shade; secondly, pure white; and thirdly, owing to another change of fashion (if I may so express myself), their present slaty, reddish, or golden-buff tints. These successive changes are intelligible only on the principle of novelty having been admired by birds for its own sake.
Several writers have objected to the whole theory of sexual selection, by assuming that with animals and savages the taste of the female for certain colours or other ornaments would not remain constant for many generations; that first one colour and then another would be admired, and consequently that no permanent effect could be produced. We may admit that taste is fluctuating, but it is not quite arbitrary. It depends much on habit, as we see in mankind; and we may infer that this would hold good with birds and other animals. Even in our own dress, the general character lasts long, and the changes are to a certain extent graduated. Abundant evidence will be given in two places in a future chapter, that savages of many races have admired for many generations the same cicatrices on the skin, the same hideously perforated lips, nostrils, or ears, distorted heads, etc.; and these deformities present some analogy to the natural ornaments of various animals. Nevertheless, with savages such fashions do not endure for ever, as we may infer from the differences in this respect between allied tribes on the same continent. So again the raisers of fancy animals certainly have admired for many generations and still admire the same breeds; they earnestly desire slight changes, which are considered as improvements, but any great or sudden change is looked at as the greatest blemish. With birds in a state of nature we have no reason to suppose that they would admire an entirely new style of coloration, even if great and sudden variations often occurred, which is far from being the case. We know that dovecot pigeons do not willingly associate with the variously coloured fancy breeds; that albino birds do not commonly get partners in marriage; and that the black ravens of the Feroe Islands chase away their piebald brethren. But this dislike of a sudden change would not preclude their appreciating slight changes, any more than it does in the case of man. Hence with respect to taste, which depends on many elements, but partly on habit and partly on a love of novelty, there seems no improbability in animals admiring for a very long period the same general style of ornamentation or other attractions, and yet appreciating slight changes in colours, form, or sound.
SUMMARY OF THE FOUR CHAPTERS ON BIRDS.
Most male birds are highly pugnacious during the breeding-season, and some possess weapons adapted for fighting with their rivals. But the most pugnacious and the best armed males rarely or never depend for success solely on their power to drive away or kill their rivals, but have special means for charming the female. With some it is the power of song, or of giving forth strange cries, or instrumental music, and the males in consequence differ from the females in their vocal organs, or in the structure of certain feathers. From the curiously diversified means for producing various sounds, we gain a high idea of the importance of this means of courtship. Many birds endeavour to charm the females by love- dances or antics, performed on the ground or in the air, and sometimes at prepared places. But ornaments of many kinds, the most brilliant tints, combs and wattles, beautiful plumes, elongated feathers, top-knots, and so forth, are by far the commonest means. In some cases mere novelty appears to have acted as a charm. The ornaments of the males must be highly important to them, for they have been acquired in not a few cases at the cost of increased danger from enemies, and even at some loss of power in fighting with their rivals. The males of very many species do not assume their ornamental dress until they arrive at maturity, or they assume it only during the breeding-season, or the tints then become more vivid. Certain ornamental appendages become enlarged, turgid, and brightly coloured during the act of courtship. The males display their charms with elaborate care and to the best effect; and this is done in the presence of the females. The courtship is sometimes a prolonged affair, and many males and females congregate at an appointed place. To suppose that the females do not appreciate the beauty of the males, is to admit that their splendid decorations, all their pomp and display, are useless; and this is incredible. Birds have fine powers of discrimination, and in some few instances it can be shewn that they have a taste for the beautiful. The females, moreover, are known occasionally to exhibit a marked preference or antipathy for certain individual males.
If it be admitted that the females prefer, or are unconsciously excited by the more beautiful males, then the males would slowly but surely be rendered more and more attractive through sexual selection. That it is this sex which has been chiefly modified, we may infer from the fact that, in almost every genus where the sexes differ, the males differ much more from one another than do the females; this is well shewn in certain closely-allied representative species, in which the females can hardly be distinguished, whilst the males are quite distinct. Birds in a state of nature offer individual differences which would amply suffice for the work of sexual selection; but we have seen that they occasionally present more strongly marked variations which recur so frequently that they would immediately be fixed, if they served to allure the female. The laws of variation must determine the nature of the initial changes, and will have largely influenced the final result. The gradations, which may be observed between the males of allied species, indicate the nature of the steps through which they have passed. They explain also in the most interesting manner how certain characters have originated, such as the indented ocelli on the tail-feathers of the peacock, and the ball-and-socket ocelli on the wing-feathers of the Argus pheasant. It is evident that the brilliant colours, top-knots, fine plumes, etc., of many male birds cannot have been acquired as a protection; indeed, they sometimes lead to danger. That they are not due to the direct and definite action of the conditions of life, we may feel assured, because the females have been exposed to the same conditions, and yet often differ from the males to an extreme degree. Although it is probable that changed conditions acting during a lengthened period have in some cases produced a definite effect on both sexes, or sometimes on one sex alone, the more important result will have been an increased tendency to vary or to present more strongly-marked individual differences; and such differences will have afforded an excellent ground- work for the action of sexual selection.
The laws of inheritance, irrespectively of selection, appear to have determined whether the characters acquired by the males for the sake of ornament, for producing various sounds, and for fighting together, have been transmitted to the males alone or to both sexes, either permanently, or periodically during certain seasons of the year. Why various characters should have been transmitted sometimes in one way and sometimes in another, is not in most cases known; but the period of variability seems often to have been the determining cause. When the two sexes have inherited all characters in common they necessarily resemble each other; but as the successive variations may be differently transmitted, every possible gradation may be found, even within the same genus, from the closest similarity to the widest dissimilarity between the sexes. With many closely-allied species, following nearly the same habits of life, the males have come to differ from each other chiefly through the action of sexual selection; whilst the females have come to differ chiefly from partaking more or less of the characters thus acquired by the males. The effects, moreover, of the definite action of the conditions of life, will not have been masked in the females, as in the males, by the accumulation through sexual selection of strongly-pronounced colours and other ornaments. The individuals of both sexes, however affected, will have been kept at each successive period nearly uniform by the free intercrossing of many individuals.
With species, in which the sexes differ in colour, it is possible or probable that some of the successive variations often tended to be transmitted equally to both sexes; but that when this occurred the females were prevented from acquiring the bright colours of the males, by the destruction which they suffered during incubation. There is no evidence that it is possible by natural selection to convert one form of transmission into another. But there would not be the least difficulty in rendering a female dull-coloured, the male being still kept bright- coloured, by the selection of successive variations, which were from the first limited in their transmission to the same sex. Whether the females of many species have actually been thus modified, must at present remain doubtful. When, through the law of the equal transmission of characters to both sexes, the females were rendered as conspicuously coloured as the males, their instincts appear often to have been modified so that they were led to build domed or concealed nests.
In one small and curious class of cases the characters and habits of the two sexes have been completely transposed, for the females are larger, stronger, more vociferous and brighter coloured than the males. They have, also, become so quarrelsome that they often fight together for the possession of the males, like the males of other pugnacious species for the possession of the females. If, as seems probable, such females habitually drive away their rivals, and by the display of their bright colours or other charms endeavour to attract the males, we can understand how it is that they have gradually been rendered, by sexual selection and sexually- limited transmission, more beautiful than the males–the latter being left unmodified or only slightly modified.
Whenever the law of inheritance at corresponding ages prevails but not that of sexually-limited transmission, then if the parents vary late in life– and we know that this constantly occurs with our poultry, and occasionally with other birds–the young will be left unaffected, whilst the adults of both sexes will be modified. If both these laws of inheritance prevail and either sex varies late in life, that sex alone will be modified, the other sex and the young being unaffected. When variations in brightness or in other conspicuous characters occur early in life, as no doubt often happens, they will not be acted on through sexual selection until the period of reproduction arrives; consequently if dangerous to the young, they will be eliminated through natural selection. Thus we can understand how it is that variations arising late in life have so often been preserved for the ornamentation of the males; the females and the young being left almost unaffected, and therefore like each other. With species having a distinct summer and winter plumage, the males of which either resemble or differ from the females during both seasons or during the summer alone, the degrees and kinds of resemblance between the young and the old are exceedingly complex; and this complexity apparently depends on characters, first acquired by the males, being transmitted in various ways and degrees, as limited by age, sex, and season.
As the young of so many species have been but little modified in colour and in other ornaments, we are enabled to form some judgment with respect to the plumage of their early progenitors; and we may infer that the beauty of our existing species, if we look to the whole class, has been largely increased since that period, of which the immature plumage gives us an indirect record. Many birds, especially those which live much on the ground, have undoubtedly been obscurely coloured for the sake of protection. In some instances the upper exposed surface of the plumage has been thus coloured in both sexes, whilst the lower surface in the males alone has been variously ornamented through sexual selection. Finally, from the facts given in these four chapters, we may conclude that weapons for battle, organs for producing sound, ornaments of many kinds, bright and conspicuous colours, have generally been acquired by the males through variation and sexual selection, and have been transmitted in various ways according to the several laws of inheritance–the females and the young being left comparatively but little modified. (57. I am greatly indebted to the kindness of Mr. Sclater for having looked over these four chapters on birds, and the two following ones on mammals. In this way I have been saved from making mistakes about the names of the species, and from stating anything as a fact which is known to this distinguished naturalist to be erroneous. But, of course, he is not at all answerable for the accuracy of the statements quoted by me from various authorities.)
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 shown 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.” (1. See Waterton’s account of two hares fighting, ‘Zoologist,’ vol. i. 1843, p. 211. On moles, Bell, ‘Hist. of British Quadrupeds,’ 1st ed., p. 100. On squirrels, Audubon and Bachman, Viviparous Quadrupeds of N. America, 1846, p. 269. On beavers, Mr. A.H. Green, in ‘Journal of Linnean Society, Zoology,’ vol. x. 1869, p. 362.) 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 their lower jaws often become distorted. (2. On the battles of seals, see Capt. C. Abbott in ‘Proc. Zool. Soc.’ 1868, p. 191; Mr. R. Brown, ibid. 1868, p. 436; also L. Lloyd, ‘Game Birds of Sweden,’ 1867, p. 412; also Pennant. On the sperm-whale see Mr. J.H. Thompson, in ‘Proc. Zool. Soc.’ 1867, p. 246.)
All male animals which are furnished with special weapons for fighting, are well known to engage in fierce battles. The courage and the desperate conflicts of stags have often been described; their skeletons have been found in various parts of the world, with the horns inextricably locked together, shewing how miserably the victor and vanquished had perished. (3. See Scrope (‘Art of Deer-stalking,’ p. 17) on the locking of the horns with the Cervus elaphus. Richardson, in ‘Fauna Bor. Americana,’ 1829, p. 252, says that the wapiti, moose, and reindeer have been found thus locked together. Sir. A. Smith found at the Cape of Good Hope the skeletons of two gnus in the same condition.) No animal in the world is so dangerous as an elephant in must. Lord Tankerville has given me a graphic description of the battles between the wild bulls in Chillingham Park, the descendants, degenerated in size but not in courage, of the gigantic Bos primigenius. In 1861 several contended for mastery; and it was observed that two of the younger bulls attacked in concert the old leader of the herd, overthrew and disabled him, so that he was believed by the keepers to be lying mortally wounded in a neighbouring wood. But a few days afterwards one of the young bulls approached the wood alone; and then the “monarch of the chase,” who had been lashing himself up for vengeance, came out and, in a short time, killed his antagonist. He then quietly joined the herd, and long held undisputed sway. Admiral Sir B.J. Sulivan informs me that, when he lived in the Falkland Islands, he imported a young English stallion, which frequented the hills near Port William with eight mares. On these hills there were two wild stallions, each with a small troop of mares; “and it is certain that these stallions would never have approached each other without fighting. Both had tried singly to fight the English horse and drive away his mares, but had failed. One day they came in TOGETHER and attacked him. This was seen by the 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 which are provided with efficient cutting or tearing teeth for the ordinary purposes of life, such as the carnivora, insectivora, and rodents, are seldom furnished with weapons especially adapted for fighting with their rivals. The case is very different with the males of many other animals. We see this in the horns of stags and of certain kinds of antelopes in which the females are hornless. With many animals the canine teeth in the upper or lower jaw, or in both, are much larger in the males than in the females, or are absent in the latter, with the exception sometimes of a hidden rudiment. Certain antelopes, the musk-deer, camel, horse, boar, various apes, seals, and the walrus, offer instances. In the females of the walrus the tusks are sometimes quite absent. (4. Mr. Lamont (‘Seasons with the Sea-Horses,’ 1861, p. 143) says that a good tusk of the male walrus weighs 4 pounds, and is longer than that of the female, which weighs about 3 pounds. The males are described as fighting ferociously. On the occasional absence of the tusks in the female, see Mr. R. Brown, ‘Proceedings, Zoological Society,’ 1868, p. 429.) In the male elephant of India and in the male dugong (5. Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 283.) the upper incisors form offensive weapons. In the male narwhal the left canine alone is developed into the well-known, spirally-twisted, so-called horn, which is sometimes from nine to ten feet in length. It is believed that the males use these horns for fighting together; for “an unbroken one can rarely be got, and occasionally one may be found with the point of another jammed into the broken place.” (6. Mr. R. Brown, in ‘Proc. Zool. Soc.’ 1869, p. 553. See Prof. Turner, in ‘Journal of Anat. and Phys.’ 1872, p. 76, on the homological nature of these tusks. Also Mr. J.W. Clarke on two tusks being developed in the males, in ‘Proceedings of the Zoological Society,’ 1871, p. 42.) The tooth on the opposite side of the head in the male consists of a rudiment about ten inches in length, which is embedded in the jaw; but sometimes, though rarely, both are equally developed on the two sides. In the female both are always rudimentary. The male cachalot has a larger head than that of the female, and it no doubt aids him in his aquatic battles. Lastly, the adult male ornithorhynchus is provided with a remarkable apparatus, namely a spur on the foreleg, closely resembling the poison-fang of a venomous snake; but according to Harting, the secretion from the gland is not poisonous; and on the leg of the female there is a hollow, apparently for the reception of the spur. (7. Owen on the cachalot and Ornithorhynchus, ibid. vol. iii. pp. 638, 641. Harting is quoted by Dr. Zouteveen in the Dutch translation of this work, vol. ii. p. 292.)
