so dexterously whilst rushing away from a pack of wolves. “During his progress he holds his nose up, so as to lay the horns horizontally back; and in this attitude cannot see the ground distinctly.”[317] The tips of the horns of the great Irish elk were actually eight feet apart! Whilst the horns are covered with velvet, which lasts with the red-deer for about twelve weeks, they are extremely sensitive to a blow; so that in Germany the stags at this time change their habits to a certain extent, and avoid dense forests, frequenting young woods and low thickets.[318] These facts remind us, that male birds have acquired ornamental plumes at the cost of retarded flight, and other ornaments at the cost of some loss of power in their battles with rival males.
With quadrupeds, when, as is often the case, the sexes differ in size, the males are, I believe, always larger and stronger. This holds good in a marked manner, as I am informed by Mr. Gould, with the marsupials of Australia, the males of which appear to continue growing until an unusually late age. But the most extraordinary case is that of one of the seals (Callorhinus ursinus), a full-grown female weighing less than one-sixth of a full-grown male.[319] The greater strength of the male is invariably displayed, as Hunter long ago remarked,[320] in those parts of the body which are brought into action in fighting with rival males,—for instance, in the massive neck of the bull. Male quadrupeds are also more courageous and pugnacious than the females. There can be little doubt that these characters have been gained, partly through sexual selection, owing to a long series of victories by the stronger and more courageous males over the weaker, and partly through the inherited effects of use. It is probable that the successive variations in strength, size, and courage, whether due to so-called spontaneous variability or to the effects of use, by the accumulation of which male quadrupeds have acquired these characteristic qualities, occurred rather late in life, and were consequently to a large extent limited in their transmission to the same sex.
Under this point of view I was anxious to obtain information in regard to the Scotch deerhound, the sexes of which differ more in size than those of any other breed (though blood-hounds differ considerably), or than in any wild canine species known to me.
Accordingly, I applied to Mr. Cupples, a well-known breeder of these dogs, who has weighed and measured many of his own dogs, and who, with great kindness, has collected for me the following facts from various sources. Superior male dogs, measured at the shoulder, range from twenty-eight inches, which is low, to thirty-three, or even thirty-four inches in height; and in weight from eighty pounds, which is low, to 120, or even more pounds. The females range in height from twenty-three to twenty-seven, or even to twenty-eight inches; and in weight from fifty to seventy, or even eighty pounds.[321] Mr. Cupples concludes that from ninety-five to one hundred pounds for the male, and seventy for the female, would be a safe average; but there is reason to believe that formerly both sexes attained a greater weight. Mr. Cupples has weighed puppies when a fortnight old; in one litter the average weight of four males exceeded that of two females by six and a half ounces; in another litter the average weight of four males exceeded that of one female by less than one ounce; the same males, when three weeks old, exceeded the female by seven and a half ounces, and at the age of six weeks by nearly fourteen ounces. Mr. Wright of Yeldersley House, in a letter to Mr. Cupples, says: “I have taken notes on the sizes and weights of puppies of many litters, and as far as my experience goes, dog-puppies as a rule differ very little from bitches till they arrive at about five or six months old; and then the dogs begin to increase, gaining upon the bitches both in weight and size. At birth, and for several weeks afterwards, a bitch-puppy will occasionally be larger than any of the dogs, but they are invariably beaten by them later.” Mr. McNeill, of Colinsay, concludes that “the males do not attain their full growth till over two years old, though the females attain it sooner.” According to Mr. Cupples’ experience, male dogs go on growing in stature till they are from twelve to eighteen months old, and in weight till from eighteen to twenty-four months old; whilst the females cease increasing in stature at the age of from nine to fourteen or fifteen months, and in weight at the age of from twelve to fifteen months. From these various statements it is clear that the full difference in size between the male and female Scotch deerhound is not acquired until rather late in life. The males are almost exclusively used for coursing, for, as Mr. McNeill informs me, the females have not sufficient strength and weight to pull down a full-grown deer. From the names used in old legends, it appears, as I hear from Mr. Cupples, that at a very ancient period the males were the most celebrated, the females being mentioned only as the mothers of famous dogs. Hence during many generations, it is the male which has been chiefly tested for strength, size, speed, and courage, and the best will have been bred from. As, however, the males do not attain their full dimensions until a rather late period in life, they will have tended, in accordance with the law often indicated, to transmit their characters to their male offspring alone; and thus the great inequality in size between the sexes of the Scotch deerhound may probably be accounted for.
