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 Müller 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 selection, though transmitted equally, or almost equally, to both sexes. With many of the Quadrumana, we have additional evidence of the action of sexual selection in the greater size and strength of the males, and in the greater development of their canine teeth, in comparison with the females.

[Fig. 77. Cercopithecus petaurista (from Brehm).]

A few instances will suffice of the strange manner in which both sexes of some species are coloured, and of the beauty of others. The face of the Cercopithecus petaurista (Fig. 77) is black, the whiskers and beard being white, with a defined, round, white spot on the nose, covered with short white hair, which gives to the animal an almost ludicrous aspect. The Semnopithecus frontatus likewise has a blackish face with a long black beard, and a large naked spot on the forehead of a bluish-white colour. The face of Macacus lasiotus is dirty flesh-coloured, with a defined red spot on each cheek. The appearance of Cercocebus aethiops is grotesque, with its black face, white whiskers and collar, chestnut head, and a large naked white spot over each eyelid. In very many species, the beard, whiskers, and crests of hair round the face are of a different colour from the rest of the head, and when different, are always of a lighter tint (45. I observed this fact in the Zoological Gardens; and many cases may be seen in the coloured plates in Geoffroy St.-Hilaire and F. Cuvier, ‘Histoire Nat. des Mammifères,’ tom. i. 1824.), being often pure white, sometimes bright yellow, or reddish. The whole face of the South American Brachyurus calvus is of a “glowing scarlet hue”; but this colour does not appear until the animal is nearly mature. (46. Bates, ‘The Naturalist on the Amazons,’ 1863, vol. ii. p. 310.) The naked skin of the face differs wonderfully in colour in the various species. It is often brown or flesh-colour, with parts perfectly white, and often as black as that of the most sooty negro. In the Brachyurus the scarlet tint is brighter than that of the most blushing Caucasian damsel. It is sometimes more distinctly orange than in any Mongolian, and in several species it is blue, passing into violet or grey. In all the species known to Mr. Bartlett, in which the adults of both sexes have strongly-coloured faces, the colours are dull or absent during early youth. This likewise holds good with the mandrill and Rhesus, in which the face and the posterior parts of the body are brilliantly coloured in one sex alone. In these latter cases we have reason to believe that the colours were acquired through sexual selection; and we are naturally led to extend the same view to the foregoing species, though both sexes when adult have their faces coloured in the same manner.

[Fig. 78. Cercopithecus diana (from Brehm).]

Although many kinds of monkeys are far from beautiful according to our taste, other species are universally admired for their elegant appearance and bright colours. The Semnopithecus nemaeus, though peculiarly coloured, is described as extremely pretty; the orange-tinted face is surrounded by long whiskers of glossy whiteness, with a line of chestnut-red over the eyebrows; the fur on the back is of a delicate grey, with a square patch on the loins, the tail and the fore-arms being of a pure white; a gorget of chestnut surmounts the chest; the thighs are black, with the legs chestnut-red. I will mention only two other monkeys for their beauty; and I have selected these as presenting slight sexual differences in colour, which renders it in some degree probable that both sexes owe their elegant appearance to sexual selection. In the moustache-monkey (Cercopithecus cephus) the general colour of the fur is mottled-greenish with the throat white; in the male the end of the tail is chestnut, but the face is the most ornamented part, the skin being chiefly bluish-grey, shading into a blackish tint beneath the eyes, with the upper lip of a delicate blue, clothed on the lower edge with a thin black moustache; the whiskers are orange-coloured, with the upper part black, forming a band which extends backwards to the ears, the latter being clothed with whitish hairs. In the Zoological Society’s Gardens I have often overheard visitors admiring the beauty of another monkey, deservedly called Cercopithecus diana (Fig. 78); the general colour of the fur is grey; the chest and inner surface of the forelegs are white; a large triangular defined space on the hinder part of the back is rich chestnut; in the male the inner sides of the thighs and the abdomen are delicate fawn-coloured, and the top of the head is black; the face and ears are intensely black, contrasting finely with a white transverse crest over the eyebrows and a long white peaked beard, of which the basal portion is black. (47. I have seen most of the above monkeys in the Zoological Society’s Gardens. The description of the Semnopithecus nemaeus is taken from Mr. W.C. Martin’s ‘Natural History of Mammalia,’ 1841, p. 460; see also pp. 475, 523.)

In these and many other monkeys, the beauty and singular arrangement of their colours, and still more the diversified and elegant arrangement of the crests and tufts of hair on their heads, force the conviction on my mind that these characters have been acquired through sexual selection exclusively as ornaments.

SUMMARY

The law of battle for the possession of the female appears to prevail throughout the whole great class of mammals.

Most naturalists will admit that the greater size, strength, courage, and pugnacity of the male, his special weapons of offence, as well as his special means of defence, have been acquired or modified through that form of selection which I have called sexual. This does not depend on any superiority in the general struggle for life, but on certain individuals of one sex, generally the male, being successful in conquering other males, and leaving a larger number of offspring to inherit their superiority than do the less successful males.

There is another and more peaceful kind of contest, in which the males endeavour to excite or allure the females by various charms. This is probably carried on in some cases by the powerful odours emitted by the males during the breeding-season; the odoriferous glands having been acquired through sexual selection. Whether the same view can be extended to the voice is doubtful, for the vocal organs of the males must have been strengthened by use during maturity, under the powerful excitements of love, jealousy or rage, and will consequently have been transmitted to the same sex. Various crests, tufts, and mantles of hair, which are either confined to the male, or are more developed in this sex than in the female, seem in most cases to be merely ornamental, though they sometimes serve as a defence against rival males. There is even reason to suspect that the branching horns of stags, and the elegant horns of certain antelopes, though properly serving as weapons of offence or defence, have been partly modified for ornament.

When the male differs in colour from the female, he generally exhibits darker and more strongly-contrasted tints. We do not in this class meet with the splendid red, blue, yellow, and green tints, so common with male birds and many other animals. The naked parts, however, of certain Quadrumana must be excepted; for such parts, often oddly situated, are brilliantly coloured in some species. The colours of the male in other cases may be due to simple variation, without the aid of selection. But when the colours are diversified and strongly pronounced, when they are not developed until near maturity, and when they are lost after emasculation, we can hardly avoid the conclusion that they have been acquired through sexual selection for the sake of ornament, and have been transmitted exclusively, or almost exclusively, to the same sex. When both sexes are coloured in the same manner, and the colours are conspicuous or curiously arranged, without being of the least apparent use as a protection, and especially when they are associated with various other ornamental appendages, we are led by analogy to the same conclusion, namely, that they have been acquired through sexual selection, although transmitted to both sexes. That conspicuous and diversified colours, whether confined to the males or common to both sexes, are as a general rule associated in the same groups and sub-groups with other secondary sexual characters serving for war or for ornament, will be found to hold good, if we look back to the various cases given in this and the last chapter.

The law of the equal transmission of characters to both sexes, as far as colour and other ornaments are concerned, has prevailed far more extensively with mammals than with birds; but weapons, such as horns and tusks, have often been transmitted either exclusively or much more perfectly to the males than to the females. This is surprising, for, as the males generally use their weapons for defence against enemies of all kinds, their weapons would have been of service to the females. As far as we can see, their absence in this sex can be accounted for only by the form of inheritance which has prevailed. Finally, with quadrupeds the contest between the individuals of the same sex, whether peaceful or bloody, has, with the rarest exceptions, been confined to the males; so that the latter have been modified through sexual selection, far more commonly than the females, either for fighting with each other or for alluring the opposite sex.