When the males are provided with weapons which in the females are absent, there can be hardly a doubt that these serve for fighting with other males; and that they were acquired through sexual selection, and were transmitted to the male sex alone. It is not probable, at least in most cases, that the females have been prevented from acquiring such weapons, on account of their being useless, superfluous, or in some way injurious. On the contrary, as they are often used by the males for various purposes, more especially as a defence against their enemies, it is a surprising fact that they are so poorly developed, or quite absent, in the females of so many animals. With female deer the development during each recurrent season of great branching horns, and with female elephants the development of immense tusks, would be a great waste of vital power, supposing that they were of no use to the females. Consequently, they would have tended to be eliminated in the female through natural selection; that is, if the successive variations were limited in their transmission to the female sex, for otherwise the weapons of the males would have been injuriously affected, and this would have been a greater evil. On the whole, and from the consideration of the following facts, it seems probable that when the various weapons differ in the two sexes, this has generally depended on the kind of transmission which has prevailed.
As the reindeer is the one species in the whole family of Deer, in which the female is furnished with horns, though they are somewhat smaller, thinner, and less branched than in the male, it might naturally be thought that, at least in this case, they must be of some special service to her. The female retains her horns from the time when they are fully developed, namely, in September, throughout the winter until April or May, when she brings forth her young. Mr. Crotch made particular enquiries for me in Norway, and it appears that the females at this season conceal themselves for about a fortnight in order to bring forth their young, and then reappear, generally hornless. In Nova Scotia, however, as I hear from Mr. H. Reeks, the female sometimes retains her horns longer. The male on the other hand casts his horns much earlier, towards the end of November. As both sexes have the same requirements and follow the same habits of life, and as the male is destitute of horns during the winter, it is improbable that they can be of any special service to the female during this season, which includes the larger part of the time during which she is horned. Nor is it probable that she can have inherited horns from some ancient progenitor of the family of deer, for, from the fact of the females of so many species in all quarters of the globe not having horns, we may conclude that this was the primordial character of the group. (8. On the structure and shedding of the horns of the reindeer, Hoffberg, ‘Amoenitates Acad.’ vol. iv. 1788, p. 149. See Richardson, ‘Fauna Bor. Americana,’ p. 241, in regard to the American variety or species: also Major W. Ross King, ‘The Sportsman in Canada,’ 1866, p. 80.
The horns of the reindeer are developed at a most unusually early age; but what the cause of this may be is not known. The effect has apparently been the transference of the horns to both sexes. We should bear in mind that horns are always transmitted through the female, and that she has a latent capacity for their development, as we see in old or diseased females. (9. Isidore Geoffroy St.-Hilaire, ‘Essais de Zoolog. Generale,’ 1841, p. 513. Other masculine characters, besides the horns, are sometimes similarly transferred to the female; thus Mr. Boner, in speaking of an old female chamois (‘Chamois Hunting in the Mountains of Bavaria,’ 1860, 2nd ed., p. 363), says, “not only was the head very male-looking, but along the back there was a ridge of long hair, usually to be found only in bucks.”) Moreover the females of some other species of deer exhibit, either normally or occasionally, rudiments of horns; thus the female of Cervulus moschatus has “bristly tufts, ending in a knob, instead of a horn”; and “in most specimens of the female wapiti (Cervus canadensis) there is a sharp bony protuberance in the place of the horn.” (10. On the Cervulus, Dr. Gray, ‘Catalogue of Mammalia in the British Museum,’ part iii. p. 220. On the Cervus canadensis or wapiti, see Hon. J.D. Caton, ‘Ottawa Academy of Nat. Sciences,’ May 1868, p. 9.) From these several considerations we may conclude that the possession of fairly well-developed horns by the female reindeer, is due to the males having first acquired them as weapons for fighting with other males; and secondarily to their development from some unknown cause at an unusually early age in the males, and their consequent transference to both sexes.
Turning to the sheath-horned ruminants: with antelopes a graduated series can be formed, beginning with species, the females of which are completely destitute of horns–passing on to those which have horns so small as to be almost rudimentary (as with the Antilocapra americana, in which species they are present in only one out of four or five females (11. I am indebted to Dr. Canfield for this information; see also his paper in the ‘Proceedings of the Zoological Society,’ 1866, p. 105.))–to those which have fairly developed horns, but manifestly smaller and thinner than in the male and sometimes of a different shape (12. For instance the horns of the female Ant. euchore resemble those of a distinct species, viz. the Ant. dorcas var. Corine, see Desmarest, ‘Mammalogie,’ p. 455.),–and ending with those in which both sexes have horns of equal size. As with the reindeer, so with antelopes, there exists, as previously shewn, a relation between the period of the development of the horns and their transmission to one or both sexes; it is therefore probable that their presence or absence in the females of some species, and their more or less perfect condition in the females of other species, depends, not on their being of any special use, but simply on inheritance. It accords with this view that even in the same restricted genus both sexes of some species, and the males alone of others, are thus provided. It is also a remarkable fact that, although the females of Antilope bezoartica are normally destitute of horns, Mr. Blyth has seen no less than three females thus furnished; and there was no reason to suppose that they were old or diseased.
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. (13. Gray, ‘Catalogue of Mammalia, the British Museum,’ part iii. 1852, p. 160.) In several domestic breeds of these two animals, the males alone are furnished with horns; and in some breeds, for instance, in the sheep of North Wales, though both sexes are properly horned, the ewes are very liable to be hornless. I have been informed by a trustworthy witness, who purposely inspected a flock of these same sheep during the lambing season, that the horns at birth are generally more fully developed in the male than in the female. Mr. J. Peel crossed his Lonk sheep, both sexes of which always bear horns, with hornless Leicesters and hornless Shropshire Downs; and the result was that the male offspring had their horns considerably reduced, whilst the females were wholly destitute of them. These several facts indicate that, with sheep, the horns are a much less firmly fixed character in the females than in the males; and this leads us to look at the horns as properly of masculine origin.
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. (14. Richardson, ‘Fauna Bor. Americana,’ p. 278.) 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.” (15. ‘Land and Water,’ 1867, p. 346.) Dr. Forsyth Major also informs me that a fossil skull, believed to be that of the female Bos etruscus, has been found in Val d’Arno, which is wholly without horns. In the Rhinoceros simus, as I may add, the horns of the female are generally longer but less powerful than in the male; and in some other species of rhinoceros they are said to be shorter in the female. (16. Sir Andrew Smith, ‘Zoology of S. Africa,’ pl. xix. Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 624.) From these various facts we may infer as probable that horns of all kinds, even when they are equally developed in the two sexes, were primarily acquired by the male in order to conquer other males, and have been transferred more or less completely to the female.
The effects of castration deserve notice, as throwing light on this same point. Stags after the operation never renew their horns. The male reindeer, however, must be excepted, as after castration he does renew them. This fact, as well as the possession of horns by both sexes, seems at first to prove that the horns in this species do not constitute a sexual character (17. This is the conclusion of Seidlitz, ‘Die Darwinsche Theorie,’ 1871, p. 47.); but as they are developed at a very early age, before the sexes differ in constitution, it is not surprising that they should be unaffected by castration, even if they were aboriginally acquired by the male. With sheep both sexes properly bear horns; and I am informed that with Welch sheep the horns of the males are considerably reduced by castration; but the degree depends much on the age at which the operation is performed, as is likewise the case with other animals. Merino rams have large horns, whilst the ewes “generally speaking are without horns”; and in this breed castration seems to produce a somewhat greater effect, so that if performed at an early age the horns “remain almost undeveloped.” (18. I am much obliged to Prof. Victor Carus, for having made enquiries for me in Saxony on this subject. H. von Nathusius (‘Viehzucht,’ 1872, p. 64) says that the horns of sheep castrated at an early period, either altogether disappear or remain as mere rudiments; but I do not know whether he refers to merinos or to ordinary breeds.) On the Guinea coast there is a breed in which the females never bear horns, and, as Mr. Winwood Reade informs me, the rams after castration are quite destitute of them. With cattle, the horns of the males are much altered by castration; for instead of being short and thick, they become longer than those of the cow, but otherwise resemble them. The Antilope bezoartica offers a somewhat analogous case: the males have long straight spiral horns, nearly parallel to each other, and directed backwards; the females occasionally bear horns, but these when present are of a very different shape, for they are not spiral, and spreading widely, bend round with the points forwards. Now it is a remarkable fact that, in the castrated male, as Mr. Blyth informs me, the horns are of the same peculiar shape as in the female, but longer and thicker. If we may judge from analogy, the female probably shews us, in these two cases of cattle and the antelope, the former condition of the horns in some early progenitor of each species. But why castration should lead to the reappearance of an early condition of the horns cannot be explained with any certainty. Nevertheless, it seems probable, that in nearly the same manner as the constitutional disturbance in the offspring, caused by a cross between two distinct species or races, often leads to the reappearance of long-lost characters (19. I have given various experiments and other evidence proving that this is the case, in my ‘Variation of Animals and Plants under Domestication,’ vol. ii. 1868, pp. 39-47.); so here, the disturbance in the constitution of the individual, resulting from castration, produces the same effect.
The tusks of the elephant, in the different species or races, differ according to sex, nearly as do the horns of ruminants. In India and Malacca the males alone are provided with well-developed tusks. The elephant of Ceylon is considered by most naturalists as a distinct race, but by some as a distinct species, and here “not one in a hundred is found with tusks, the few that possess them being exclusively males.” (20. Sir J. Emerson Tennent, ‘Ceylon,’ 1859, vol. ii. p. 274. For Malacca, ‘Journal of Indian Archipelago,’ vol. iv. p. 357.) 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, and their frequent absence in the female of Antilocapra americana–the presence of two tusks in some few male narwhals–the complete absence of tusks in some female walruses–are all instances of the extreme variability of secondary sexual characters, and of their liability to differ in closely-allied forms.
Although tusks and horns appear in all cases to have been primarily developed as sexual weapons, they often serve other purposes. The elephant uses his tusks in attacking the tiger; according to Bruce, he scores the trunks of trees until they can be thrown down easily, and he likewise thus extracts the farinaceous cores of palms; in Africa he often uses one tusk, always the same, to probe the ground and thus ascertain whether it will bear his weight. The common bull defends the herd with his horns; and the elk in Sweden has been known, according to Lloyd, to strike a wolf dead with a single blow of his great horns. Many similar facts could be given. One of the most curious secondary uses to which the horns of an animal may be occasionally put is that observed by Captain Hutton (21. ‘Calcutta Journal of Natural History,’ vol. ii, 1843, p. 526.) with the wild goat (Capra aegagrus) of the Himalayas and, as it is also said with the ibex, namely that when the male accidentally falls from a height he bends inwards his head, and by alighting on his massive horns, breaks the shock. The female cannot thus use her horns, which are smaller, but from her more quiet disposition she does not need this strange kind of shield so much.