The males of some few quadrupeds possess organs or parts developed solely as a means of defence against the attacks of other males. Some kinds of deer use, as we have seen, the upper branches of their horns chiefly or exclusively for defending themselves; and the Oryx antelope, as I am informed by Mr. Bartlett, fences most skilfully with his long, gently curved horns; but these are likewise used as organs of offence. Rhinoceroses, as the same observer remarks, in fighting parry each other’s sidelong blows with their horns, which loudly clatter together, as do the tusks of boars. Although wild boars fight desperately together, they seldom, according to Brehm, receive fatal blows, as these fall on each other’s tusks, or on the layer of gristly skin covering the shoulder, which the German hunters call the shield; and here we have a part specially modified for defence. With boars in the prime
Fig. 63. Head of common wild boar, in prime of life (from Brehm). of life (see fig. 63) the tusks in the lower jaw are used for fighting but they become in old age, as Brehm states, so much curved inwards and upwards, over the snout, that they can no longer be thus used. They may, however, still continue to serve, and even in a still more effective manner, as a means of defence. In compensation for the loss of the lower tusks as weapons of offence, those in the upper jaw, which always project a little laterally, increase so much in length during old age, and curve so much upwards, that they can be used as a means of attack. Nevertheless an old boar is not so dangerous to man as one at the age of six or seven years.[322]
Fig. 64. Skull of the Babirusa Pig (from Wallace’s ‘Malay Archipelago’).
In the full-grown male Babirusa pig of Celebes (fig. [64]), the lower tusks are formidable weapons, like those of the European boar in the prime of life, whilst the upper tusks are so long and have their points so much curled inwards, sometimes even touching the forehead, that they are utterly useless as weapons of attack. They more nearly resemble horns than teeth, and are so manifestly useless as teeth that the animal was formerly supposed to rest his head by hooking them on to a branch. Their convex surfaces would, however, if the head were held a little laterally, serve as an excellent guard; and hence, perhaps it is that in old animals they “are generally broken off, as if by fighting.”[323] Here, then, we have the curious case of the upper tusks of the Babirusa regularly assuming during the prime of life, a structure which apparently renders them fitted only for defence; whilst in the European boar the lower and opposite tusks assume in a less degree and only during old age nearly the same form, and then serve in like manner solely for defence.
Fig. 65. Head of Æthiopian Wart-hog, from ‘Proc. Zool. Soc.’ 1869. (I now find that this drawing represents the head of a female, but it serves to shew, on a reduced scale, the characters of the male.)
In the wart-hog (Phacochœrus æthiopicus, fig. 65) the tusks in the upper jaw of the male curve upwards during the prime of life, and from being pointed, serve as formidable weapons. The tusks in the lower jaw are sharper than those in the upper, but from their shortness it seems hardly possible that they can be used as weapons of attack. They must, however, greatly strengthen those in the upper jaw, from being ground so as to fit closely against their bases. Neither the upper nor the lower tusks appear to have been specially modified to act as guards, though, no doubt, they are thus used to a certain extent. But the wart-hog is not destitute of other special means of protection, for there exists, on each side of the face, beneath the eyes, a rather stiff, yet flexible, cartilaginous, oblong pad (fig. [65]), which projects two or three inches outwards; and it appeared to Mr. Bartlett and myself, when viewing the living animal, that these pads, when struck from beneath by the tusks of an opponent, would be turned upwards, and would thus protect in an admirable manner the somewhat prominent eyes. These boars, as I may add on the authority of Mr. Bartlett, when fighting together, stand directly face to face.
