Secondary Sexual Characters Of Mammals

Quadrupeds use their voices to:

• signal danger
• call from one member of a troop to another
• call for lost offspring
• call for protection to their mother

But I am concerned only with the difference between the voices of the sexes, between:

• lion and lioness
• bull and cow

Almost all male animals use their voices much more during the rutting-season than at any other time.

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 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 Säugethiere 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 Säugethiere 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, ‘Säugethiere,’ 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, étant plus terne, le blanc moins pur, les raies ayant moins de largeur et les taches moins de diamètre.” (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 Mammifères’).]

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.