Each male animal uses his weapons in his own peculiar fashion. The common ram makes a charge and butts with such force with the bases of his horns, that I have seen a powerful man knocked over like a child. Goats and certain species of sheep, for instance the Ovis cycloceros of Afghanistan (22. Mr. Blyth, in ‘Land and Water,’ March, 1867, p. 134, on the authority of Capt. Hutton and others. For the wild Pembrokeshire goats, see the ‘Field,’ 1869, p. 150.), rear on their hind legs, and then not only butt, but “make a cut down and a jerk up, with the ribbed front of their scimitar-shaped horn, as with a sabre. When the O. cycloceros attacked a large domestic ram, who was a noted bruiser, he conquered him by the sheer novelty of his mode of fighting, always closing at once with his adversary, and catching him across the face and nose with a sharp drawing jerk of the head, and then bounding out of the way before the blow could be returned.” In Pembrokeshire a male goat, the master of a flock which during several generations had run wild, was known to have killed several males in single combat; this goat possessed enormous horns, measuring thirty-nine inches in a straight line from tip to tip. The common bull, as every one knows, gores and tosses his opponent; but the Italian buffalo is said never to use his horns: he gives a tremendous blow with his convex forehead, and then tramples on his fallen enemy with his knees–an instinct which the common bull does not possess. (23. M. E.M. Bailly, “Sur l’usage des cornes,” etc., .Annal des Sciences Nat.’ tom. ii. 1824, p. 369.) Hence a dog who pins a buffalo by the nose is immediately crushed. We must, however, remember that the Italian buffalo has been long domesticated, and it is by no means certain that the wild parent-form had similar horns. Mr. Bartlett informs me that when a female Cape buffalo (Bubalus caffer) was turned into an enclosure with a bull of the same species, she attacked him, and he in return pushed her about with great violence. But it was manifest to Mr. Bartlett that, had not the bull shewn dignified forbearance, he could easily have killed her by a single lateral thrust with his immense horns. The giraffe uses his short, hair-covered horns, which are rather longer in the male than in the female, in a curious manner; for, with his long neck, he swings his head to either side, almost upside down, with such force that I have seen a hard plank deeply indented by a single blow.
[Fig. 63. 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 springboc (Ant. euchore) has rather short upright horns, with the sharp points bent inwards almost at right angles, so as to face each other; Mr. Bartlett does not know how they are used, but suggests that they would inflict a fearful wound down each side of the face of an antagonist. The slightly-curved horns of the Oryx leucoryx (Fig. 63) are directed backwards, and are of such length that their points reach beyond the middle of the back, over which they extend in almost parallel lines. Thus they seem singularly ill-fitted for fighting; but Mr. Bartlett informs me that when two of these animals prepare for battle, they kneel down, with their beads between their fore 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 each endeavours to get the upturned points under the body of the other; if one succeeds in doing this, he suddenly springs up, throwing up his head at the same time, and can thus wound or perhaps even transfix his antagonist. Both animals always kneel down, so as to guard as far as possible against this manoeuvre. It has been recorded that one of these antelopes has used his horn with effect even against a lion; yet from being forced to place his head between the 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 branches of the horns offer a curious case of difficulty; for certainly a single straight point would inflict a much more serious wound than several diverging ones. In Sir Philip Egerton’s museum there is a horn of the red-deer (Cervus elaphus), thirty inches in length, with “not fewer than fifteen snags or branches”; and at Moritzburg there is still preserved a pair of antlers of a red-deer, shot in 1699 by Frederick I., one of which bears the astonishing number of thirty-three branches and the other twenty-seven, making altogether sixty branches. Richardson figures a pair of antlers of the wild reindeer with twenty-nine points. (24. On the horns of red-deer, Owen, ‘British Fossil Mammals,’ 1846, p. 478; Richardson on the horns of the reindeer, ‘Fauna Bor. Americana,’ 1829, p. 240. I am indebted to Prof. Victor Carus, for the Moritzburg case.) 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 (25. Hon. J.D. Caton (‘Ottawa Acad. of Nat. Science,’ May 1868, p. 9) says that the American deer fight with their fore-feet, after “the question of superiority has been once settled and acknowledged in the herd.” Bailly, ‘Sur l’Usage des cornes,’ ‘Annales des Sciences Nat.’ tom. ii. 1824, p. 371.), M. Bailly actually comes to the conclusion that their horns are more injurious than useful to them. But this author overlooks the pitched battles between rival males. As I felt much perplexed about the use or advantage of the branches, I applied to Mr. McNeill of Colonsay, who has long and carefully observed the habits of red-deer, and he informs me that he has never seen some of the branches brought into use, but that the brow antlers, from inclining downwards, are a great protection to the forehead, and their points are likewise used in attack. Sir Philip Egerton also informs me both as to red-deer and fallow-deer that, in fighting, they suddenly dash together, and getting their horns fixed against each other’s bodies, a desperate struggle ensues. When one is at last forced to yield and turn round, the victor endeavours to plunge his brow antlers into his defeated foe. It thus appears that the upper branches are used chiefly or exclusively for pushing and fencing. Nevertheless in some species the upper branches are used as weapons of offence; when a man was attacked by a wapiti deer (Cervus canadensis) in Judge Caton’s park in Ottawa, and several men tried to rescue him, the stag “never raised his head from the ground; in fact he kept his face almost flat on the ground, with his nose nearly between his fore feet, except when he rolled his head to one side to take a new observation preparatory to a plunge.” In this position the ends of the horns were directed against his adversaries. “In rolling his head he necessarily raised it somewhat, because his antlers were so long that he could not roll his head without raising them on one side, while, on the other side they touched the ground.” The stag by this procedure gradually drove the party of rescuers backwards to a distance of 150 or 200 feet; and the attacked man was killed. (26. See a most interesting account in the Appendix to Hon. J.D. Caton’s paper, as above quoted.)
[Fig. 64. Strepsiceros Kudu (from Sir Andrew Smith’s ‘Zoology of South Africa.’]
Although the horns of stags are efficient weapons, there can, I think be no doubt that a single point would have been much more dangerous than a branched antler; and Judge Caton, who has had large experience with deer, fully concurs in this conclusion. Nor do the branching horns, though highly important as a means of defence against rival stags, appear perfectly well adapted for this purpose, as they are liable to become interlocked. The suspicion has therefore crossed my mind that they may serve in part as ornaments. That the branched antlers of stags as well as the elegant lyrated horns of certain antelopes, with their graceful double curvature (Fig. 64), are ornamental in our eyes, no one will dispute. If, then, the horns, like the splendid accoutrements of the knights of old, add to the noble appearance of stags and antelopes, they may have been modified partly for this purpose, though mainly for actual service in battle; but I have no evidence in favour of this belief.
An interesting case has lately been published, from which it appears that the horns of a deer in one district in the United States are now being modified through sexual and natural selection. A writer in an excellent American Journal (27. The ‘American Naturalist,’ Dec. 1869, p. 552.) says, that he has hunted for the last twenty-one years in the Adirondacks, where the Cervus virginianus abounds. About fourteen years ago he first heard of SPIKE-HORN BUCKS. These became from year to year more common; about five years ago he shot one, and afterwards another, and now they are frequently killed. “The spike-horn differs greatly from the common antler of the C. virginianus. It consists of a single spike, more slender than the antler, and scarcely half so long, projecting forward from the brow, and terminating in a very sharp point. It gives a considerable advantage to its possessor over the common buck. Besides enabling him to run more swiftly through the thick woods and underbrush (every hunter knows that does and yearling bucks run much more rapidly than the large bucks when armed with their cumbrous antlers), the spike-horn is a more effective weapon than the common antler. With this advantage the spike-horn bucks are gaining upon the common bucks, and may, in time, entirely supersede them in the Adirondacks. Undoubtedly, the first spike-horn buck was merely an accidental freak of nature. But his spike-horns gave him an advantage, and enabled him to propagate his peculiarity. His descendants having a like advantage, have propagated the peculiarity in a constantly increasing ratio, till they are slowly crowding the antlered deer from the region they inhabit.” A critic has well objected to this account by asking, why, if the simple horns are now so advantageous, were the branched antlers of the parent-form ever developed? To this I can only answer by remarking, that a new mode of attack with new weapons might be a great advantage, as shewn by the case of the Ovis cycloceros, who thus conquered a domestic ram famous for his fighting power. Though the branched antlers of a stag are well adapted for fighting with his rivals, and though it might be an advantage to the prong-horned variety slowly to acquire long and branched horns, if he had to fight only with others of the same kind, yet it by no means follows that branched horns would be the best fitted for conquering a foe differently armed. In the foregoing case of the Oryx leucoryx, it is almost certain that the victory would rest with an antelope having short horns, and who therefore did not need to kneel down, though an oryx might profit by having still longer horns, if he fought only with his proper rivals.
Male quadrupeds, which are furnished with tusks, use them in various ways, as in the case of horns. The boar strikes laterally and upwards; the musk- deer downwards with serious effect. (28. Pallas, ‘Spicilegia Zoologica,’ fasc. xiii. 1779, p. 18.) The walrus, though having so short a neck and so unwieldy a body, “can strike either upwards, or downwards, or sideways, with equal dexterity.” (29. Lamont, ‘Seasons with the Sea-Horses,’ 1861, p. 141.) I was informed by the late Dr. Falconer, that the Indian elephant fights in a different manner according to the position and curvature of his tusks. When they are directed forwards and upwards he is able to fling a tiger to a great distance–it is said to even thirty feet; when they are short and turned downwards he endeavours suddenly to pin the tiger to the ground and, in consequence, is dangerous to the rider, who is liable to be jerked off the howdah. (30. See also Corse (‘Philosophical Transactions,’ 1799, p. 212) on the manner in which the short-tusked Mooknah variety attacks other elephants.)
Very few male quadrupeds possess weapons of two distinct kinds specially adapted for fighting with rival males. The male muntjac-deer (Cervulus), however, offers an exception, as he is provided with horns and exserted canine teeth. But we may infer from what follows that one form of weapon has often been replaced in the course of ages by another. With ruminants the development of horns generally stands in an inverse relation with that of even moderately developed canine teeth. Thus camels, guanacoes, chevrotains, and musk-deer, are hornless, and they have efficient canines; these teeth being “always of smaller size in the females than in the males.” The Camelidae have, in addition to their true canines, a pair of canine-shaped incisors in their upper jaws. (31. Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 349.) Male deer and antelopes, on the other hand, possess horns, and they rarely have canine teeth; and these, when present, are always of small size, so that it is doubtful whether they are of any service in their battles. In Antilope montana they exist only as rudiments in the young male, disappearing as he grows old; and they are absent in the female at all ages; but the females of certain other antelopes and of certain deer have been known occasionally to exhibit rudiments of these teeth. (32. See Ruppell (in ‘Proc. Zoolog. Soc.’ Jan. 12, 1836, p. 3) on the canines in deer and antelopes, with a note by Mr. Martin on a female American deer. See also Falconer (‘Palaeont. Memoirs and Notes,’ vol. i. 1868, p. 576) on canines in an adult female deer. In old males of the musk-deer the canines (Pallas, ‘Spic. Zoolog.’ fasc. xiii. 1779, p. 18) sometimes grow to the length of three inches, whilst in old females a rudiment projects scarcely half an inch above the gums.) Stallions have small canine teeth, which are either quite absent or rudimentary in the mare; but they do not appear to be used in fighting, for stallions bite with their incisors, and do not open their mouths wide like camels and guanacoes. Whenever the adult male possesses canines, now inefficient, whilst the female has either none or mere rudiments, we may conclude that the early male progenitor of the species was provided with efficient canines, which have been partially transferred to the females. The reduction of these teeth in the males seems to have followed from some change in their manner of fighting, often (but not in the horse) caused by the development of new weapons.
