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 supérieures de leurs cuisses et le grand espace nu de leurs fesses sont également colorés du rouge le plus vif, avec un mélange de bleu qui ne manque reellement pas d’élégance.” (31. Gervais, ‘Hist. Nat. des Mammifères,’ 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 Mammifères,’ 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).