Lastly, the African river-hog (Potamochoerus penicillatus) has a hard cartilaginous knob on each side of the face beneath the eyes, which answers to the flexible pad of the wart-hog; it has also two bony prominences on the upper jaw above the nostrils. A boar of this species in the Zoological Gardens recently broke into the cage of the wart-hog. They fought all night-long, and were found in the morning much exhausted, but not seriously wounded. It is a significant fact, as shewing the purpose of the above-described projections and excrescences, that these were covered with blood, and were scored and abraded in an extraordinary manner.
The mane of the lion forms a good defence against the one danger to which he is liable, namely the attacks of rival lions: for the males, as Sir. A. Smith informs me, engage in terrible battles, and a young lion dares not approach an old one. In 1857 a tiger at Bromwich broke into the cage of a lion, and a fearful scene ensued; “the lion’s mane saved his neck and head from being much injured, but the tiger at last succeeded in ripping up his belly, and in a few minutes he was dead.”[324] The broad ruff round the throat and chin of the Canadian lynx (Felis Canadensis) is much longer in the male than in the female; but whether it serves as a defence I do not know. Male seals are well known to fight desperately together, and the males of certain kinds (Otaria jubata)[325] have great manes, whilst the females have small ones or none. The male baboon of the Cape of Good Hope (Cynocephalus porcarius) has a much longer mane and larger canine teeth than the female; and the mane probably serves as a protection, for on asking the keepers in the Zoological Gardens, without giving them any clue to my object, whether any of the monkeys especially attacked each other by the nape of the neck, I was answered that this was not the case, excepting with the above baboon. In the Hamadryas baboon, Ehrenberg compares the mane of the adult male to that of a young lion, whilst in the young of both sexes and in the female the mane is almost absent.
It appeared to me probable that the immense woolly mane of the male American bison, which reaches almost to the ground, and is much more developed in the males than in the females, served as a protection to them in their terrible battles; but an experienced hunter told Judge Caton that he had never observed anything which favoured this belief. The stallion has a thicker and fuller mane than the mare; and I have made particular inquiries of two great trainers and breeders who have had charge of many entire horses, and am assured that they “invariably endeavour to seize one another by the neck.” It does not, however, follow from the foregoing statements, that when the hair on the neck serves as a defence, that it was originally developed for this purpose, though this is probable in some cases, as in that of the lion. I am informed by Mr. McNeill that the long hairs on the throat of the stag (Cervus elephas) serve as a great protection to him when hunted, for the dogs generally endeavour to seize him by the throat; but it is not probable that these hairs were specially developed for this purpose; otherwise the young and the females would, as we may feel assured, have been equally protected.
On Preference or Choice in Pairing, as shewn by either sex of Quadrupeds.—Before describing, in the next chapter, the differences between the sexes in voice, odour emitted, and ornamentation, it will be convenient here to consider whether the sexes exert any choice in their unions. Does the female prefer any particular male, either before or after the males may have fought together for supremacy; or does the male, when not a polygamist, select any particular female? The general impression amongst breeders seems to be that the male accepts any female; and this, owing to his eagerness, is, in most cases, probably the truth. Whether the female as a general rule indifferently accepts any male is much more doubtful. In the fourteenth chapter, on Birds, a considerable body of direct and indirect evidence was advanced, shewing that the female selects her partner; and it would be a strange anomaly if female quadrupeds, which stand higher in the scale of organisation and have higher mental powers, did not generally, or at least often, exert some choice. The female could in most cases escape, if wooed by a male that did not please or excite her; and when pursued, as so incessantly occurs, by several males, she would often have the opportunity, whilst they were fighting together, of escaping with, or at least of temporarily pairing with, some one male. This latter contingency has often been observed in Scotland with female red-deer, as I have been informed by Sir Philip Egerton.[326]
It is scarcely possible that much should be known about female quadrupeds exerting in a state of nature any choice in their marriage unions. The following very curious details on the courtship of one of the eared seals, Callorhinus ursinus, are given[327] 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.”