Tusks and horns are manifestly of high importance to their possessors, for their development consumes much organised matter. A single tusk of the Asiatic elephant–one of the extinct woolly species–and of the African elephant, have been known to weigh respectively 150, 160, and 180 pounds; and even greater weights have been given by some authors. (33. Emerson Tennent, ‘Ceylon,’ 1859, vol. ii. p. 275; Owen, ‘British Fossil Mammals,’ 1846, p. 245.) With deer, in which the horns are periodically renewed, the drain on the constitution must be greater; the horns, for instance, of the moose weigh from fifty to sixty pounds, and those of the extinct Irish elk from sixty to seventy pounds–the skull of the latter weighing on an average only five pounds and a quarter. Although the horns are not periodically renewed in sheep, yet their development, in the opinion of many agriculturists, entails a sensible loss to the breeder. Stags, moreover, in escaping from beasts of prey are loaded with an additional weight for the race, and are greatly retarded in passing through a woody country. The moose, for instance, with horns extending five and a half feet from tip to tip, although so skilful in their use that he will not touch or break a twig when walking quietly, cannot act so dexterously whilst rushing away from a pack of wolves. “During his progress he holds his nose up, so as to lay the horns horizontally back; and in this attitude cannot see the ground distinctly.” (34. Richardson, ‘Fauna Bor. Americana,’ on the moose, Alces palmata, pp. 236, 237; on the expanse of the horns, ‘Land and Water,’ 1869, p. 143. See also Owen, ‘British Fossil Mammals,’ on the Irish elk, pp. 447, 455.) The tips of the horns of the great Irish elk were actually eight feet apart! Whilst the horns are covered with velvet, which lasts with red-deer for about twelve weeks, they are extremely sensitive to a blow; so that in Germany the stags at this time somewhat change their habits, and avoiding dense forests, frequent young woods and low thickets. (35. ‘Forest Creatures,’ by C. Boner, 1861, p. 60.) 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 mammals, when, as is often the case, the sexes differ in size, the males are almost always larger and stronger. I am informed by Mr. Gould that this holds good in a marked manner with the marsupials of Australia, the males of which appear to continue growing until an unusually late age. But the most extraordinary case is that of one of the seals (Callorhinus ursinus), a full-grown female weighing less than one-sixth of a full-grown male. (36. See the very interesting paper by Mr. J.A. Allen in ‘Bull. Mus. Comp. Zoology of Cambridge, United States,’ vol. ii. No. 1, p. 82. The weights were ascertained by a careful observer, Capt. Bryant. Dr. Gill in ‘The American Naturalist,’ January, 1871, Prof. Shaler on the relative size of the sexes of whales, ‘American Naturalist,’ January, 1873.) Dr. Gill remarks that it is with the polygamous seals, the males of which are well known to fight savagely together, that the sexes differ much in size; the monogamous species differing but little. Whales also afford evidence of the relation existing between the pugnacity of the males and their large size compared with that of the female; the males of the right-whales do not fight together, and they are not larger, but rather smaller, than their females; on the other hand, male sperm-whales fight much together, and their bodies are “often found scarred with the imprint of their rival’s teeth,” and they are double the size of the females. The greater strength of the male, as Hunter long ago remarked (37. ‘Animal Economy,’ p. 45.), is invariably displayed in those parts of the body which are brought into action in fighting with rival males–for instance, in the massive neck of the bull. Male quadrupeds are also more courageous and pugnacious than the females. There can be little doubt that these characters have been gained, partly through sexual selection, owing to a long series of victories, by the stronger and more courageous males over the weaker, and partly through the inherited effects of use. It is probable that the successive variations in strength, size, and courage, whether due to mere variability or to the effects of use, by the accumulation of which male quadrupeds have acquired these characteristic qualities, occurred rather late in life, and were consequently to a large extent limited in their transmission to the same sex.
From these considerations I was anxious to obtain information as to the Scotch deer-hound, the sexes of which differ more in size than those of any other breed (though blood-hounds differ considerably), or than in any wild canine species known to me. Accordingly, I applied to Mr. Cupples, well known for his success with this breed, who has weighed and measured many of his own dogs, and who has with great kindness collected for me the following facts from various sources. Fine male dogs, measured at the shoulder, range from 28 inches, which is low, to 33 or even 34 inches in height; and in weight from 80 pounds, which is light, to 120 pounds, or even more. The females range in height from 23 to 27, or even to 28 inches; and in weight from 50 to 70, or even 80 pounds. (38. See also Richardson’s ‘Manual on the Dog,’ p. 59. Much valuable information on the Scottish deer-hound is given by Mr. McNeill, who first called attention to the inequality in size between the sexes, in Scrope’s ‘Art of Deer- Stalking.’ I hope that Mr. Cupples will keep to his intention of publishing a full account and history of this famous breed.) Mr. Cupples concludes that from 95 to 100 pounds for the male, and 70 for the female, would be a safe average; but there is reason to believe that formerly both sexes attained a greater weight. Mr. Cupples has weighed puppies when a fortnight old; in one litter the average weight of four males exceeded that of two females by six and a half ounces; in another litter the average weight of four males exceeded that of one female by less than one ounce; the same males when three weeks old, exceeded the female by seven and a half ounces, and at the age of six weeks by nearly fourteen ounces. Mr. Wright of Yeldersley House, in a letter to Mr. Cupples, says: “I have taken notes on the sizes and weights of puppies of many litters, and as far as my experience goes, dog-puppies as a rule differ very little from bitches till they arrive at about five or six months old; and then the dogs begin to increase, gaining upon the bitches both in weight and size. At birth, and for several weeks afterwards, a bitch-puppy will occasionally be larger than any of the dogs, but they are invariably beaten by them later.” Mr. McNeill, of Colonsay, concludes that “the males do not attain their full growth till over two years old, though the females attain it sooner.” According to Mr. Cupples’ experience, male dogs go on growing in stature till they are from twelve to eighteen months old, and in weight till from eighteen to twenty-four months old; whilst the females cease increasing in stature at the age of from nine to fourteen or fifteen months, and in weight at the age of from twelve to fifteen months. From these various statements it is clear that the full difference in size between the male and female Scotch deer-hound is not acquired until rather late in life. The males almost exclusively are used for coursing, for, as Mr. McNeill informs me, the females have not sufficient strength and weight to pull down a full-grown deer. From the names used in old legends, it appears, as I hear from Mr. Cupples, that, at a very ancient period, the males were the most celebrated, the females being mentioned only as the mothers of famous dogs. Hence, during many generations, it is the male which has been chiefly tested for strength, size, speed, and courage, and the best will have been bred from. As, however, the males do not attain their full dimensions until rather late in life, they will have tended, in accordance with the law often indicated, to transmit their characters to their male offspring alone; and thus the great inequality in size between the sexes of the Scotch deer-hound may probably be accounted for.
[Fig. 65. Head of Common wild boar, in prime of life (from Brehm).]
The males of some few quadrupeds possess organs or parts developed solely as a means of defence against the attacks of other males. Some kinds of deer use, as we have seen, the upper branches of their horns chiefly or exclusively for defending themselves; and the Oryx antelope, as I am informed by Mr. Bartlett, fences most skilfully with his long, gently curved horns; but these are likewise used as organs of offence. The same observer remarks that rhinoceroses in fighting, parry each other’s sidelong blows with their horns, which clatter loudly together, as do the tusks of boars. Although wild boars fight desperately, they seldom, according to Brehm, receive fatal wounds, as the blows fall on each other’s tusks, or on the layer of gristly skin covering the shoulder, called by the German hunters, the shield; and here we have a part specially modified for defence. With boars in the prime of life (Fig. 65) the tusks in the lower jaw are used for fighting, but they become in old age, as Brehm states, so much curved inwards and upwards over the snout that they can no longer be used in this way. They may, however, still serve, and even more effectively, as a means of defence. In compensation for the loss of the lower tusks as weapons of offence, those in the upper jaw, which always project a little laterally, increase in old age so much in length and curve so much upwards that they can be used for attack. Nevertheless, an old boar is not so dangerous to man as one at the age of six or seven years. (39. Brehm, ‘Thierleben,’ B. ii. ss. 729-732.)
[Fig. 66. Skull of the Babirusa Pig (from Wallace’s ‘Malay Archipelago’).]
In the full-grown male Babirusa pig of Celebes (Fig. 66), the lower tusks are formidable weapons, like those of the European boar in the prime of life, whilst the upper tusks are so long and have their points so much curled inwards, sometimes even touching the forehead, that they are utterly useless as weapons of attack. They more nearly resemble horns than teeth, and are so manifestly useless as teeth that the animal was formerly supposed to rest his head by hooking them on to a branch! Their convex surfaces, however, if the head were held a little laterally, would serve as an excellent guard; and hence, perhaps, it is that in old animals they “are generally broken off, as if by fighting.” (40. See Mr. Wallace’s interesting account of this animal, ‘The Malay Archipelago,’ 1869, vol. i. p. 435.) Here, then, we have the curious case of the upper tusks of the Babirusa regularly assuming during the prime of life a structure which apparently renders them fitted only for defence; whilst in the European boar the lower tusks assume in a less degree and only during old age nearly the same form, and then serve in like manner solely for defence.
[Fig. 67. Head of female Aethopian wart-hog, from ‘Proc. Zool. Soc.’ 1869, shewing the same characters as the male, though on a reduced scale. N.B. When the engraving was first made, I was under the impression that it represented the male.]
In the wart-hog (see Phacochoerus aethiopicus, Fig. 67) the tusks in the upper jaw of the male curve upwards during the prime of life, and from being pointed serve as formidable weapons. The tusks in the lower jaw are sharper than those in the upper, but from their shortness it seems hardly possible that they can be used as weapons of attack. They must, however, greatly strengthen those in the upper jaw, from being ground so as to fit closely against their bases. Neither the upper nor the lower tusks appear to have been specially modified to act as guards, though no doubt they are to a certain extent used for this purpose. But the wart-hog is not destitute of other special means of protection, for it has, on each side of the face, beneath the eyes, a rather stiff, yet flexible, cartilaginous, oblong pad (Fig. 67), which projects two or three inches outwards; and it appeared to Mr. Bartlett and myself, when viewing the living animal, that these pads, when struck from beneath by the tusks of an opponent, would be turned upwards, and would thus admirably protect the somewhat prominent eyes. I may add, on the authority of Mr. Bartlett, that these boars when fighting stand directly face to face.
Lastly, the African river-hog (Potomochoerus penicillatus) has a hard cartilaginous knob on each side of the face beneath the eyes, which answers to the flexible pad of the wart-hog; it has also two bony prominences on the upper jaw above the nostrils. A boar of this species in the Zoological Gardens recently broke into the cage of the wart-hog. They fought all night long, and were found in the morning much exhausted, but not seriously wounded. It is a significant fact, as shewing the purposes of the above- described projections and excrescences, that these were covered with blood, and were scored and abraded in an extraordinary manner.
Although the males of so many members of the pig family are provided with weapons, and as we have just seen with means of defence, these weapons seem to have been acquired within a rather late geological period. Dr. Forsyth Major specifies (41. ‘Atti della Soc. Italiana di Sc. Nat.’ 1873, vol. xv. fasc. iv.) several miocene species, in none of which do the tusks appear to have been largely developed in the males; and Professor Rutimeyer was formerly struck with this same fact.
The mane of the lion forms a good defence against the attacks of rival lions, the one danger to which he is liable; for the males, as Sir A. Smith informs me, engage in terrible battles, and a young lion dares not approach an old one. In 1857 a tiger at Bromwich broke into the cage of a lion and a fearful scene ensued: “the lion’s mane saved his neck and head from being much injured, but the tiger at last succeeded in ripping up his belly, and in a few minutes he was dead.” (42. ‘The Times,’ Nov. 10, 1857. In regard to the Canada lynx, see Audubon and Bachman, ‘Quadrupeds of North America,’ 1846, p. 139.) 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) (43. Dr. Murie, on Otaria, ‘Proc. Zoolog. Soc.’ 1869, p. 109. Mr. J.A. Allen, in the paper above quoted (p. 75), doubts whether the hair, which is longer on the neck in the male than in the female, deserves to be called a mane.) have great manes, whilst the females have small ones or none. The male baboon of the Cape of Good Hope (Cynocephalus porcarius) has a much longer mane and larger canine teeth than the female; and the mane probably serves as a protection, for, on asking the keepers in the Zoological Gardens, without giving them any clue to my object, whether any of the monkeys especially attacked each other by the nape of the neck, I was answered that this was not the case, except with the above baboon. In the Hamadryas baboon, Ehrenberg compares the mane of the adult male to that of a young lion, whilst in the young of both sexes and in the female the mane is almost absent.
It appeared to me probable that the immense woolly mane of the male American bison, which reaches almost to the ground, and is much more developed in the males than in the females, served as a protection to them in their terrible battles; but an experienced hunter told Judge Caton that he had never observed anything which favoured this belief. The stallion has a thicker and fuller mane than the mare; and I have made particular inquiries of two great trainers and breeders, who have had charge of many entire horses, and am assured that they “invariably endeavour to seize one another by the neck.” It does not, however, follow from the foregoing statements, that when the hair on the neck serves as a defence, that it was originally developed for this purpose, though this is probable in some cases, as in that of the lion. I am informed by Mr. McNeill that the long hairs on the throat of the stag (Cervus elaphus) serve as a great protection to him when hunted, for the dogs generally endeavour to seize him by the throat; but it is not probable that these hairs were specially developed for this purpose; otherwise the young and the females would have been equally protected.
CHOICE IN PAIRING BY EITHER SEX OF QUADRUPEDS.
Before describing in the next chapter, the differences between the sexes in voice, odours emitted, and ornaments, it will be convenient here to consider whether the sexes exert any choice in their unions. Does the female prefer any particular male, either before or after the males may have fought together for supremacy; or does the male, when not a polygamist, select any particular female? The general impression amongst breeders seems to be that the male accepts any female; and this owing to his eagerness, is, in most cases, probably the truth. Whether the female as a general rule indifferently accepts any male is much more doubtful. In the fourteenth chapter, on Birds, a considerable body of direct and indirect evidence was advanced, shewing that the female selects her partner; and it would be a strange anomaly if female quadrupeds, which stand higher in the scale and have higher mental powers, did not generally, or at least often, exert some choice. The female could in most cases escape, if wooed by a male that did not please or excite her; and when pursued by several males, as commonly occurs, she would often have the opportunity, whilst they were fighting together, of escaping with some one male, or at least of temporarily pairing with him. This latter contingency has often been observed in Scotland with female red-deer, as I am informed by Sir Philip Egerton and others. (44. Mr. Boner, in his excellent description of the habits of the red-deer in Germany (‘Forest Creatures,’ 1861, p. 81) says, “while the stag is defending his rights against one intruder, another invades the sanctuary of his harem, and carries off trophy after trophy.” Exactly the same thing occurs with seals; see Mr. J.A. Allen, ibid. p. 100.)
It is scarcely possible that much should be known about female quadrupeds in a state of nature making any choice in their marriage unions. The following curious details on the courtship of one of the eared seals (Callorhinus ursinus) are given (45. Mr. J.A. Allen in ‘Bull. Mus. Comp. Zoolog. of Cambridge, United States,’ vol. ii. No. 1, p. 99.) 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 a large size. (46. ‘Dogs: their Management,’ by E. Mayhew, M.R.C.V.S., 2nd ed., 1864, pp. 187-192.) The well-known veterinary Blaine states (47. Quoted by Alex. Walker, ‘On Intermarriage,’ 1838, p. 276; see also p. 244.) that his own female pug dog became so attached to a spaniel, and a female setter to a cur, that in neither case would they pair with a dog of their own breed until several weeks had elapsed. Two similar and trustworthy accounts have been given me in regard to a female retriever and a spaniel, both of which became enamoured with terrier-dogs.
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 in 1868, a female deerhound in his kennel thrice produced puppies, and on each occasion shewed a marked preference for one of the largest and handsomest, but not the most eager, of four deerhounds living with her, all in the prime of life. Mr. Cupples has observed that the female generally favours a dog whom she has associated with and knows; her shyness and timidity at first incline her against a strange dog. The male, on the contrary, seems rather inclined towards strange females. It appears to be rare when the male refuses any particular female, but Mr. Wright, of Yeldersley House, a great breeder of dogs, informs me that he has known some instances; he cites the case of one of his own deerhounds, who would not take any notice of a particular female mastiff, so that another deerhound had to be employed. It would be superfluous to give, as I could, other instances, and I will only add that Mr. Barr, who has carefully bred many bloodhounds, states that in almost every instance particular individuals of opposite sexes shew a decided preference for each other. Finally, Mr. Cupples, after attending to this subject for another year, has written to me, “I have had full confirmation of my former statement, that dogs in breeding form decided preferences for each other, being often influenced by size, bright colour, and individual characters, as well as by the degree of their previous familiarity.”
In regard to horses, Mr. Blenkiron, the greatest breeder of race-horses in the world, informs me that stallions are so frequently capricious in their choice, rejecting one mare and without any apparent cause taking to another, that various artifices have to be habitually used. The famous Monarque, for instance, would never consciously look at the dam of Gladiateur, and a trick had to be practised. We can partly see the reason why valuable race-horse stallions, which are in such demand as to be exhausted, should be so particular in their choice. Mr. Blenkiron has never known a mare reject a horse; but this has occurred in Mr. Wright’s stable, so that the mare had to be cheated. Prosper Lucas (48. ‘Traite de l’Hered. Nat.’ tom. ii. 1850, p. 296.) quotes various statements from French authorities, and remarks, “On voit des etalons qui s’eprennent d’une jument, et negligent toutes les autres.” He gives, on the authority of Baelen, similar facts in regard to bulls; and Mr. H. Reeks assures me that a famous short-horn bull belonging to his father “invariably refused to be matched with a black cow.” Hoffberg, in describing the domesticated reindeer of Lapland says, “Foeminae majores et fortiores mares prae caeteris admittunt, ad eos confugiunt, a junioribus agitatae, qui hos in fugam conjiciunt.” (49. ‘Amoenitates Acad.’ vol. iv. 1788, p. 160.) A clergyman, who has bred many pigs, asserts that sows often reject one boar and immediately accept another.
From these facts there can be no doubt that, with most of our domesticated quadrupeds, strong individual antipathies and preferences are frequently exhibited, and much more commonly by the female than by the male. This being the case, it is improbable that the unions of quadrupeds in a state of nature should be left to mere chance. It is much more probable that the females are allured or excited by particular males, who possess certain characters in a higher degree than other males; but what these characters are, we can seldom or never discover with certainty.
CHAPTER XVIII.
SECONDARY SEXUAL CHARACTERS OF MAMMALS–continued.
Voice–Remarkable sexual peculiarities in seals–Odour–Development of the hair–Colour of the hair and skin–Anomalous case of the female being more ornamented than the male–Colour and ornaments due to sexual selection– Colour acquired for the sake of protection–Colour, though common to both sexes, often due to sexual selection–On the disappearance of spots and stripes in adult quadrupeds–On the colours and ornaments of the Quadrumana–Summary.
Quadrupeds use their voices for various purposes, as a signal of danger, as a call from one member of a troop to another, or from the mother to her lost offspring, or from the latter for protection to their mother; but such uses need not here be considered. We are concerned only with the difference between the voices of the sexes, for instance between that of the lion and lioness, or of the bull and cow. Almost all male animals use their voices much more during the rutting-season than at any other time; and some, as the giraffe and porcupine (1. Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 585.), are said to be completely mute excepting at this season. As the throats (i.e. the larynx and thyroid bodies (2. Ibid. p. 595.)) of stags periodically become enlarged at the beginning of the breeding-season, it might be thought that their powerful voices must be somehow of high importance to them; but this is very doubtful. From information given to me by two experienced observers, Mr. McNeill and Sir P. Egerton, it seems that young stags under three years old do not roar or bellow; and that the old ones begin bellowing at the commencement of the breeding-season, at first only occasionally and moderately, whilst they restlessly wander about in search of the females. Their battles are prefaced by loud and prolonged bellowing, but during the actual conflict they are silent. Animals of all kinds which habitually use their voices utter various noises under any strong emotion, as when enraged and preparing to fight; but this may merely be the result of nervous excitement, which leads to the spasmodic contraction of almost all the muscles of the body, as when a man grinds his teeth and clenches his fists in rage or agony. No doubt stags challenge each other to mortal combat by bellowing; but those with the more powerful voices, unless at the same time the stronger, better-armed, and more courageous, would not gain any advantage over their rivals.
It is possible that the roaring of the lion may be of some service to him by striking terror into his adversary; for when enraged he likewise erects his mane and thus instinctively tries to make himself appear as terrible as possible. But it can hardly be supposed that the bellowing of the stag, even if it be of service to him in this way, can have been important enough to have led to the periodical enlargement of the throat. Some writers suggest that the bellowing serves as a call to the female; but the experienced observers above quoted inform me that female deer do not search for the male, though the males search eagerly for the females, as indeed might be expected from what we know of the habits of other male quadrupeds. The voice of the female, on the other hand, quickly brings to her one or more stags (3. See, for instance, Major W. Ross King (‘The Sportsman in Canada,’ 1866, pp. 53, 131) on the habits of the moose and wild reindeer.), as is well known to the hunters who in wild countries imitate her cry. If we could believe that the male had the power to excite or allure the female by his voice, the periodical enlargement of his vocal organs would be intelligible on the principle of sexual selection, together with inheritance limited to the same sex and season; but we have no evidence in favour of this view. As the case stands, the loud voice of the stag during the breeding-season does not seem to be of any special service to him, either during his courtship or battles, or in any other way. But may we not believe that the frequent use of the voice, under the strong excitement of love, jealousy, and rage, continued during many generations, may at last have produced an inherited effect on the vocal organs of the stag, as well as of other male animals? This appears to me, in our present state of knowledge, the most probable view.
The voice of the adult male gorilla is tremendous, and he is furnished with a laryngeal sack, as is the adult male orang. (4. Owen ‘Anatomy of Vertebrates,’ vol. iii. p. 600.) The gibbons rank among the noisiest of monkeys, and the Sumatra species (Hylobates syndactylus) is also furnished with an air sack; but Mr. Blyth, who has had opportunities for observation, does not believe that the male is noisier than the female. Hence, these latter monkeys probably use their voices as a mutual call; and this is certainly the case with some quadrupeds, for instance the beaver. (5. Mr. Green, in ‘Journal of Linnean Society,’ vol. x. ‘Zoology,’ 1869, note 362.) Another gibbon, the H. agilis, is remarkable, from having the power of giving a complete and correct octave of musical notes (6. C.L. Martin, ‘General Introduction to the Natural History of Mamm. Animals,’ 1841, p. 431.), which we may reasonably suspect serves as a sexual charm; but I shall have to recur to this subject in the next chapter. The vocal organs of the American Mycetes caraya are one-third larger in the male than in the female, and are wonderfully powerful. These monkeys in warm weather make the forests resound at morning and evening with their overwhelming voices. The males begin the dreadful concert, and often continue it during many hours, the females sometimes joining in with their less powerful voices. An excellent observer, Rengger (7. ‘Naturgeschichte der Saugethiere von Paraguay,’ 1830, ss. 15, 21.), could not perceive that they were excited to begin by any special cause; he thinks that, like many birds, they delight in their own music, and try to excel each other. Whether most of the foregoing monkeys have acquired their powerful voices in order to beat their rivals and charm the females–or whether the vocal organs have been strengthened and enlarged through the inherited effects of long-continued use without any particular good being thus gained–I will not pretend to say; but the former view, at least in the case of the Hylobates agilis, seems the most probable.
I may here mention two very curious sexual peculiarities occurring in seals, because they have been supposed by some writers to affect the voice. The nose of the male sea-elephant (Macrorhinus proboscideus) becomes greatly elongated during the breeding-season, and can then be erected. In this state it is sometimes a foot in length. The female is not thus provided at any period of life. The male makes a wild, hoarse, gurgling noise, which is audible at a great distance and is believed to be strengthened by the proboscis; the voice of the female being different. Lesson compares the erection of the proboscis, with the swelling of the wattles of male gallinaceous birds whilst courting the females. In another allied kind of seal, the bladder-nose (Cystophora cristata), the head is covered by a great hood or bladder. This is supported by the septum of the nose, which is produced far backwards and rises into an internal crest seven inches in height. The hood is clothed with short hair, and is muscular; can be inflated until it more than equals the whole head in size! The males when rutting, fight furiously on the ice, and their roaring “is said to be sometimes so loud as to be heard four miles off.” When attacked they likewise roar or bellow; and whenever irritated the bladder is inflated and quivers. Some naturalists believe that the voice is thus strengthened, but various other uses have been assigned to this extraordinary structure. Mr. R. Brown thinks that it serves as a protection against accidents of all kinds; but this is not probable, for, as I am assured by Mr. Lamont who killed 600 of these animals, the hood is rudimentary in the females, and it is not developed in the males during youth. (8. On the sea-elephant, see an article by Lesson, in ‘Dict. Class. Hist. Nat.’ tom. xiii. p. 418. For the Cystophora, or Stemmatopus, see Dr. Dekay, ‘Annals of Lyceum of Nat. Hist.’ New York, vol. i. 1824, p. 94. Pennant has also collected information from the sealers on this animal. The fullest account is given by Mr. Brown, in ‘Proc. Zoolog. Soc.’ 1868, p. 435.)
ODOUR.
With some animals, as with the notorious skunk of America, the overwhelming odour which they emit appears to serve exclusively as a defence. With shrew-mice (Sorex) both sexes possess abdominal scent-glands, and there can be little doubt, from the rejection of their bodies by birds and beasts of prey, that the odour is protective; nevertheless, the glands become enlarged in the males during the breeding-season. In many other quadrupeds the glands are of the same size in both sexes (9. As with the castoreum of the beaver, see Mr. L.H. Morgan’s most interesting work, ‘The American Beaver,’ 1868, p. 300. Pallas (‘Spic. Zoolog.’ fasc. viii. 1779, p. 23) has well discussed the odoriferous glands of mammals. Owen (‘Anat. of Vertebrates,’ vol. iii. p. 634) also gives an account of these glands, including those of the elephant, and (p. 763) those of shrew-mice. On bats, Mr. Dobson in ‘Proceedings of the Zoological Society’ 1873, p. 241.), but their uses are not known. In other species the glands are confined to the males, or are more developed than in the females; and they almost always become more active during the rutting-season. At this period the glands on the sides of the face of the male elephant enlarge, and emit a secretion having a strong musky odour. The males, and rarely the females, of many kinds of bats have glands and protrudable sacks situated in various parts; and it is believed that these are odoriferous.