As so little is known about the courtship of animals in a state of nature, I have endeavoured to discover how far our domesticated quadrupeds evince any choice in their unions. Dogs offer the best opportunity for observation, as they are carefully attended to and well understood. Many breeders have expressed a strong opinion on this head. Thus Mr. Mayhew remarks, “The females are able to bestow their affections; and tender recollections are as potent over them as they are known to be in other cases, where higher animals are concerned. Bitches are not always prudent in their loves, but are apt to fling themselves away on curs of low degree. If reared with a companion of vulgar appearance, there often springs up between the pair a devotion which no time can afterwards subdue. The passion, for such it really is, becomes of a more than romantic endurance.” Mr. Mayhew, who attended chiefly to the smaller breeds, is convinced that the females are strongly attracted by males of large size.[328] The well-known veterinary Blaine states[329] that his own female pug became so attached to a spaniel, and a female setter to a cur, that in neither case would they pair with a dog of their own breed until several weeks had elapsed. Two similar and trustworthy accounts have been given me in regard to a female retriever and a spaniel, both of which became enamoured with terrier-dogs.
Mr. Cupples informs me that he can personally vouch for the accuracy of the following more remarkable case, in which a valuable and wonderfully-intelligent female terrier loved a retriever, belonging to a neighbour, to such a degree that she had often to be dragged away from him. After their permanent separation, although repeatedly shewing milk in her teats, she would never acknowledge the courtship of any other dog, and to the regret of her owner, never bore puppies. Mr. Cupples also states that a female deerhound now (1868) in his kennel has thrice produced puppies, and on each occasion shewed a marked preference for one of the largest and handsomest, but not the most eager, of four deerhounds living with her, all in the prime of life. Mr. Cupples has observed that the female generally favours a dog whom she has associated with and knows; her shyness and timidity at first incline her against a strange dog. The male, on the contrary, seems rather inclined towards strange females. It appears to be rare when the male refuses any particular female, but Mr. Wright, of Yeldersley House, a great breeder of dogs, informs me that he has known some instances; he cites the case of one of his own deerhounds, who would not take any notice of a particular female mastiff, so that another deerhound had to be employed. It would be superfluous to give other cases, and I will only add that Mr. Barr, who has carefully bred many blood-hounds, states that in almost every instance particular individuals of the opposite sex shew a decided preference for each other. Finally Mr. Cupples, after attending to this subject for another year, has recently written to me, “I have had full confirmation of my former statement, that dogs in breeding form decided preferences for each other, being often influenced by size, bright colour, and individual character, as well as by the degree of their previous familiarity.”
In regard to horses, Mr. Blenkiron, the greatest breeder of race-horses in the world, informs me that stallions are so frequently capricious in their choice, rejecting one mare and without any apparent cause taking to another, that various artifices have to be habitually used. The famous Monarque, for instance, would never consciously look at the dam of Gladiateur, and a trick had to be practised. We can partly see the reason why valuable race-horse stallions, which are in such demand, should be so particular in their choice. Mr. Blenkiron has never known a mare to reject a horse; but this has occurred in Mr. Wright’s stable, so that the mare had to be cheated. Prosper Lucas[330] quotes various statements from French authorities, and remarks, “On voit des étalons qui s’éprennent d’une jument, et négligent toutes les autres.” He gives, on the authority of Baëlen, similar facts in regard to bulls. Hoffberg, in describing the domesticated reindeer of Lapland, says, “Fœmina majores et fortiores mares præ cæteris admittunt, ad eos confugiunt, a junioribus agitatæ, qui hos in fugam conjiciunt.”[331] A clergy-man, who has bred many pigs, assures me that sows often reject one boar and immediately accept another.
From these facts there can be no doubt that with most of our domesticated quadrupeds strong individual antipathies and preferences are frequently exhibited, and much more commonly by the female than by the male. This being the case, it is improbable that the unions of quadrupeds in a state of nature should be left to mere chance. It is much more probable that the females are allured or excited by particular males, who possess certain characters in a higher degree than other males; but what these characters are, we can seldom or never discover with certainty.