The rank effluvium of the male goat is well known, and that of certain male deer is wonderfully strong and persistent. On the banks of the Plata I perceived the air tainted with the odour of the male Cervus campestris, at half a mile to leeward of a herd; and a silk handkerchief, in which I carried home a skin, though often used and washed, retained, when first unfolded, traces of the odour for one year and seven months. This animal does not emit its strong odour until more than a year old, and if castrated whilst young never emits it. (10. Rengger, ‘Naturgeschichte der Saugethiere von Paraguay,’ 1830, s. 355. This observer also gives some curious particulars in regard to the odour.) Besides the general odour, permeating the whole body of certain ruminants (for instance, Bos moschatus) in the breeding-season, many deer, antelopes, sheep, and goats possess odoriferous glands in various situations, more especially on their faces. The so-called tear-sacks, or suborbital pits, come under this head. These glands secrete a semi-fluid fetid matter which is sometimes so copious as to stain the whole face, as I have myself seen in an antelope. They are “usually larger in the male than in the female, and their development is checked by castration.” (11. Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 632. See also Dr. Murie’s observations on those glands in the ‘Proc. Zoolog. Soc.’ 1870, p. 340. Desmarest, ‘On the Antilope subgutturosa, ‘Mammalogie,’ 1820, p. 455.) According to Desmarest they are altogether absent in the female of Antilope subgutturosa. Hence, there can be no doubt that they stand in close relation with the reproductive functions. They are also sometimes present, and sometimes absent, in nearly allied forms. In the adult male musk-deer (Moschus moschiferus), a naked space round the tail is bedewed with an odoriferous fluid, whilst in the adult female, and in the male until two years old, this space is covered with hair and is not odoriferous. The proper musk- sack of this deer is from its position necessarily confined to the male, and forms an additional scent-organ. It is a singular fact that the matter secreted by this latter gland, does not, according to Pallas, change in consistence, or increase in quantity, during the rutting-season; nevertheless this naturalist admits that its presence is in some way connected with the act of reproduction. He gives, however, only a conjectural and unsatisfactory explanation of its use. (12. Pallas, ‘Spicilegia Zoolog.’ fasc. xiii. 1799, p. 24; Desmoulins, ‘Dict. Class. d’Hist. Nat.’ tom. iii. p. 586.)
In most cases, when only the male emits a strong odour during the breeding- season, it probably serves to excite or allure the female. We must not judge on this head by our own taste, for it is well known that rats are enticed by certain essential oils, and cats by valerian, substances far from agreeable to us; and that dogs, though they will not eat carrion, sniff and roll on it. From the reasons given when discussing the voice of the stag, we may reject the idea that the odour serves to bring the females from a distance to the males. Active and long-continued use cannot here have come into play, as in the case of the vocal organs. The odour emitted must be of considerable importance to the male, inasmuch as large and complex glands, furnished with muscles for everting the sack, and for closing or opening the orifice, have in some cases been developed. The development of these organs is intelligible through sexual selection, if the most odoriferous males are the most successful in winning the females, and in leaving offspring to inherit their gradually perfected glands and odours.
DEVELOPMENT OF THE HAIR.
We have seen that male quadrupeds often have the hair on their necks and shoulders much more developed than the females; and many additional instances could be given. This sometimes serves as a defence to the male during his battles; but whether the hair in most cases has been specially developed for this purpose, is very doubtful. We may feel almost certain that this is not the case, when only a thin and narrow crest runs along the back; for a crest of this kind would afford scarcely any protection, and the ridge of the back is not a place likely to be injured; nevertheless such crests are sometimes confined to the males, or are much more developed in them than in the females. Two antelopes, the Tragelaphus scriptus (13. Dr. Gray, ‘Gleanings from the Menagerie at Knowsley,’ pl. 28.) (Fig. 70) and Portax picta may be given as instances. When stags, and the males of the wild goat, are enraged or terrified, these crests stand erect (14. Judge Caton on the Wapiti, ‘Transact. Ottawa Acad. Nat. Sciences,’ 1868, pp. 36, 40; Blyth, ‘Land and Water,’ on Capra aegagrus 1867, p. 37.); but it cannot be supposed that they have been developed merely for the sake of exciting fear in their enemies. One of the above-named antelopes, the Portax picta, has a large well-defined brush of black hair on the throat, and this is much larger in the male than in the female. In the Ammotragus tragelaphus of North Africa, a member of the sheep-family, the fore-legs are almost concealed by an extraordinary growth of hair, which depends from the neck and upper halves of the legs; but Mr. Bartlett does not believe that this mantle is of the least use to the male, in whom it is much more developed than in the female.
[Fig. 68. Pithecia satanas, male (from Brehm).]
Male quadrupeds of many kinds differ from the females in having more hair, or hair of a different character, on certain parts of their faces. Thus the bull alone has curled hair on the forehead. (15. Hunter’s ‘Essays and Observations,’ edited by Owen, 1861. vol. i. p. 236.) In three closely- allied sub-genera of the goat family, only the males possess beards, sometimes of large size; in two other sub-genera both sexes have a beard, but it disappears in some of the domestic breeds of the common goat; and neither sex of the Hemitragus has a beard. In the ibex the beard is not developed during the summer, and is so small at other times that it may be called rudimentary. (16. See Dr. Gray’s ‘Catalogue of Mammalia in the British Museum,’ part iii. 1852, p. 144.) With some monkeys the beard is confined to the male, as in the orang; or is much larger in the male than in the female, as in the Mycetes caraya and Pithecia satanas (Fig. 68). So it is with the whiskers of some species of Macacus (17. Rengger, ‘Saugthiere,’ etc., s. 14; Desmarest, ‘Mammalogie,’ p. 86.), and, as we have seen, with the manes of some species of baboons. But with most kinds of monkeys the various tufts of hair about the face and head are alike in both sexes.
The males of various members of the ox family (Bovidae), and of certain antelopes, are furnished with a dewlap, or great fold of skin on the neck, which is much less developed in the female.
Now, what must we conclude with respect to such sexual differences as these? No one will pretend that the beards of certain male goats, or the dewlaps of the bull, or the crests of hair along the backs of certain male antelopes, are of any use to them in their ordinary habits. It is possible that the immense beard of the male Pithecia, and the large beard of the male orang, may protect their throats when fighting; for the keepers in the Zoological Gardens inform me that many monkeys attack each other by the throat; but it is not probable that the beard has been developed for a distinct purpose from that served by the whiskers, moustache, and other tufts of hair on the face; and no one will suppose that these are useful as a protection. Must we attribute all these appendages of hair or skin to mere purposeless variability in the male? It cannot be denied that this is possible; for in many domesticated quadrupeds, certain characters, apparently not derived through reversion from any wild parent form, are confined to the males, or are more developed in them than in the females– for instance, the hump on the male zebu-cattle of India, the tail of fat- tailed rams, the arched outline of the forehead in the males of several breeds of sheep, and lastly, the mane, the long hairs on the hind legs, and the dewlap of the male of the Berbura goat. (18. See the chapters on these several animals in vol. i. of my ‘Variation of Animals under Domestication;’ also vol. ii. p. 73; also chap. xx. on the practice of selection by semi-civilised people. For the Berbura goat, see Dr. Gray, ‘Catalogue,’ ibid. p. 157.) The mane, which occurs only in the rams of an African breed of sheep, is a true secondary sexual character, for, as I hear from Mr. Winwood Reade, it is not developed if the animal be castrated. Although we ought to be extremely cautious, as shewn in my work on ‘Variation under Domestication,’ in concluding that any character, even with animals kept by semi-civilised people, has not been subjected to selection by man, and thus augmented, yet in the cases just specified this is improbable; more especially as the characters are confined to the males, or are more strongly developed in them than in the females. If it were positively known that the above African ram is a descendant of the same primitive stock as the other breeds of sheep, and if the Berbura male-goat with his mane, dewlap, etc., is descended from the same stock as other goats, then, assuming that selection has not been applied to these characters, they must be due to simple variability, together with sexually- limited inheritance.
Hence it appears reasonable to extend this same view to all analogous cases with animals in a state of nature. Nevertheless I cannot persuade myself that it generally holds good, as in the case of the extraordinary development of hair on the throat and fore-legs of the male Ammotragus, or in that of the immense beard of the male Pithecia. Such study as I have been able to give to nature makes me believe that parts or organs which are highly developed, were acquired at some period for a special purpose. With those antelopes in which the adult male is more strongly-coloured than the female, and with those monkeys in which the hair on the face is elegantly arranged and coloured in a diversified manner, it seems probable that the crests and tufts of hair were gained as ornaments; and this I know is the opinion of some naturalists. If this be correct, there can be little doubt that they were gained or at least modified through sexual selection; but how far the same view may be extended to other mammals is doubtful.
COLOUR OF THE HAIR AND OF THE NAKED SKIN.
I will first give briefly all the cases known to me of male quadrupeds differing in colour from the females. With Marsupials, as I am informed by Mr. Gould, the sexes rarely differ in this respect; but the great red kangaroo offers a striking exception, “delicate blue being the prevailing tint in those parts of the female which in the male are red.” (19. Osphranter rufus, Gould, ‘Mammals of Australia,’ 1863, vol. ii. On the Didelphis, Desmarest, ‘Mammalogie,’ p. 256.) In the Didelphis opossum of Cayenne the female is said to be a little more red than the male. Of the Rodents, Dr. Gray remarks: “African squirrels, especially those found in the tropical regions, have the fur much brighter and more vivid at some seasons of the year than at others, and the fur of the male is generally brighter than that of the female.” (20. ‘Annals and Magazine of Natural History,’ Nov. 1867, p. 325. On the Mus minutus, Desmarest, ‘Mammalogie,’ p. 304.) Dr. Gray informs me that he specified the African squirrels, because, from their unusually bright colours, they best exhibit this difference. The female of the Mus minutus of Russia is of a paler and dirtier tint than the male. In a large number of bats the fur of the male is lighter than in the female. (21. J.A. Allen, in ‘Bulletin of Mus. Comp. Zoolog. of Cambridge, United States,’ 1869, p. 207. Mr. Dobson on sexual characters in the Chiroptera, ‘Proceedings of the Zoological Society,’ 1873, p. 241. Dr. Gray on Sloths, ibid. 1871, p. 436.) Mr. Dobson also remarks, with respect to these animals: “Differences, depending partly or entirely on the possession by the male of fur of a much more brilliant hue, or distinguished by different markings or by the greater length of certain portions, are met only, to any appreciable extent, in the frugivorous bats in which the sense of sight is well developed.” This last remark deserves attention, as bearing on the question whether bright colours are serviceable to male animals from being ornamental. In one genus of sloths, it is now established, as Dr. Gray states, “that the males are ornamented differently from the females–that is to say, that they have a patch of soft short hair between the shoulders, which is generally of a more or less orange colour, and in one species pure white. The females, on the contrary, are destitute of this mark.”
The terrestrial Carnivora and Insectivora rarely exhibit sexual differences of any kind, including colour. The ocelot (Felis pardalis), however, is exceptional, for the colours of the female, compared with those of the male, are “moins apparentes, le fauve, etant plus terne, le blanc moins pur, les raies ayant moins de largeur et les taches moins de diametre.” (22. Desmarest, ‘Mammalogie,’ 1820, p. 220. On Felis mitis, Rengger, ibid. s. 194.) The sexes of the allied Felis mitis also differ, but in a less degree; the general hues of the female being rather paler than in the male, with the spots less black. The marine Carnivora or seals, on the other hand, sometimes differ considerably in colour, and they present, as we have already seen, other remarkable sexual differences. Thus the male of the Otaria nigrescens of the southern hemisphere is of a rich brown shade above; whilst the female, who acquires her adult tints earlier in life than the male, is dark-grey above, the young of both sexes being of a deep chocolate colour. The male of the northern Phoca groenlandica is tawny grey, with a curious saddle-shaped dark mark on the back; the female is much smaller, and has a very different appearance, being “dull white or yellowish straw-colour, with a tawny hue on the back”; the young at first are pure white, and can “hardly be distinguished among the icy hummocks and snow, their colour thus acting as a protection.” (23. Dr. Murie on the Otaria, ‘Proceedings Zoological Society,’ 1869, p. 108. Mr. R. Brown on the P. groenlandica, ibid. 1868, p. 417. See also on the colours of seals, Desmarest, ibid. pp. 243, 249.)
With Ruminants sexual differences of colour occur more commonly than in any other order. A difference of this kind is general in the Strepsicerene antelopes; thus the male nilghau (Portax picta) is bluish-grey and much darker than the female, with the square white patch on the throat, the white marks on the fetlocks, and the black spots on the ears all much more distinct. We have seen that in this species the crests and tufts of hair are likewise more developed in the male than in the hornless female. I am informed by Mr. Blyth that the male, without shedding his hair, periodically becomes darker during the breeding-season. Young males cannot be distinguished from young females until about twelve months old; and if the male is emasculated before this period, he never, according to the same authority, changes colour. The importance of this latter fact, as evidence that the colouring of the Portax is of sexual origin, becomes obvious, when we hear (24. Judge Caton, in ‘Transactions of the Ottawa Academy of Natural Sciences,’ 1868, p. 4.) that neither the red summer-coat nor the blue winter-coat of the Virginian deer is at all affected by emasculation. With most or all of the highly-ornamented species of Tragelaphus the males are darker than the hornless females, and their crests of hair are more fully developed. In the male of that magnificent antelope, the Derbyan eland, the body is redder, the whole neck much blacker, and the white band which separates these colours broader than in the female. In the Cape eland, also, the male is slightly darker than the female. (25. Dr. Gray, ‘Cat. of Mamm. in Brit. Mus.’ part iii. 1852, pp. 134-142; also Dr. Gray, ‘Gleanings from the Menagerie of Knowsley,’ in which there is a splendid drawing of the Oreas derbianus: see the text on Tragelaphus. For the Cape eland (Oreas canna), see Andrew Smith, ‘Zoology of S. Africa,’ pl. 41 and 42. There are also many of these Antelopes in the Zoological Gardens.)
In the Indian black-buck (A. bezoartica), which belongs to another tribe of antelopes, the male is very dark, almost black; whilst the hornless female is fawn-coloured. We meet in this species, as Mr. Blyth informs me, with an exactly similar series of facts, as in the Portax picta, namely, in the male periodically changing colour during the breeding-season, in the effects of emasculation on this change, and in the young of both sexes being indistinguishable from each other. In the Antilope niger the male is black, the female, as well as the young of both sexes, being brown; in A. sing-sing the male is much brighter coloured than the hornless female, and his chest and belly are blacker; in the male A. caama, the marks and lines which occur on various parts of the body are black, instead of brown as in the female; in the brindled gnu (A. gorgon) “the colours of the male are nearly the same as those of the female, only deeper and of a brighter hue.” (26. On the Ant. niger, see ‘Proc. Zool. Soc.’ 1850, p. 133. With respect to an allied species, in which there is an equal sexual difference in colour, see Sir S. Baker, ‘The Albert Nyanza,’ 1866, vol. ii. p. 627. For the A. sing-sing, Gray, ‘Cat. B. Mus.’ p. 100. Desmarest, ‘Mammalogie,’ p. 468, on the A. caama. Andrew Smith, ‘Zoology of S. Africa,’ on the Gnu.) Other analogous cases could be added.
The Banteng bull (Bos sondaicus) of the Malayan Archipelago is almost black, with white legs and buttocks; the cow is of a bright dun, as are the young males until about the age of three years, when they rapidly change colour. The emasculated bull reverts to the colour of the female. The female Kemas goat is paler, and both it and the female Capra aegagrus are said to be more uniformly tinted than their males. Deer rarely present any sexual differences in colour. Judge Caton, however, informs me that in the males of the wapiti deer (Cervus canadensis) the neck, belly, and legs are much darker than in the female; but during the winter the darker tints gradually fade away and disappear. I may here mention that Judge Caton has in his park three races of the Virginian deer, which differ slightly in colour, but the differences are almost exclusively confined to the blue winter or breeding-coat; so that this case may be compared with those given in a previous chapter of closely-allied or representative species of birds, which differ from each other only in their breeding plumage. (27. ‘Ottawa Academy of Sciences,’ May 21, 1868, pp. 3, 5.) The females of Cervus paludosus of S. America, as well as the young of both sexes, do not possess the black stripes on the nose and the blackish-brown line on the breast, which are characteristic of the adult males. (28. S. Muller, on the Banteng, ‘Zoog. Indischen Archipel.’ 1839-1844, tab. 35; see also Raffles, as quoted by Mr. Blyth, in ‘Land and Water,’ 1867, p. 476. On goats, Dr. Gray, ‘Catalogue of the British Museum,’ p. 146; Desmarest, ‘Mammalogie,’ p. 482. On the Cervus paludosus, Rengger, ibid. s. 345.) Lastly, as I am informed by Mr. Blyth, the mature male of the beautifully coloured and spotted axis deer is considerably darker than the female: and this hue the castrated male never acquires.
The last Order which we need consider is that of the Primates. The male of the Lemur macaco is generally coal-black, whilst the female is brown. (29. Sclater, ‘Proc. Zool. Soc.’ 1866, p. i. The same fact has also been fully ascertained by MM. Pollen and van Dam. See, also, Dr. Gray in ‘Annals and Magazine of Natural History,’ May 1871, p. 340.) Of the Quadrumana of the New World, the females and young of Mycetes caraya are greyish-yellow and like each other; in the second year the young male becomes reddish-brown; in the third, black, excepting the stomach, which, however, becomes quite black in the fourth or fifth year. There is also a strongly-marked difference in colour between the sexes of Mycetes seniculus and Cebus capucinus; the young of the former, and I believe of the latter species, resembling the females. With Pithecia leucocephala the young likewise resemble the females, which are brownish-black above and light rusty-red beneath, the adult males being black. The ruff of hair round the face of Ateles marginatus is tinted yellow in the male and white in the female. Turning to the Old World, the males of Hylobates hoolock are always black, with the exception of a white band over the brows; the females vary from whity-brown to a dark tint mixed with black, but are never wholly black. (30. On Mycetes, Rengger, ibid. s. 14; and Brehm, ‘Thierleben,’ B. i. s. 96, 107. On Ateles Desmarest, ‘Mammalogie,’ p. 75. On Hylobates, Blyth, ‘Land and Water,’ 1867, p. 135. On the Semnopithecus, S. Muller, ‘Zoog. Indischen Archipel.’ tab. x.) In the beautiful Cercopithecus diana, the head of the adult male is of an intense black, whilst that of the female is dark grey; in the former the fur between the thighs is of an elegant fawn- colour, in the latter it is paler. In the beautiful and curious moustache monkey (Cercopithecus cephus) the only difference between the sexes is that the tail of the male is chestnut and that of the female grey; but Mr. Bartlett informs me that all the hues become more pronounced in the male when adult, whilst in the female they remain as they were during youth. According to the coloured figures given by Solomon Muller, the male of Semnopithecus chrysomelas is nearly black, the female being pale brown. In the Cercopithecus cynosurus and griseo-viridis one part of the body, which is confined to the male sex, is of the most brilliant blue or green, and contrasts strikingly with the naked skin on the hinder part of the body, which is vivid red.
[Fig. 69. Head of male Mandrill (from Gervais, ‘Hist. Nat. des Mammiferes’).]
Lastly, in the baboon family, the adult male of Cynocephalus hamadryas differs from the female not only by his immense mane, but slightly in the colour of the hair and of the naked callosities. In the drill (C. leucophaeus) the females and young are much paler-coloured, with less green, than the adult males. No other member in the whole class of mammals is coloured in so extraordinary a manner as the adult male mandrill (C. mormon). The face at this age becomes of a fine blue, with the ridge and tip of the nose of the most brilliant red. According to some authors, the face is also marked with whitish stripes, and is shaded in parts with black, but the colours appear to be variable. On the forehead there is a crest of hair, and on the chin a yellow beard. “Toutes les parties superieures de leurs cuisses et le grand espace nu de leurs fesses sont egalement colores du rouge le plus vif, avec un melange de bleu qui ne manque reellement pas d’elegance.” (31. Gervais, ‘Hist. Nat. des Mammiferes,’ 1854, p. 103. Figures are given of the skull of the male. Also Desmarest, ‘Mammalogie,’ p. 70. Geoffroy St.-Hilaire and F. Cuvier, ‘Hist. Nat. des Mammiferes,’ 1824, tom. i.) When the animal is excited all the naked parts become much more vividly tinted. Several authors have used the strongest expressions in describing these resplendent colours, which they compare with those of the most brilliant birds. Another remarkable peculiarity is that when the great canine teeth are fully developed, immense protuberances of bone are formed on each cheek, which are deeply furrowed longitudinally, and the naked skin over them is brilliantly- coloured, as just-described. (Fig. 69.) In the adult females and in the young of both sexes these protuberances are scarcely perceptible; and the naked parts are much less bright coloured, the face being almost black, tinged with blue. In the adult female, however, the nose at certain regular intervals of time becomes tinted with red.
In all the cases hitherto given the male is more strongly or brighter coloured than the female, and differs from the young of both sexes. But as with some few birds it is the female which is brighter coloured than the male, so with the Rhesus monkey (Macacus rhesus), the female has a large surface of naked skin round the tail, of a brilliant carmine red, which, as I was assured by the keepers in the Zoological Gardens, periodically becomes even yet more vivid, and her face also is pale red. On the other hand, in the adult male and in the young of both sexes (as I saw in the Gardens), neither the naked skin at the posterior end of the body, nor the face, shew a trace of red. It appears, however, from some published accounts, that the male does occasionally, or during certain seasons, exhibit some traces of the red. Although he is thus less ornamented than the female, yet in the larger size of his body larger canine teeth, more developed whiskers, more prominent superciliary ridges, he follows the common rule of the male excelling the female.
I have now given all the cases known to me of a difference in colour between the sexes of mammals. Some of these may be the result of variations confined to one sex and transmitted to the same sex, without any good being gained, and therefore without the aid of selection. We have instances of this with our domesticated animals, as in the males of certain cats being rusty-red, whilst the females are tortoise-shell coloured. Analogous cases occur in nature: Mr. Bartlett has seen many black varieties of the jaguar, leopard, vulpine phalanger, and wombat; and he is certain that all, or nearly all these animals, were males. On the other hand, with wolves, foxes, and apparently American squirrels, both sexes are occasionally born black. Hence it is quite possible that with some mammals a difference in colour between the sexes, especially when this is congenital, may simply be the result, without the aid of selection, of the occurrence of one or more variations, which from the first were sexually limited in their transmission. Nevertheless it is improbable that the diversified, vivid, and contrasted colours of certain quadrupeds, for instance, of the above monkeys and antelopes, can thus be accounted for. We should bear in mind that these colours do not appear in the male at birth, but only at or near maturity; and that unlike ordinary variations, they are lost if the male be emasculated. It is on the whole probable that the strongly-marked colours and other ornamental characters of male quadrupeds are beneficial to them in their rivalry with other males, and have consequently been acquired through sexual selection. This view is strengthened by the differences in colour between the sexes occurring almost exclusively, as may be collected from the previous details, in those groups and sub-groups of mammals which present other and strongly-marked secondary sexual characters; these being likewise due to sexual selection.
Quadrupeds manifestly take notice of colour. Sir S. Baker repeatedly observed that the African elephant and rhinoceros attacked white or grey horses with special fury. I have elsewhere shewn (32. The ‘Variation of Animals and Plants under Domestication,’ 1868, vol. ii. pp. 102, 103.) that half-wild horses apparently prefer to pair with those of the same colour, and that herds of fallow-deer of different colours, though living together, have long kept distinct. It is a more significant fact that a female zebra would not admit the addresses of a male ass until he was painted so as to resemble a zebra, and then, as John Hunter remarks, “she received him very readily. In this curious fact, we have instinct excited by mere colour, which had so strong an effect as to get the better of everything else. But the male did not require this, the female being an animal somewhat similar to himself, was sufficient to rouse him.” (33. ‘Essays and Observations,’ by J. Hunter, edited by Owen, 1861, vol. i. p. 194.)
In an earlier chapter we have seen that the mental powers of the higher animals do not differ in kind, though greatly in degree, from the corresponding powers of man, especially of the lower and barbarous races; and it would appear that even their taste for the beautiful is not widely different from that of the Quadrumana. As the negro of Africa raises the flesh on his face into parallel ridges “or cicatrices, high above the natural surface, which unsightly deformities are considered great personal attractions” (34. Sir S. Baker, ‘The Nile Tributaries of Abyssinia,’ 1867.);–as negroes and savages in many parts of the world paint their faces with red, blue, white, or black bars,–so the male mandrill of Africa appears to have acquired his deeply-furrowed and gaudily-coloured face from having been thus rendered attractive to the female. No doubt it is to us a most grotesque notion that the posterior end of the body should be coloured for the sake of ornament even more brilliantly than the face; but this is not more strange than that the tails of many birds should be especially decorated.
With mammals we do not at present possess any evidence that the males take pains to display their charms before the female; and the elaborate manner in which this is performed by male birds and other animals is the strongest argument in favour of the belief that the females admire, or are excited by, the ornaments and colours displayed before them. There is, however, a striking parallelism between mammals and birds in all their secondary sexual characters, namely in their weapons for fighting with rival males, in their ornamental appendages, and in their colours. In both classes, when the male differs from the female, the young of both sexes almost always resemble each other, and in a large majority of cases resemble the adult female. In both classes the male assumes the characters proper to his sex shortly before the age of reproduction; and if emasculated at an early period, loses them. In both classes the change of colour is sometimes seasonal, and the tints of the naked parts sometimes become more vivid during the act of courtship. In both classes the male is almost always more vividly or strongly coloured than the female, and is ornamented with larger crests of hair or feathers, or other such appendages. In a few exceptional cases the female in both classes is more highly ornamented than the male. With many mammals, and at least in the case of one bird, the male is more odoriferous than the female. In both classes the voice of the male is more powerful than that of the female. Considering this parallelism, there can be little doubt that the same cause, whatever it may be, has acted on mammals and birds; and the result, as far as ornamental characters are concerned, may be attributed, as it appears to me, to the long-continued preference of the individuals of one sex for certain individuals of the opposite sex, combined with their success in leaving a larger number of offspring to inherit their superior attractions.
EQUAL TRANSMISSION OF ORNAMENTAL CHARACTERS TO BOTH SEXES.
With many birds, ornaments, which analogy leads us to believe were primarily acquired by the males, have been transmitted equally, or almost equally, to both sexes; and we may now enquire how far this view applies to mammals. With a considerable number of species, especially of the smaller kinds, both sexes have been coloured, independently of sexual selection, for the sake of protection; but not, as far as I can judge, in so many cases, nor in so striking a manner, as in most of the lower classes. Audubon remarks that he often mistook the musk-rat (35. Fiber zibethicus, Audubon and Bachman, ‘The Quadrupeds of North America,’ 1846, p. 109.), whilst sitting on the banks of a muddy stream, for a clod of earth, so complete was the resemblance. The hare on her form is a familiar instance of concealment through colour; yet this principle partly fails in a closely-allied species, the rabbit, for when running to its burrow, it is made conspicuous to the sportsman, and no doubt to all beasts of prey, by its upturned white tail. No one doubts that the quadrupeds inhabiting snow-clad regions have been rendered white to protect them from their enemies, or to favour their stealing on their prey. In regions where snow never lies for long, a white coat would be injurious; consequently, species of this colour are extremely rare in the hotter parts of the world. It deserves notice that many quadrupeds inhabiting moderately cold regions, although they do not assume a white winter dress, become paler during this season; and this apparently is the direct result of the conditions to which they have long been exposed. Pallas (36. ‘Novae species Quadrupedum e Glirium ordine,’ 1778, p. 7. What I have called the roe is the Capreolus sibiricus subecaudatus of Pallas.) states that in Siberia a change of this nature occurs with the wolf, two species of Mustela, the domestic horse, the Equus hemionus, the domestic cow, two species of antelopes, the musk- deer, the roe, elk, and reindeer. The roe, for instance, has a red summer and a greyish-white winter coat; and the latter may perhaps serve as a protection to the animal whilst wandering through the leafless thickets, sprinkled with snow and hoar-frost. If the above-named animals were gradually to extend their range into regions perpetually covered with snow, their pale winter-coats would probably be rendered through natural selection, whiter and whiter, until they became as white as snow.
Mr. Reeks has given me a curious instance of an animal profiting by being peculiarly coloured. He raised from fifty to sixty white and brown piebald rabbits in a large walled orchard; and he had at the same time some similarly coloured cats in his house. Such cats, as I have often noticed, are very conspicuous during day; but as they used to lie in watch during the dusk at the mouths of the burrows, the rabbits apparently did not distinguish them from their parti-coloured brethren. The result was that, within eighteen months, every one of these parti-coloured rabbits was destroyed; and there was evidence that this was effected by the cats. Colour seems to be advantageous to another animal, the skunk, in a manner of which we have had many instances in other classes. No animal will voluntarily attack one of these creatures on account of the dreadful odour which it emits when irritated; but during the dusk it would not easily be recognised and might be attacked by a beast of prey. Hence it is, as Mr. Belt believes (37. ‘The Naturalist in Nicaragua,’ p. 249.), that the skunk is provided with a great white bushy tail, which serves as a conspicuous warning.
[Fig. 70. Tragelaphus scriptus, male (from the Knowsley Menagerie).
Fig. 71. Damalis pygarga, male (from the Knowsley Menagerie).]
Although we must admit that many quadrupeds have received their present tints either as a protection, or as an aid in procuring prey, yet with a host of species, the colours are far too conspicuous and too singularly arranged to allow us to suppose that they serve for these purposes. We may take as an illustration certain antelopes; when we see the square white patch on the throat, the white marks on the fetlocks, and the round black spots on the ears, all more distinct in the male of the Portax picta, than in the female;–when we see that the colours are more vivid, that the narrow white lines on the flank and the broad white bar on the shoulder are more distinct in the male Oreas derbyanus than in the female;–when we see a similar difference between the sexes of the curiously-ornamented Tragelaphus scriptus (Fig. 70),–we cannot believe that differences of this kind are of any service to either sex in their daily habits of life. It seems a much more probable conclusion that the various marks were first acquired by the males and their colours intensified through sexual selection, and then partially transferred to the females. If this view be admitted, there can be little doubt that the equally singular colours and marks of many other antelopes, though common to both sexes, have been gained and transmitted in a like manner. Both sexes, for instance, of the koodoo (Strepsiceros kudu) (Fig. 64) have narrow white vertical lines on their hind flanks, and an elegant angular white mark on their foreheads. Both sexes in the genus Damalis are very oddly coloured; in D. pygarga the back and neck are purplish-red, shading on the flanks into black; and these colours are abruptly separated from the white belly and from a large white space on the buttocks; the head is still more oddly coloured, a large oblong white mask, narrowly-edged with black, covers the face up to the eyes (Fig. 71); there are three white stripes on the forehead, and the ears are marked with white. The fawns of this species are of a uniform pale yellowish-brown. In Damalis albifrons the colouring of the head differs from that in the last species in a single white stripe replacing the three stripes, and in the ears being almost wholly white. (38. See the fine plates in A. Smith’s ‘Zoology of South Africa,’ and Dr. Gray’s ‘Gleanings from the Menagerie of Knowsley.’) After having studied to the best of my ability the sexual differences of animals belonging to all classes, I cannot avoid the conclusion that the curiously-arranged colours of many antelopes, though common to both sexes, are the result of sexual selection primarily applied to the male.
The same conclusion may perhaps be extended to the tiger, one of the most beautiful animals in the world, the sexes of which cannot be distinguished by colour, even by the dealers in wild beasts. Mr. Wallace believes (39. ‘Westminster Review,’ July 1, 1867, p. 5.) that the striped coat of the tiger “so assimilates with the vertical stems of the bamboo, as to assist greatly in concealing him from his approaching prey.” But this view does not appear to me satisfactory. We have some slight evidence that his beauty may be due to sexual selection, for in two species of Felis the analogous marks and colours are rather brighter in the male than in the female. The zebra is conspicuously striped, and stripes cannot afford any protection in the open plains of South Africa. Burchell (40. ‘Travels in South Africa,’ 1824, vol. ii. p. 315.) in describing a herd says, “their sleek ribs glistened in the sun, and the brightness and regularity of their striped coats presented a picture of extraordinary beauty, in which probably they are not surpassed by any other quadruped.” But as throughout the whole group of the Equidae the sexes are identical in colour, we have here no evidence of sexual selection. Nevertheless he who attributes the white and dark vertical stripes on the flanks of various antelopes to this process, will probably extend the same view to the Royal Tiger and beautiful Zebra.
We have seen in a former chapter that when young animals belonging to any class follow nearly the same habits of life as their parents, and yet are coloured in a different manner, it may be inferred that they have retained the colouring of some ancient and extinct progenitor. In the family of pigs, and in the tapirs, the young are marked with longitudinal stripes, and thus differ from all the existing adult species in these two groups. With many kinds of deer the young are marked with elegant white spots, of which their parents exhibit not a trace. A graduated series can be followed from the axis deer, both sexes of which at all ages and during all seasons are beautifully spotted (the male being rather more strongly coloured than the female), to species in which neither the old nor the young are spotted. I will specify some of the steps in this series. The Mantchurian deer (Cervus mantchuricus) is spotted during the whole year, but, as I have seen in the Zoological Gardens, the spots are much plainer during the summer, when the general colour of the coat is lighter, than during the winter, when the general colour is darker and the horns are fully developed. In the hog-deer (Hyelaphus porcinus) the spots are extremely conspicuous during the summer when the coat is reddish-brown, but quite disappear during the winter when the coat is brown. (41. Dr. Gray, ‘Gleanings from the Menagerie of Knowsley,’ p. 64. Mr. Blyth, in speaking (‘Land and Water,’ 1869, p. 42) of the hog-deer of Ceylon, says it is more brightly spotted with white than the common hog-deer, at the season when it renews its horns.) In both these species the young are spotted. In the Virginian deer the young are likewise spotted, and about five per cent. of the adult animals living in Judge Caton’s park, as I am informed by him, temporarily exhibit at the period when the red summer coat is being replaced by the bluish winter coat, a row of spots on each flank, which are always the same in number, though very variable in distinctness. From this condition there is but a very small step to the complete absence of spots in the adults at all seasons; and, lastly, to their absence at all ages and seasons, as occurs with certain species. From the existence of this perfect series, and more especially from the fawns of so many species being spotted, we may conclude that the now living members of the deer family are the descendants of some ancient species which, like the axis deer, was spotted at all ages and seasons. A still more ancient progenitor probably somewhat resembled the Hyomoschus aquaticus–for this animal is spotted, and the hornless males have large exserted canine teeth, of which some few true deer still retain rudiments. Hyomoschus, also, offers one of those interesting cases of a form linking together two groups, for it is intermediate in certain osteological characters between the pachyderms and ruminants, which were formerly thought to be quite distinct. (42. Falconer and Cautley, ‘Proc. Geolog. Soc.’ 1843; and Falconer’s ‘Pal. Memoirs,’ vol. i. p. 196.)
A curious difficulty here arises. If we admit that coloured spots and stripes were first acquired as ornaments, how comes it that so many existing deer, the descendants of an aboriginally spotted animal, and all the species of pigs and tapirs, the descendants of an aboriginally striped animal, have lost in their adult state their former ornaments? I cannot satisfactorily answer this question. We may feel almost sure that the spots and stripes disappeared at or near maturity in the progenitors of our existing species, so that they were still retained by the young; and, owing to the law of inheritance at corresponding ages, were transmitted to the young of all succeeding generations. It may have been a great advantage to the lion and puma, from the open nature of their usual haunts, to have lost their stripes, and to have been thus rendered less conspicuous to their prey; and if the successive variations, by which this end was gained, occurred rather late in life, the young would have retained their stripes, as is now the case. As to deer, pigs, and tapirs, Fritz Muller has suggested to me that these animals, by the removal of their spots or stripes through natural selection, would have been less easily seen by their enemies; and that they would have especially required this protection, as soon as the carnivora increased in size and number during the tertiary periods. This may be the true explanation, but it is rather strange that the young should not have been thus protected, and still more so that the adults of some species should have retained their spots, either partially or completely, during part of the year. We know that, when the domestic ass varies and becomes reddish-brown, grey, or black, the stripes on the shoulders and even on the spine frequently disappear, though we cannot explain the cause. Very few horses, except dun-coloured kinds, have stripes on any part of their bodies, yet we have good reason to believe that the aboriginal horse was striped on the legs and spine, and probably on the shoulders. (43. The ‘Variation of Animals and Plants under Domestication,’ 1868, vol. i. pp. 61-64.) Hence the disappearance of the spots and stripes in our adult existing deer, pigs, and tapirs, may be due to a change in the general colour of their coats; but whether this change was effected through sexual or natural selection, or was due to the direct action of the conditions of life, or to some other unknown cause, it is impossible to decide. An observation made by Mr. Sclater well illustrates our ignorance of the laws which regulate the appearance and disappearance of stripes; the species of Asinus which inhabit the Asiatic continent are destitute of stripes, not having even the cross shoulder-stripe, whilst those which inhabit Africa are conspicuously striped, with the partial exception of A. taeniopus, which has only the cross shoulder-stripe and generally some faint bars on the legs; and this species inhabits the almost intermediate region of Upper Egypt and Abyssinia. (44. ‘Proc. Zool. Soc.’ 1862, p. 164. See, also, Dr. Hartmann, ‘Ann. d. Landw.’ Bd. xliii. s. 222.)
QUADRUMANA.
[Fig. 72. Head of Semnopithecus rubicundus. This and the following figures (from Prof. Gervais) are given to shew the odd arrangement and development of the hair on the head.
Fig. 73. Head of Semnopithecus comatus.
Fig. 74. Head of Cebus capucinus.
Fig. 75. Head of Ateles marginatus.
Fig. 76. Head of Cebus vellerosus.]
Before we conclude, it will be well to add a few remarks on the ornaments of monkeys. In most of the species the sexes resemble each other in colour, but in some, as we have seen, the males differ from the females, especially in the colour of the naked parts of the skin, in the development of the beard, whiskers, and mane. Many species are coloured either in so extraordinary or so beautiful a manner, and are furnished with such curious and elegant crests of hair, that we can hardly avoid looking at these characters as having been gained for the sake of ornament. The accompanying figures (Figs. 72 to 76) serve to shew the arrangement of the hair on the face and head in several species. It is scarcely conceivable that these crests of hair, and the strongly contrasted colours of the fur and skin, can be the result of mere variability without the aid of selection; and it is inconceivable that they can be of use in any ordinary way to these animals. If so, they have probably been gained through sexual